Psephosauriscus
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''Psephosauriscus'' 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
placodont Placodonts (" tablet teeth") are an extinct order of marine reptiles that lived during the Triassic period, becoming extinct at the end of the period. They were part of Sauropterygia, the group that includes plesiosaurs. Placodonts were generall ...
reptile from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
of Israel and Egypt. It is known from bony armor plates that have been found from
Makhtesh Ramon Makhtesh Ramon (; ''lit.'' Ramon Crater/ Makhtesh; ; ''lit.'' The Ruman Wadi) is a geological feature of Israel's Negev desert. Located some 85 km south of Beersheba, the landform is the world's largest "erosion cirque" (steephead valley ...
in Israel's
Negev The Negev ( ; ) or Naqab (), is a desert and semidesert region of southern Israel. The region's largest city and administrative capital is Beersheba (pop. ), in the north. At its southern end is the Gulf of Aqaba and the resort town, resort city ...
desert and Araif en Naqua on Egypt's
Sinai Peninsula The Sinai Peninsula, or simply Sinai ( ; ; ; ), is a peninsula in Egypt, and the only part of the country located in Asia. It is between the Mediterranean Sea to the north and the Red Sea to the south, and is a land bridge between Asia and Afri ...
. The genus was erected in 2002 as a replacement name for several species of the genus ''Psephosaurus'', which was named in 1957. It includes the species ''P. mosis'', ''P. ramonensis'', ''P. sinaiticus'', and a possible fourth species, ''P. rhombifer''. All species, with the exception of ''P. ramonensis'', were once assigned to the genus ''Psephosaurus''. Remains of ''P. mosis'' and ''P. ramonensis'' were found in Makhtesh Ramon, while ''P. sinaiticus'' and ''P. rhombifer'' were found in Araif en Naqua.


History

Armor of ''Psephosauriscus'' is abundant in Middle Triassic limestone in the Sinai and Negev regions, which is comparable to the
Muschelkalk The Muschelkalk (German for "shell-bearing limestone"; ) is a sequence of sedimentary rock, sedimentary rock strata (a lithostratigraphy, lithostratigraphic unit) in the geology of central and western Europe. It has a Middle Triassic (240 to 230 m ...
of European rock sequences. While the German Muschelkalk contains many nearly complete skeletons of placodonts, the only non-armor skeletal bones from the Middle East are two partial skulls and a fragmentary lower jaw, which cannot be assigned to ''Psephosauriscus'' or any other placodont because of their lack of diagnostic features. Remains of ''Psephosauriscus'' come from two deposits called the '' Beneckeia'' beds and the ''
Ceratites ''Ceratites'' is an extinct genus of ammonite cephalopods. These nektonic carnivores lived in marine habitats in what is now Europe, during the Triassic, from the upper-most Anisian In the geologic timescale, the Anisian is the lower stage (st ...
'' beds, named after their most common
ammonite Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
fossils. The ''Beneckeia'' beds date back to the
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
stage of the Middle Triassic, and the ''Ceratites'' beds date back to the end of the Anisian and beginning of the
Ladinian The Ladinian is a stage and age in the Middle Triassic series or epoch. It spans the time between Ma and ~237 Ma (million years ago). The Ladinian was preceded by the Anisian and succeeded by the Carnian (part of the Upper or Late Triassic ...
stage. Swedish paleontologist F. Brotzen described placodont armor from Makhtesh Ramon in 1957, naming the species ''Psephosaurus mosis'' and ''Psephosaurus picardi'' from the ''Beneckeia'' and ''Ceratites'' beds, respectively. ''P. picardi'' is now considered a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'' because it was based on an impression of the inner surface of the carapace that did not possess any distinct features. In 1959, Austrian paleontologist Georg Haas named ''P. sinaiticus'' and ''P. rhombifer'' from Araif en Naqua. In 2002, paleontologist Olivier Rieppel erected the genus ''Psephosauriscus'' to include most of the species named by Brotzen and Haas, which he considered distinct from the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of ''Psephosaurus'', ''P. suevicus'', named by
Eberhard Fraas Eberhard Fraas (26 June 1862 – 6 March 1915) was a German scientist, geologist and paleontologist. He worked as a curator at the Stuttgarter Naturaliensammlung and discovered the dinosaurs of the Tendaguru formation in then German East Africa (n ...
from the Middle Triassic of Germany. Rieppel noted differences between the armor plates of ''P. suevicus'' and the Middle Eastern species that warranted a new genus. However, ''P. rhombifer'' could not be assigned with certainty to ''Psephosauriscus'' because 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 ...
specimen described by Haas had since been lost. Rieppel reported additional material from Araif en Naqua that showed similarities to ''P. rhombifer'' and the species of ''Psephosauriscus'', which he tentatively described as a species within ''Psephosauriscus'', ''Psephosauriscus cf. rhombifer''. Rieppel also named ''Psephosauriscus ramonensis'' as an entirely new species.


Species

The type species of ''Psephosauriscus'', ''P. mosis'', is known from a single specimen including portions of the
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
and
plastron The turtle shell is a shield for the ventral and dorsal parts of turtles (the Order (biology), order Testudines), completely enclosing all the turtle's vital organs and in some cases even the head. It is constructed of modified bony elements such ...
. The
scute A scute () or scutum (Latin: ''scutum''; plural: ''scuta'' "Scutum (shield), shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of Bird anatomy#Scales, birds. The ter ...
s that cover the armor plates do not have as well-defined a shape as the hexagonal osteoderms that lay underneath them. The osteoderms that form the plastron are relatively large. Some osteoderms have a raised keel. Two distinct ridges along either side of the shell separate the carapace, a lateral wall of smaller osteoderms, and the plastron. ''P. ramonensis'' is known from a partial carapace and connected plastron. The osteoderms of the carapace have smoother edges than most other species of ''Psephosauriscus'', and lack the keel of species like ''P. mosis''. The carapace curves into the lateral wall of the body without a separating ridge as in ''P. mosis'', but a lower ridge does separate the lateral wall from the plastron. ''P. sinaiticus'' is known from several armor fragments and larger pieces of the carapace and plastron. The osteoderms of the carapace and plastron are smaller than those of other species. As in ''P. mosis'', two ridges separate run along the side of the shell. ''P. cf. rhombifer'' can be distinguished from the three other species of ''Psephosauriscus'' by the rectangular shape of its scutes. Unlike the smooth shells of other species, the carapace of ''P. cf. rhombifer'' has a bumpy surface; each osteoderm is covered in radiating grooves and has a small depression at its center.


References

{{Taxonbar, from=Q20722690 Middle Triassic reptiles of Africa Fossils of Israel Fossils of Egypt Placodontia Anisian life Anisian genus first appearances Ladinian genus extinctions Sauropterygian genera Fossil taxa described in 2002