Parahelicoprion

Estimates of the extent of the complete whorl, body size, and ecology of Parahelicoprion are speculative as a result of its incomplete fossils, although it is assumed to have been very large, predatory, and potentially pelagic.

The type specimen of Parahelicoprion clerci was found in the Ural Mountains region of Russia,[1][2] in strata dated to the Artinskian stage of the Cisuralian epoch (early Permian).

[5] The first five portions of the specimen to be found, which consist of several teeth, were discovered incidentally by a miner and were badly damaged as a result poor handling during collection.

[16][17] The specific name, P. clerci, honors Onésime Clerc, who at the time of its description was the president of the Ural Society of Natural Science Lovers.

The species is named in honor of Dr. Mario Suarez-Riglos,[18] and the type specimen is currently housed in the collection of the Noel Kempff Mercado Natural History Museum.

Both Parahelicoprion species are very incompletely known,[23] and the only material which has been assigned confidently to the genus consists of fragments of the symphyseal (midline) tooth whorl.

[5] The description of P. mariosuarezi suggests that it likely possessed a very short tooth whorl situated at the tip of a greatly elongated pair of jaws,[18][28] based on both the partial skull of Sarcoprion edax and the well-preserved fossils of the related caseodonts.

[30] Karpinsky himself did not provide body length estimates in his descriptions,[30]: S1  but did note that the tooth crowns of Parahelicoprion clerci were significantly larger than those of any then-known Helicoprion specimens and suggested that the animal must have been very large to accommodate them.

Ellis states that, in spite of the fragmentary nature of the known material, "... unless it [Parahelicoprion] was an animal with a gigantic head or outlandishly oversized teeth, it had to have been a monster, at least 100 feet long and maybe more."

[10] Lengths of 11–13 meters (36–43 ft) have more recently been suggested for Parahelicoprion by online sources,[36][30]: S1  although these numbers originate from non-academic amateur researchers and are not supported by scientific literature.

[30]: S1  Due to the fragmentary nature of the known material, it has been considered unreasonable by some researchers to give precise total length estimates for Helicoprion, Parahelicoprion, or any other members of Edestoidea.

[4][27] In a 1925 publication, Karpinsky suggested that P. clerci may represent a directly intermediate, transitional form between the "primitive" genus Campodus (based on material now assigned to Agassizodus)[27] and the more derived Helicoprion,[6] a conclusion agreed with by Egil Nielsen in his 1952 description of Sarcoprion and Parahelicampodus.

[42] Svend Erik Bendix-Almgreem, in a 1976 paper, suggested that Parahelicoprion may have been part of a radiation of whorl-toothed cartilaginous fish unrelated to the helicoprionids and edestids.

[9] Rainer Zangerl (1981) considered Parahelicoprion to be the sister taxon to Campyloprion in his morphological analysis of all (then known) members of the newly proposed order Eugeneodontida, which united the edestids, helicoprionids, and the caseodonts.

[27] Following Zangerl's analysis, paleontologists Dagmar Merino-Rodo and Philippe Janvier concluded in their 1986 description of P. mariosuarezi that the genus Parahelicoprion may lack defining derived characteristics, which puts its status as a monophyletic group into question and complicates the matter of assigning new species.

[18] In a 2018 publication, paleontologist Serge Naugholnykh proposed that P. clerci simply represents an especially large individual of Helicoprion and that the two genera are synonymous,[3][29] although subsequent papers have continued to recognize Parahelicoprion as a valid genus of helicoprionid eugeneodont.

[1][5][6] While multiple feeding styles are thought to have been present among different genera of edestoids,[33][31] it has been hypothesized that members of the Helicoprionidae were molluscivorous and fed primarily on ammonoid and coleoid cephalopods,[33][45] with smaller cartilaginous fish potentially constituting a portion of their diet as well.

[3][8][46] During the early Permian, reef habitats made up of crinoids, sponges,[6] bryozoans and rugose corals were present, which were inhabited by a diverse assemblage of trilobites,[8] goniatite and nautiloid cephalopods,[46] and fishes.

[3] In addition to Parahelicoprion clerci, the Divya Formation yields a large variety of chondrichthyan fossils, including the remains of euselachian sharks, hybodonts, petalodonts, cochliodonts and cladodonts.

[5][6] The Copacabana Formation represented a shallow marine habitat, somewhat older than the Arta Beds and dated to the boundary between the Carboniferous and the Permian.

While lower fish-bearing strata of the Copacabana Formation are believed to represent a benthic reef community, Merino-Rodo and Janvier suggest that the sandstones which compose the upper fish-bearing layer and which preserve the whorl of Parahelicoprion may have formed in an even shallower, intertidal habitat, and that the type of P. mariosuarezi represents the remains of an animal which stranded on the shore.

[43] Karpinsky has suggested that the disappearance of the seaway connecting the Arctic and Tethys oceans was directly responsible for the extinction of the Uralian edestoids.

[6] Alongside Parahelicoprion, many other cartilaginous fish genera of the Divya Formation disappear from the fossil record at the close of the Artinskian stage.

Fragments of the large spine-like organ which Karpinsky assigned to P. clerci . [ 6 ] This specimen has since been recognized as belonging to the form-genera Physonemus or Xystracanthus [ 21 ] [ 22 ]
Speculative life reconstruction of Parahelicoprion clerci , similar to that proposed for P. mariosuarezi in its 1986 description and illustrated in Philippe Janvier's textbook Early Vertebrates (1996) [ 18 ] [ 28 ]
Three re-assembled crowns of the type specimen of P. clerci , photographed in lateral (1a) and transverse (1b) views
Additional fragments of the P. clerci holotype, including the presumed foremost tooth crown (fig. 2)
Illustration of the rearmost known crown tip of P. clerci in transverse (a) and lateral (b) views
Tip of the "wing" or "rib" of the tooth of P. clerci , displaying this species' unique arrangement of serrations (above) and crenulations (below)
Reconstructed teeth of P. clerci in lateral (1a, 2a), transverse (1b, 2b), and superior (2c) views, compared with the teeth of Helicoprion bessonowi (3a, 3b, 3c). Drawn to scale [ 5 ]
Whorl segments of Parahelicoprion clerci illustrated in lateral (3) and transverse (4) views, compared with those of the related eugeneodonts Campodus ( Agassizodus ?) , [ 27 ] Helicoprion, and Edestus [ 12 ]
The whorls of the related Helicoprion bessonowi (left) and Sarcoprion edax (right) displayed at the Moscow Paleontological museum
The teeth of Artiodus prominens , a euselachian stem-group shark known from the Arta Beds of Krasnoufimsk [ 8 ]