Some Vertebrate Types (Chondrichthyes, Actinopterygii, Sarcopterygii, and Tetrapoda) from Two Paleozoic Lagerstätten of Ohio, U.S.A.

ABSTRACT The type specimens of eight Paleozoic vertebrate species from Ohio, U.S.A., originally described by J. S. Newberry and subsequently reposited in the Orton Geological Museum, are reviewed and illustrated photographically. They include four actinopterygian fish species from the Linton Konservat-Lagerstätte (Upper Freeport Coal; Carboniferous) in Jefferson County, Ohio, described in 1857 under the basionyms Mecolepis corrugatus (now Haplolepis corrugata), M. insculptus (synonymized as Parahaplolepis tuberculata), M. lineatus (now Pyritocephalus lineatus), and M. ovoideus (now Microhaplolepis ovoidea). One actinopterygian species, Eurylepis striolatus, a junior synonym of Haplolepis corrugata, was described in 1873 from the Linton deposit; and one chondrichthyan species, Orthacanthus gracilis, now replaced as Orthacanthus adamas Babcock, 2024, was first described by Newberry in 1875 from Linton. Lectotypes are designated from the syntypic series of M. insculptus and O. adamas. A tooth from the Linton Lagerstätte, originally described as a fish in 1857 under the basionym Rhizodus lancifer, is the holotype of the tetrapod (anthracosaur) Anthracosaurus lancifer. One Devonian sarcopterygian fish species, Onychodus ortoni, was described in 1889 from the Ohio Shale, Huron Shale Member, a Konservat-Lagerstätte occurring in Franklin County, Ohio.


INTRODUCTION
John Strong Newberry (1822Newberry ( -1892)), who was a major figure in American science during the latter half of the 19th century (Berg, 1996;White, 1909), served as Chief Geologist of the second Geological Survey of Ohio between 1869 and 1882 (Hansen & Collins, 1979).During that time, he played an important role in the early growth of the Geological Museum of The Ohio State University (Babcock et al., 2023), which was founded in 1874 by Edward Orton, the university's first Professor of Geology and President.Between 1869 and 1882, Orton also served as an Assistant Geologist of the Geological Survey of Ohio under Newberry (Hansen & Collins, 1979).
Prior to 1873, Geological Survey of Ohio collections were deposited in various museums and colleges.Beginning around 1874, however, some rock, mineral, and fossil collections in the possession of the Geological Survey of Ohio, including legacy collections, began to be transferred to the Geological Museum (later renamed the Orton Geological Museum).Notably, the museum received fossil collections produced or studied by officers of the Geological Survey (J. S. Newberry, E. B. Andrews, Edward Orton, and F. B. Meek), and from the local and special assistants of the Survey (among them, E. D. Cope, G. K. Gilbert, James Hall, Herman Hertzer, O. C. Marsh, R. P. Whitfield, and N. H. Winchell).Transfer of voucher specimens from the Geological Survey of Ohio continued through Edward Orton's term as Chief Geologist of the Survey (1882-1899), and well into the 20th century.
Beginning in 1873, the second Geological Survey of Ohio published summary reports of studies on Ohio geology and paleontology, and owing to Newberry's initiative, many of the fossils described and illustrated in these reports are now in the Orton Museum.The early reports include seminal papers on the fossils of Ohio and some neighboring states, including Ordovician to Carboniferous invertebrates (Hall & Whitfield, 1875;Meek, 1873Meek, , 1875;;Nicholson, 1875), Devonian and Carboniferous fishes and conodonts (Newberry, 1873a(Newberry, , 1873b(Newberry, , 1874(Newberry, , 1875a(Newberry, , 1875b)), Carboniferous amphibians (Cope, 1875), and Carboniferous plants (Andrews, 1875;Newberry, 1873b).In addition, Newberry (1889) published a compilation of Paleozoic fishes as a U.S. Geological Survey Monograph, incorporating a substantial number of Ohio specimens.Some line-art illustrations from these various works were reprinted in subsequent 19th century publications such as Lesley (1889aLesley ( , 1889bLesley ( , 1890)).The impact of Newberry's work on the developing understanding of fossil fishes was reviewed by Woodward (1890).
One important set of fossils reposited in the Orton Museum is a portion of the Paleozoic vertebrates, mostly fishes, studied by Newberry between about 1855 and 1889.The museum retains more than 100 specimens amassed by the Geological Survey of Ohio, including primary types of at least eight species.Many of the other described fishes from Newberry's collection, plus a large number of undescribed specimens, are reposited in the American Museum of Natural History (Hook & Baird, 1986, Lowney, 1986;Westoll, 1944).Some studied fish specimens were placed in the Department of Geology at Ohio Wesleyan University, Delaware, Ohio, as early as the 1850s.At least one specimen was placed with Princeton University, and has since been transferred to the Yale Peabody Museum (Baird, 1962b).
The purpose of this communication is to illustrate the type Paleozoic vertebrates described by J. S. Newberry (1857Newberry ( , 1875aNewberry ( , 1889) ) that are reposited in the Orton Geological Museum (OSU), and to update, clarify, and stabilize their nomenclatural status.This is not intended to be a complete systematic treatment, but rather a starting point for other work.In addition to papers specific to the materials treated here, the reader is referred to broader or related discussions such as Ginter et al. (2010) for chondrichthyans, and Elliot (2014, 2018) and Schultze et al. (2021) for haplolepids (actinopterygians).
Species indicated here as Newberry (1857) have been erroneously cited with an 1856 date in most earlier publications.The volume in which the names first appear are proceedings for 1856, but the title page of the volume states that it was published in 1857 (see Babcock, 2024).Thus, the names became available in the sense of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 2000) in 1857.
The specimens discussed here are important partly because of their type status and historical value, but they also derive from two recognized Konservat-Lagerstätten (conservation deposits; see Babcock, 2024;Babcock et al., 2023;Hurey et al., 2017;Seilacher, 1970;Seilacher et al., 1985).Seilacher et al. (1985:17) mentioned the Ohio Shale as a Konservat-Lagerstätte, comparing it to a stagnant basin deposit of Holzmaden-type, and characterized by the presence of marine bituminous shales.The thin cannel coal interval below the main Upper Freeport Coal seam fits into the non-marine, lignitic shale facies of the paleoenvironmental model advocated by Seilacher et al. (1985:fig. 1).Studies addressing the paleoenvironmental setting and taphonomic conditions associated with the Linton Lagerstätte include Hook and Baird (1988), Hook and Ferm (1985), Hurey et al. (2017), andWitzmann et al. (2017).

LOCALITY INFORMATION AND GEOLOGIC SETTINGS
Locality: Ohio Shale, Perry Township, Ohio One sarcopterygian species, Onychodus ortoni Newberry, 1889, was described from a specimen collected from the Ohio Shale, Huron Shale Member (Upper Devonian: Famennian), in Perry Township, Franklin County, Ohio, U.S.A. Numerous other specimens, including placoderm fish and Protosalvinia spores, from the Huron Shale Member in the Orton Geological Museum, collected between about 1875 and 1965, are from the lower reaches of Flint Run, a tributary to the Olentangy River, which in some early reports was referred to as the Whetstone River (e.g., Orton, 1878).This locality, just north of the city of Worthington, was referred to by Stauffer et al. (1911:27, pl.IV) as "the Narrows;" it was re-illustrated by Westgate (1926:pl. III b).Excellent exposures of the Huron Shale Member along Flint Run are now included in Camp Mary Orton (see Hellstrom & Babcock, 2000;Stauffer, 1911, for stratigraphic sections).Almost certainly the Onychodus specimen was collected from Flint Run or nearby exposures along the Olentangy River.
The lower part of the Huron Shale Member of the Ohio Shale yields fossil fish remains at various localities in Franklin, Delaware, Huron, and Lorain counties, Ohio.Specimens are derived both from thinly laminated black shale layers and from large, ferroan carbonate concretions.Most remains are disarticulated, but occasionally articulated material is found in the concretions.The fish fauna of the Huron Shale Member was summarized by Eastman (1907).
When categorizing the Ohio Shale as a stagnant basin deposit of Holzmaden-type, Seilacher et al. (1985:17) did not specify whether they were referring to black shales of the Huron Member, in the lower part of the formation, or of the Cleveland Shale Member, in the upper part of the formation.A much more diverse fauna has been recorded from the Cleveland Shale Member (e.g., Eastman, 1907).Nevertheless, the two members record similar laminated black shales, both have layers of large Fe-rich carbonate concretions, and both have yielded hard-part plus nonbiomineralized-part preservation of fish remains.As the Huron Member and Cleveland Member both represent similar organic-rich lithofacies, both can be considered Holzmaden-type deposits.Both members record relatively little physical or biological disturbance of fine-grained sediment at the sediment-water interface apart from simple traces (Schieber, 2003).Many authors have discussed models for the origin of organicrich facies in the Ohio Shale and correlative units, and anoxia or dysoxia in either the sediment or lower part of the water column is a common theme (e.g., Raiswell & Berner, 1985;Raiswell et al., 1988;Rhoads & Morse, 1971;Rich, 1951;Sageman et al., 2003;Schieber, 2003).Long (1991) interpreted the onychodontiform species Onychodus jandemarrai Andrews et al., 2006, as a nektobenthic predator that would lie in wait for its prey close to the sediment surface, and then lunge at its prey.Assuming that the related species O. ortoni had the same predatory strategy, the presence of Onychodus in the Huron Member of the Ohio Shale has implications for bottom water oxygenation in this black shale unit.Sageman et al. (2003) and Schieber (2003) provided evidence for oxygenation of water near the sediment-water interface.To support the presence of Onychodus and its prey species, some oxygenation of bottom waters would have been required.

Locality: Upper Freeport Coal, Linton, Ohio
Seven species discussed here were described from specimens collected from a thin layer of cannel coal underlying the main seam of the Upper Freeport Coal of the Allegheny Group (Upper Carboniferous: Pennsylvanian), at the Diamond Coal Mine in Linton, Jefferson County, Ohio, U.S.A. (Newberry, 1871(Newberry, :41, 1872:39:39; see also Hook & Ferm, 1985;Orton, 1884;Witzmann et al., 2017).Newberry (1871:41;1872:39) reported the thickness of the cannel coal layer as 4 cm, whereas, much later, Hook and Ferm (1985:104) indicated that the cannel coal averages 30 cm in thickness in the vicinity of the Diamond Coal Mine.The underground mine was located about 1 km Babcock-Paleozoic vertebrates from Ohio Lagerstätten (e2308621-2) southwest of the mouth of Yellow Creek, where it empties into the Ohio River (Hook & Baird, 1986;Hook & Ferm, 1985).
The Orton Museum has primary types of seven vertebrate species that Newberry (1857Newberry ( , 1873aNewberry ( , 1874Newberry ( , 1875aNewberry ( , 1875b) ) described from the Linton deposit.Some species erected by Newberry, including some reported here, have been synonymized.Hook andBaird (1986, 1988) recognized 12 species of chondrichthyan, actinopterygian, and sarcopterygian fishes from the Linton Lagerstätte.Babcock (2024) noted that two chondrichthyan species that Newberry (1857, 1875a) named from Linton were junior homonyms of Orthacanthus gracilis (Giebel, 1848) from the Carboniferous of Germany, and published replacement names for them.The syntypes of Orthacanthus adamas Babcock, 2024, a replacement name for Orthacanthus gracilis Newberry, 1875a, are in the Orton Museum.
Although the Orton Museum received a significant portion of Newberry's vertebrate collection from Linton, specimens that he collected or examined have been widely distributed.As shown by Baird (1962b) and Hook and Baird (1986), his specimens, including ones described by Cope (1873Cope ( , 1875)), are also known to have been deposited in the American Museum of Natural History, Columbia College School of Mines (now in the American Museum of Natural History), Princeton University (now in the Yale Peabody Museum), and the U.S. National Museum of Natural History.Newberry disseminated Linton specimens among contemporaries (Hook & Baird, 1986), and some of these specimens may have ended up in museums, universities, or colleges.The Royal Scottish Museum, the Field Museum of Natural History, and the Humboldt Natural History Museum, Germany, have specimens that may have been acquired from Newberry.At least seven other museums hold Linton material (e.g., see Westoll, 1944;Baird, 1962b;Baum & Lund, 1974;Panchen, 1977;Hook & Baird, 1986;Lowney, 1986;Witzmann et al., 2017).In the years since the Diamond Coal Mine ceased operations, the spoil piles have been collected by numerous field parties and a number of publications have resulted from these efforts (e.g., Baum & Lund, 1974;Holmes & Baird, 2011;Hurey et al., 2017;Witzmann et al., 2017).

MATERIALS AND METHODS
Specimens representing eight primary types, seven of fishes, and one of a tetrapod (anthracosaur) originally described as a fish, are illustrated here (Figs.1-3).Most were originally identified as types in the Orton Museum's type collection, but search of the vertebrate collections resulted in identification of one previously unrecognized syntype (Fig. 1B) and the counterpart of a holotype (Fig. 3A).
Specimens were photographed with a Canon Rebel Xsi, and photographs were processed in Adobe Photoshop.Most studied fish specimens from the Linton Lagerstätte were coated with shellac as a preservative in earlier years.In most cases the shellac is thin and relatively easy to photograph through (e.g., Figs.3A, B), but one specimen (Fig. 3C) is coated with a thick layer and difficult to image well.In all cases, the shellac introduces light reflection and refraction.Efflorescence in the coal beneath the shellac also produces distracting coloration (e.g., Fig. 3C).Specimens were photographed without preparation.
Locality, Horizon, and Age-Upper Freeport Coal of the Allegheny Group (Upper Carboniferous: Pennsylvanian), at the Diamond Coal Mine of Linton, Jefferson County, Ohio.
Orthacanthus gracilis Newberry, 1875a, was erected on the basis of two specimens (Fig. 1A, B), both dorsal spines, illustrated as a composite (Newberry, 1875a:pl.59, fig.7; reprinted 1875b:pl.59, fig.7).Both specimens were originally numbered OSU 4467.The original published description and composite illustration (Newberry, 1875a:56; pl.59, fig.7) show that both specimens were intended to be syntypes, although Morningstar (1924) listed the larger, more complete spine, now OSU 4467A (Fig. 1A), as the "holotype" in a catalog of type specimens.The word "cotype" is printed on the label accompanying the two specimens.Morningstar's (1924) reference to OSU 4467A as the "holotype" does not satisfy the provisions of Article 74.6 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 2000) for fixation of a lectotype by inference of holotype because the original description and illustration require that there were syntypes.OSU 4467A formed the basis for part of Newberry's figure, but it lacks the distinct denticles referred to in the description and appearing in the figure.The denticles are distinct in the smaller, broken specimen (OSU 4467B; Fig. 1B), and a paper label attached to the slab, handwritten by Newberry, states this to be a "Summitshowing denticles."The larger specimen, OSU 4467A, has the word "Type," handwritten by Newberry on a paper label attached to the back of the slab.Herein, OSU 4467A is designated as the lectotype of the species, and OSU 4467B is designated as the paralectotype.
Locality, Horizon, and Age-Upper Freeport Coal of the Allegheny Group (Upper Carboniferous: Pennsylvanian), at the Diamond Coal Mine of Linton, Jefferson County, Ohio.
Eurylepis striolatus Newberry, 1873a, was erected on the basis of a single, small specimen.The holotype (OSU 4508) is illustrated here for the first time (Fig. 2C).A paper label attached to the specimen, signed "JSN," states this is to be the "Type."Westoll (1944) synonymized Eurylepis striolatus Newberry, 1873a, with H. (H ).corrugata (a new combination), although there is no indication that Westoll observed the holotype.The holotype of E. striolatus (Fig. 2C) lacks the cranial plates, so it is not possible to determine whether the species has the "corrugations" characteristic of H. corrugata.Newberry (1873a:355) stated that the posterior margins of the scales "seem not to be serrated, but they are perhaps slightly crenulated by shallow scallops."The holotype of E. striolatus (Fig. 2C) is much smaller than the holotype of O. corrugata (Fig. 2A) but examination of the lateral scales indicates both specimens have similar serrations.This observation is consistent with Westoll's (1944) interpretation of E. striolatus as a junior synonym of H. corrugata.The "scallops" described by Newberry (1873a) in E. striolatus may be serrations, with the descriptive difference being a function of proportional differences in the sizes of specimens he studied.PARAHAPLOLEPIS Westoll, 1944 Type Species-Mecolepis tuberculatus Newberry, 1857.Notes-Mecolepis tuberculatus Newberry, 1857, is the type species of the subgenus Haplolepis (Parahaplolepis) Westoll, 1944. Lowney (1986:944) raised Parahaplolepis to genus level.
Notes-Mecolepis insculptus Newberry, 1857, was erected on the basis of a series of syntypes (illustrated by Newberry, 1873a:97, pl. 38, figs. 2, 2a-2c, 3, 3a).According to Baird (1962a:27-28) three specimens, OSU 4514, AMNH 448-G (now AMNH FF 448), and PU 1767 (now reposited at YPM) are labeled "Eurylepis insculptus N. Type" in Newberry's handwriting.None of the specimens agrees with Newberry's (1873, pl. 39, fig.2) illustration.The illustration may be a composite (compare with Orthacanthus adamas, discussed above), or may be based on another specimen.Westoll (1944:32) reported AMNH 448-G (now AMNH FF 448) as the "type," but overlooked Morningstar's (1924:24) previous statement that OSU 4514 is the "holotype."Morningstar (1924), the first reviser of this species, was apparently unaware that M. insculptus was based on a syntypic series, so reference to a "holotype" was mistaken.This act also cannot be considered designation of a lectotype, as it does not satisfy the provisions of Article 74.6 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 2000) for fixation of a lectotype by inference of holotype because Newberry's specimen labels clearly indicate the designation of at least three syntypes.To stabilize the nomenclature of M. insculptus, OSU 4514 is designated the lectotype of the species.Other specimens labeled by Newberry as types of M. insculptus are designated here as paralectotypes.
MICROHAPLOLEPIS OVOIDEA (Newberry, 1857) (Fig. 3A, B) Notes-Mecolepis ovoideus Newberry, 1857, was erected on the basis of a single, small specimen.According to Newberry (1857:97), this species is characterized by a small, ovoid body, corrugated and finely granulated cranial plates, threadlike corrugations on the opercula and lower parts of the head, granulated scales in the anterior abdomen, and serrated scales laterally.The holotype (part, OSU 4563, Fig. 3B; counterpart OSU 4496, Fig. 3A) was redescribed with the new combination Eurylepis ovoideus (Newberry) and illustrated for the first time by Newberry (1873a:351, pl. 39, fig. 1).A paper label attached to the part, handwritten by Newberry, identifies the specimen as "E.ovoideus N," "Type," and a similar label attached to the counterpart states this is to be the "Type Specimen."The counterpart, which is more complete, was not cited by Morningstar (1924), and is reported here for the first time.The counterpart generally agrees with the first illustration (Newberry, 1873a:pl. 39, fig. 1), although there has been some artistic restoration.A thin coating of shellac is present on the specimen.
Locality, Horizon, and Age-Upper Freeport Coal of the Allegheny Group (Upper Carboniferous: Pennsylvanian), at the Diamond Coal Mine of Linton, Jefferson County, Ohio.
Notes-Mecolepis lineatus Newberry, 1857 was erected using a single specimen.According to Newberry (1857:97), this species is characterized by a robust, fusiform body, with threadlike ornamentation on the cephalic bones, concentric thread lines on the anterior abdominal scales and on the margins of the lateral scales, and lateral scales located low on the body.The holotype (part, OSU 4565A; counterpart, AMNH FF 454) was redescribed with the new combination Eurylepis lineatus (Newberry) and illustrated for the first time by Newberry (1873a:353, pl. 39, figs. 7, 7a).A paper label attached to the part (OSU 4565A), handwritten by Newberry, states this is to be the "Type," and it generally agrees with his illustration (Newberry, 1873a:pl. 39, fig.7), although the illustration is reversed.A thick coating of shellac is present on the specimen.The counterpart, AMNH FF 454, is less complete than the part; a plaster cast is reposited as OSU 4565B.SARCOPTERYGII Romer, 1955ONYCHODONTIDA Andrews, 1973sensu Ciudad Real, Mondéjar Fernández, Vidal, & Botella, 2022ONYCHODONTIDAE Woodward, 1891ONYCHODUS Newberry, 1857 Type Species-Onychodus sigmoides Newberry, 1857, Devonian of Ohio, U.S.A., by original designation (Newberry, 1857:297).
Notes-Newberry (1857:124) described the genus Onychodus without specifically designating a type species.As he originally included two species in the genus and later changed his identification of some specimens, interpretation of the designation of the type species is a confusing matter.Here, statements made by Newberry (1857; see also Newberry, 1870Newberry, , 1873aNewberry, , 1889) ) are construed to indicate that O. sigmoides Newberry, 1857, from the "Corniferous limestone" (Columbus-Delaware limestones) of Ohio, was intended to be the type species of Onychodus Newberry, 1857.
In describing the new genus Onychodus, Newberry (1857:124) stated "The detached teeth are frequently found in the Cliff limestone … ."The term "Cliff limestone" was a term for the strata later known as the "Corniferous limestone," and today recognized as the Columbus Limestone-Delaware Limestone succession.Two new species were originally included in Onychodus, O. hopkinsi Newberry, 1857 and O. sigmoides Newberry, 1857.Both species were described on the same page (Newberry, 1857:297), and O. hopkinsi preceded O. sigmoides.Although no statement of the occurrence of O. hopkinsi was given, under O. sigmoides, Newberry stated "These teeth are not uncommon in the Cliff limestone of Ohio." Further evidence that Newberry intended O. sigmoides to be the type species of Onychodus can be gleaned from his earlier and later publications.Before proposing the species-group Babcock-Paleozoic vertebrates from Ohio Lagerstätten (e2308621-7) name Onychodus sigmoides, he (Newberry, 1853:fig.2) illustrated two undetermined teeth from the "Corniferous Limestone" that clearly belong to O. sigmoides.A list of fossils from the Corniferous limestone of Ohio (Newberry, 1862:74), derived from his 1857 paper describing Onychodus, listed both O. hopkinsi and O. sigmoides but, afterward, Newberry (1873a:301-302, 1889:28) stated that his inclusion of O. hopkinsi in the list was erroneous.In another paper, Newberry (1870:18) stated that he named Onychodus from remains in the Corniferous limestone, but did not cite a species in connection with the generic name.Finally, Newberry (1873a:297) wrote that he erected the genus Onychodus in January 1857 to "receive certain conical, curved teeth found in considerable abundance in the Corniferous limestone."Elsewhere in the same paper, Newberry (1873a:301-302) noted that he recognized O. hopkinsi as occurring only in the "Chemung of New York" (see also Newberry, 1889:99-100).
Previous to Newberry's (1889) report, Orton (1878:628) stated in text that remains of Onychodus, and perhaps O. sigmoides Newberry, 1857, range through a considerable stratigraphic range in Franklin County, Ohio, from the "middle Corniferous" (Columbus Limestone) through the Huron Shale Member of the Ohio Shale.OSU 14154 is the only known specimen of Onychodus from the Huron Shale Member in collections made by the Geological Survey of Ohio prior to 1889, and Orton's (1878) reference to Onychodus from the Huron Shale probably pertains to this specimen.Andrews et al. (2006:200), Mann et al. (2017:235), and Mondéjar-Fernández (2020:574) mentioned O. ortoni, remarking that it is known from too little material to be considered a properly defined species.TETRAPODA Jaekel, 1909(fide Sues, 2019) ANTHRACOSAURIA Cope, 1875EMBOLOMERI Cope, 1885ANTHRACOSAURUS Huxley, 1863 Type Species-Anthracosaurus russelli Huxley, 1863, Carboniferous of Scotland.
Holotype-Tooth (palatal tusk), OSU 4500.Locality, Horizon, and Age-Upper Freeport Coal of the Allegheny Group (Upper Carboniferous: Pennsylvanian), at the Diamond Coal Mine of Linton, Jefferson County, Ohio.
Abbreviations-AMNH, American Museum of Natural History, New York, NY, U.S.A.; OSU, Orton Geological Museum, The Ohio State University, Columbus, OH, U.S.A.; PU, Princeton University collection, now incorporated as part of YPM, Yale Peabody Museum, New Haven, CT, U.S.A.