REVISION OF THE AQUATIC ERYOPID TEMNOSPONDYL GLAUKERPETON AVINOFFI ROMER, 1952, FROM THE UPPER PENNSYLVANIAN OF NORTH AMERICA

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1 ANNALS OF CARNEGIE MUSEUM vol. 81, number 1, PP DecembeR 2012 REVISION OF THE AQUATIC ERYOPID TEMNOSPONDYL GLAUKERPETON AVINOFFI ROMER, 1952, FROM THE UPPER PENNSYLVANIAN OF NORTH AMERICA Ralf WeRnebuRg Naturhistorisches Museum Schloss Bertholdsburg Schleusingen, Burgstrasse 6, D Schleusingen, Germany DaviD S berman Curator, Section of Vertebrate Paleontology, Carnegie Museum of Natural History 4400 Forbes Avenue, Pittsburgh, Pennsylvania, bermand@carnegiemnh.org ABSTRACT The taxonomic validity of the eryopid temnospondyl Glaukerpeton avinoffi Romer, 1952, from the Upper Pennsylvanian Conemaugh Group of North America is confirmed on the basis of a detailed restudy of two specimens: the holotype, based on a partial skull roof from Pittsburgh, Pennsylvania; and the skull and postcranial material of a second specimen from an approximately equivalent stratigraphic level in West Virginia that was originally referred to Eryops cf. E. avinoffi (Romer), but is reassessed here as G. avinoffi. This contradicts a previous redescription of the holotype of G. avinoffi as referable to Eryops Cope, A single unique feature, the presence of three large, fang-like tusks on the ectopterygoid, distinguishes Glaukerpeton from all other eryopids. A cladistic analysis was performed using 19 cranial and two postcranial characters to clarify the phylogenetic relationships between Glaukerpeton Romer, 1952, and the only other eryopids in which the skull anatomies are well known: the Late Pennsylvanian Early Permian Eryops and the Early Permian Onchiodon Geinitz, The resultant cladogram indicates that Eryops and Onchiodon share a more recent common ancestor than either does with Glaukerpeton. The position of Glaukerpeton as a basalmost taxon may indicate that the ancestry of Eryopidae predates the Late Pennsylvanian. The possession of lateral line sulci, ossified ceratobranchials, and a thinly ossified skull roof indicates a probable aquatic habitus of the adult Glaukerpeton. Reconstructions are presented for the first time of the holotypic skull roof in dorsal view and the cranium and mandible of the referred specimen in various views. Key WoRDS: aquatic habitus, eryopid relationships, revised diagnosis, taxonomic status Primarily on the basis of a partial, fragmented skull roof (CM 8539) from the Upper Pennsylvanian Conemaugh Group in Pittsburgh, Pennsylvania, Romer (1952) described the eryopid temnospondyl Glaukerpeton avinoffi. Despite a strong resemblance to the Lower Permian Eryops Cope, 1882, Romer (1952) argued that three features distinguish it from Eryops: absence of an interfrontal bone, smaller size, and a finer reticulate sculpturing pattern of the skull roofing bones. Vaughn (1958) reassigned Glaukerpeton Romer, 1952, to Eryops, arguing that the latter two features are of doubtful value at the generic level, noting that they may simply reflect an immaturity of the smaller holotypic skull compared to the much larger skulls of Eryops available to him. Most significantly, on the basis of a restoration of the broken and dislocated elements of the holotypic skull roof, he concluded that the interfrontal bone, a hallmark feature of Eryops, was present. Unfortunately, an illustration was not provided that would have allowed the opportunity to confirm or challenge this observation. Yet, Glaukerpeton was subsequently accepted widely as a junior synonym of Eryops, thus extending its geologic range from the Upper Pennsylvanian to the Lower Permian. The validity of Glaukerpeton was further confused with the description by Murphy (1971) of a well-preserved skull and some disarticulated postcranial bones of INTRODUCTION an eryopid (CMNH 11025) from the Upper Pennsylvanian Conemaugh Group of West Virginia, which he assigned to Eryops cf. E. avinoffi. However, as he noted, the skull is slightly smaller than the holotypic skull of Glaukerpeton and considerably smaller than adult species of Eryops, and the sculpturing of the skull roof consists of a fine reticular pattern of pits. Unfortunately, he was unable to confirm or deny the presence of an interfrontal bone. Further limiting a comparison with either genus, the specimen was illustrated with poor quality photographs only and without indication of anatomical features. More recently, Lucas et al. (2005) and Werneburg et al. (2010a) described a large eryopid skull from the Upper Pennsylvanian El Cobre Canyon Formation of New Mexico that clearly belongs to the genus Eryops. It possesses an interfrontal, as well as other distinctive features of the genus. Werneburg et al. (2010a) briefly discussed the taxonomic status of Glaukerpeton, declaring it to be a valid taxon. Therefore, the specimen of Eryops sp. from New Mexico is the first to document firmly the genus from the Pennsylvanian. The primary objectives of the present study of G. avinoffi are to 1) provide a detailed description of the skull and postcranial elements, 2) demonstrate its taxonomic distinction from Eryops, and 3) determine its relationship to Onchiodon Geinitz, 1861, and Eryops. The only other recognized eryopids, Actinodon Gaudry, 1866,

2 34 annals of carnegie museum vol. 81 Osteophorus Meyer, 1856, and Clamorosaurus Gubin, 1983, are excluded from the analysis of relationships, as they are currently being redescribed. In addition, reconstructions of G. avinoffi are presented for the first time that include the skull roof in dorsal view of the holotype CM 8535 and various views of the cranium and mandible of the referred specimen CMNH The following acronyms are used to refer to institutional repositories: AMNH, American Museum of Natural History, New York; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; CMNH, Cleveland Museum of Natural History, Cleveland, Ohio; FMNH, Field Museum of Natural History, Chicago, Illinois; MCZ, Museum of Comparative Zoology of Harvard University, Cambridge, Massachusetts; MMG, Museum für Mineralogie und Geologie, Dresden, Germany; NHMS, Naturhistorisches Museum Schloss Bertholdsburg, Schleusingen, Germany; MNHUB, Museum für Naturkunde der Humboldt-Universität, Berlin, Germany; NMMNH, New Mexico Museum of Natural History, Albuquerque, New Mexico; SMNS, Staatliches Museum für Naturkunde Stuttgart, Germany; TUBAF, Technische Universität Bergakademie Freiberg, Germany; USNM, United States National Museum, Washington, D.C. Anatomical abbreviations: a, angular; ahk, anterior humeral keel; ap, anterior palatal depression; ar, articular; bb, basibranchial; bo, basioccipital; c1 3, coronoids 1 3; cb1 4, ceratobranchials 1 4; cg, groove for carotid artery; cgr, clavicular groove; ch, choane; chy, ceratohyal; cl, clavicle; clr, clavicular recess; d, dentary; dcp, dorsal clavicular process; dcr, dorsal clavicular rod; delt, deltoid crest; d.sm, dorsal part of septomaxilla; ec, ectopterygoid; ect, ectepicondyle; ectn, ectepicondylar notch; ectr, ectepicondylar ridge; ent, entepicondyle; entr, entepicondylar ridge; eo, exoccipital; f, frontal; fgl, glenoid foramen; for, foramen; f.pq, paraquadrate foramen; fsgl, supraglenoid foramen; f.stp, stapedial foramen; gl, glenoid fossa; hum, humerus; humc, humeral head; ic, intercentrum; ifc. pt, postorbital part of infraorbital sulcus; ifc.sb, suborbital part of infraorbital sulcus; igb, intraglenoid buttress; j, jugal; la, lacrimal; ldp, latissimus dorsi process; lsr, lateral supraglenoid ridge; m, maxilla; md, mandible; mf, Meckelian fenestra; mfo, Meckelian foramen; n, nasal; nvii, cranial nerve VII; na, neural arch; occ.lam, occipital lamella; op, otic plate; p, parietal; pa, prearticular; pc, pleurocentrum; p.c, cultriform process of parasphenoid; pect, pectoral crest; phr, proximal humeral ridge; pl, palatine; pm, premaxilla; po, postorbital; poc, preopercular sulcus; pp, postparietal; prf, prefrontal; prsp, presplenial; ps, parasphenoid; psp, postsplenial; ps.p, basipterygoid process of parasphenoid; pt, pterygoid; ptf, postfrontal; q, quadrate; qj, quadratojugal; r, rib; rac, radial condyle; sa, surangular; sc, scapulocoracoid; sctor, scapular torus (scapular blade); sgb, supraglenoid buttress; sgf, supraglenoid fossa; shr, scapulohumeral ridge; sm, septomaxilla; soc, supraoccipital sulcus; sph, sphenethmoid; sq, squamosal; ssf, subscapular fossa; st, supratemporal; stp, stapes; sup, supinator process; t, tabular; ulc, ulnar condyle; vd.sm, ventrally directed part of septomaxilla; v, vomer. Measurement abbreviations used in Fig. 5 and Appendices 1 3: as w, anterior width of skull at level of maxillapremaxilla sutures; H l, postorbital midline length of skull from level of posterior margins of orbits; H w, postorbital width of skull between lateral margins of supratemporals; IN w, minimum internarial width; IO w, minimum interorbital width; Ju w, transverse width of jugal at maximum lateral lacrimal extent of orbit; La l, maximum length of lacrimal; La w, maximum transverse width of lacrimal; ms w, midlength width of skull at midlength level of orbits; O l, maximum length of orbit; p, number of dermal skull pits or valleys per in 2 mainly from frontal and jugal at midlength level of orbits; PO l, preorbital midline length of skull from level of anterior margins of orbits; Po l, maximum posterior length of postorbital from posteriormost extent of orbit; Po w, maximum transverse width of postorbital at contribution to orbital margin; ps w, maximum posterior width of skull at level of posterolateral margins of cheeks; S l, midline skull length; Th l, length of tabular horn region between levels of posterior tabular corner and occipital midline margin; W w, maximum transverse width of cheek from lateral margin of supratemporal anterior to otic notch. SYSTEMATIC PALEONTOLOGY Order Temnospondyli Zittel, 1888 Family Eryopidae Cope, 1882 Genus Glaukerpeton Romer, 1952 Type species. Glaukerpeton avinoffi Romer, Glaukerpeton avinoffi Romer, 1952 (Figs. 1 18, 20) Glaukerpeton avinoffi Romer, Werneburg et al., 2010a. Eryops avinoffi: Vaughn, 1958; Moulton, Eryops cf. E. avinoffi: Murphy, Eryops avinoffi: Werneburg, Revised Diagnosis. An eryopid temnospondyl amphibian that can be distinguished from all other eryopids by the unique feature of three large, fang-like tusks on the ectopterygoid and from the eryopids Onchiodon and Eryops by combinations of the following primitive characters: 1) lateral line sulci present in adults as short, discontinuous segments on the prefrontal, postorbital, jugal, and squamosal; 2) number of dermal sculpturing pits/in 2 in interorbital area to midline skull length proportion in adults 2.6 to 3.3; 3) absence of strut-like ridges on skull roof; 4) skull-roof bones 30 50% thinner; 5) internarial width narrower than interorbital width when expressed as a proportion to skull length; 6) prenarial, midline length of premaxilla about 7% of midline skull length; 7) postorbital width of skull table to midline skull length proportion 0.56; 8) width of cheek to midline skull length proportion

3 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 35 Fig. 1. Holotype of Glaukerpeton avinoffi, Romer, CM A, photograph; and B, illustration of skull roof with fragments of palate and mandible in dorsal view. C, reconstruction of skull roof in dorsal view with outline based on Figs. 3 and 11.

4 36 annals of carnegie museum vol. 81 Fig. 2. Skull and right mandible of Glaukerpeton avinoffi, referred specimen CMNH 11025, in dorsal view. A, photograph; and B, illustration.

5 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 37 Fig. 3. Reconstruction of skull of Glaukerpeton avinoffi, CMNH 11025, in dorsal view ; 9) maximum length of orbit to midline skull length proportion ; 10) interfrontal absent; 11) margin of pineal foramen sculptured; 12) anterior extent of parietals to level of posterior margin of orbits; 13) contribution of supratemporal to dorsal rim of otic notch approximately 20 40%; 14) paraquadrate foramen visible in lateral, dorsal, and occipital views of skull; 15) internal or basal process of pterygoid suturally overlaps the ventral surface of parasphenoidal plate of basipterygoid process; 16) free lateral margin of proximal region of palatal ramus of pterygoid extends smoothly anterolaterally except for slight angular expansion into adductor chamber at contact with ectopterygoid; 17) longitudinal distance between levels of occipital midline margin of skull table and posterior margin of cheek less than 18% of midline length of the skull; 18) ceratohyal and ceratobranchials ossified in adults; 19) marginal dentitions of skull and mandible lack caniniform teeth; 20) supinator process directed normal to humeral shaft; and 21) angle between supraglenoid buttress and anterior margin of scapular blade less than 90º. Holotype. CM 8539, consisting almost entirely of the greater portion of skull roof with many of the bones fragmented and dislocated, and some fragments of palate and mandible. Holotypic Horizon and Locality. Pittsburgh Limestone Member, at the top of the Upper Conemaugh Group, Upper Pennsylvanian, within Pittsburgh City, Pennsylvania (locality 1 of Moran 1952). Referred Specimen. CMNH 11025, well-preserved, essentially complete skull that includes both mandibles, pectoral girdle, and limb and vertebral elements. Referred Horizon and Locality. Siltstones at top of the Morgantown Sandstone Member, mid-portion of Conemaugh Group, Upper Pennsylvanian, at road cut along U.S. Route 19 between 1 and 1.1 mile southeast of southern end of Elk River bridge at Sutton, Braxton County, West Virginia (Murphy 1971). Possible Referable Specimens. CM 8591, postparietal, other skull fragments, and Eryops-like phalanx (Romer 1952) from the same locality as that of the holotype; CM 1944, 1945, 1947, and 1948, dorsal vertebra and rib fragments, neural arch and caudal vertebra, neural spine, and vertebral fragments, respectively, from approximately the same level as the holotypic horizon (locality C of Moran 1952), the Round Knob Shale Member of the Conemaugh Group at Pitcairn, Pennsylvania. These elements were originally figured and referred to Eryops by Case (1908), but considered by Romer (1952) as more probably pertaining to Glaukerpeton; CM 8538, cranial bone fragments with a sculpturing pattern comparable to that in the holotype, vertebrae and ribs, partial ulna and femur, and ventral dermal scales (Romer 1952) from a nearby locality 2 of Moran (1952) and at practically the same horizon as that of the holotype. Due to the taxonomic uncertainty of these

6 38 annals of carnegie museum vol. 81 Fig. 4. Skull of Glaukerpeton avinoffi, referred specimen CMNH 11025, in lateral view. A, photograph; B, illustration; and C, reconstruction of skull. specimens, however, they are not included in this study, which relies solely on the holotype CM 8539 and referred specimen CMNH DESCRIPTION AND COMPARISONS Skull Roof The holotypic skull of G. avinoffi, CM 8539, consists almost entirely of the greater portion of the roof in dorsal view with many of the bones fragmented, incomplete, and dislocated (Figs. 1A B). However, a tentative reconstruction of CM 8539 (Fig. 1C) has been possible using the well-preserved, nearly complete skull of the referred specimen of G. avinoffi, CMNH 11025, as a guide. Although many of the skull roofing bones of CMNH are displaced slightly or marginally incomplete or unexposed (Figs. 2A B), an accurate reconstruction of the skull roof in dorsal view has been possible (Fig. 3). The skull is little disturbed in left lateral view (Figs. 4A B), also allowing an accurate reconstruction in this view (Fig. 4C). Unless otherwise stated, the description of G. avinoffi that follows is based on the referred specimen CMNH In addition, comparisons with the skulls of the eryopids Onchiodon and Eryops are based, unless stated otherwise, almost entirely on descriptions of the Lower Permian Eryops megacephalus Cope, 1882, by Sawin (1941), and Onchiodon thuringiensis Werneburg, 2007, by Werneburg (2007), and their repeated citation in the text below has been eliminated as an unnecessary redundancy. The Glaukerpeton skulls CM 8539 and CMNH have nearly the same estimated midline lengths of 23.0 and 22.8 cm, respectively. Both skulls can be distinguished from those of Onchiodon and Eryops on the basis of several proportional differences, the measurements of which are based, as examples, on two adult specimens each of Onchiodon and Eryops. Almost all of the proportions discussed below are presented in Appendix 1, whereas Fig. 5 illustrates how the skull-roof measurements were taken. Glaukerpeton, Onchiodon, and Eryops exhibit a progressive

7 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 39 Fig. 5. Diagram of generalized eryopid skull roof illustrating key to positions and abbreviations of measurements used in text. increase in the longitudinal distance between the levels of the occipital midline margin of the skull table and posterior margin of the cheek, which when expressed as a percentage of the midline length of the skull yields values of 16, 18, and a maximum 29%, respectively. Differences in this feature are perhaps more dramatically expressed as the proportion between the longitudinal distances from the level of the anterior margin of the otic notch to the posterolateral corner of the tabular and to the level of the posterior margin of the skull cheek, which yields decreasing values of 0.86, 0.58, and 0.37, respectively. Possibly related to these decreasing values is a marked decrease in the contribution of the supratemporal to the dorsal margin of the otic notch, which in Glaukerpeton and Onchiodon is about 39 and 18%, respectively, whereas in Eryops the supratemporal contribution varies from less than 10% to being excluded entirely from the otic notch by a narrow squamosal-tabular contact. In dorsal view the lateral margin of the skull of Glaukerpeton describes a wide parabolic curve with a broad, bluntly rounded snout and a proportionally expanded postorbital region. In these features it closely approaches the larger adult skulls of Onchiodon and Eryops. However, although the width of the postorbital skull table is greater in Glaukerpeton than in Onchiodon and Eryops, the cheek region is narrower, resulting in a net posterior skull width that is similar in all three eryopids. The orbits in Glaukerpeton are proportionally larger when compared to those of the larger skulls of Onchiodon and Eryops, which likely reflects negative allometric growth of the orbit. However, the internarial distance in Glaukerpeton is proportionally much smaller than the interorbital distance, which distinguishes it from Onchiodon and Eryops. The skull roofing bones in Glaukerpeton are relatively very thin, about 30 50% thinner than those in Onchiodon and Eryops, and, as such, more comparable to those of the stem-stereospondylomorphs Sclerocephalus Goldfuss, 1847, and Glanochthon Schoch and Witzmann, 2009 (Goldfuss 1847; Schoch and Witzmann 2009b).

8 40 annals of carnegie museum vol. 81 Fig. 6. Right naris of Glaukerpeton avinoffi, referred specimen CMNH 11025, indicating short dorsal, sculptured portion (d.sm) and large, smooth, ventrally directed portion (vd.sm) of septomaxilla. An accessory internasal, present at the sutural intersection of the premaxillae and nasals in Onchiodon thuringiensis, is absent in all other eryopids. A traceable sutural outline, even partial, of an interfrontal, a diagnostic feature of Eryops, is not detectable in either the holotype or the referred specimen of Glaukerpeton. In Eryops the interfrontal (internasofrontal of Sawin 1941) occupies the midline of the skull roof, where it separates approximately the posterior thirds of the paired nasals and the anterior halves of the paired frontals. In the holotype of Glaukerpeton the medial margins of the posterior half of the right nasal and approximately the anterior half of the frontal are straight and extended along or very close to the midline suture of the skull, thus strongly suggesting the absence of an interfrontal. Similarly, in Glaukerpeton CMNH the nasal-frontal sutural contact on both sides of the skull is preserved and extends to or very close to the skull midline without any indication of accommodating an interfrontal. Although Romer (1952) also reached the conclusion that the holotype lacked an interfrontal, Vaughn (1958) and Murphy (1971) were unable to provide irrefutable evidence of its presence or absence in Glaukerpeton. Its absence is clearly demonstrated in specimens of Onchiodon. The frontals in CMNH are proportionally as wide as those in Eryops, although they may appear narrower due to the displacement of the left frontal over the medial margin of the right frontal and the displacement of the left prefrontal over the lateral margin of the left frontal. The lengths of the frontals in Glaukerpeton also appear to be variable and more elongated anteriorly in the holotype than in CMNH 11025, and yet, the combined lengths of the frontals and nasals in both specimens are subequal. The width of the nasals in Glaukerpeton is proportionally narrower than those in Eryops, which is reflected in the shorter distance between the narial openings and the anteromedial narrowing of the lacrimals. The bluntly rounded snout in Glaukerpeton is a feature shared with Onchiodon and Eryops, as is the narrow, distinct alary process of the premaxilla overlapping the lateral margin of the nasal. The prenarial midline length of the premaxilla, as in Onchiodon (Werneburg 2007: figs. 2a, 7), is about 7% of the midline length of the skull, which is typical of temnospondyls. In Eryops, on the other hand, the prenarial midline length of the premaxilla is noticeably greater, as is well documented in E. megacephalus (Sawin 1941: fig. 1 and table on p. 410; Werneburg 2007: fig. 6), which ranges from 18 to 25% of the midline length of the skull. The greater prenarial length of the premaxillae in Eryops is reflected in a shallow, broadly concave narrowing of the skull margin at the level of the nares. The naris has an oval to triangular outline. Crescentshaped septomaxillae, preserved only in CMNH (Fig. 6), lie along the ventral margin of the naris proper and contact the premaxilla, maxilla, and lacrimal, blocking completely the latter two bones from the opening. Therefore, the narial opening is restricted dorsal to the septomaxilla, where it is bordered dorsally by the premaxilla and nasal. The anterior two-thirds of the septomaxilla are smooth, whereas the posterior third is sculptured in a pattern like that of the adjoining maxilla, with which it is smoothly incorporated into the skull roof. In Onchiodon, on the other hand, the septomaxillae are completely sculptured and incorporated into the skull roof (Boy 1990). The entire septomaxilla in Eryops is smooth with one known exception, Eryops sp., AMNH 4673, from the Lower Permian Arroyo Formation, which exhibits a sculptured posterior portion. The septomaxilla in Eryops also differs from those in Glaukerpeton and Onchiodon in occupying almost the entire naris proper, restricting the narial opening to two small openings, one at the anterior margin that is bordered by the premaxilla and one at the posterior margin that is bordered by the nasal. In all three eryopids the lacrimal has roughly an elongated diamond outline and is separated from the orbit by a dorsal antorbital process of the jugal that contacts the prefrontal. However, in Glaukerpeton and Onchiodon the dorsal antorbital process of the jugal is approximately half the width of that in Eryops and extends to the midheight level of the orbit compared to the prefrontal forming nearly the entire anterior orbital margin in Eryops. The jugal in Glaukerpeton is proportionally slightly narrower than

9 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 41 Fig. 7. Posterior portion of left cheek of Glaukerpeton avinoffi, referred specimen CMNH A, dorsolateral; B, posterodorsal; and C, ventral views. Fig. 8. Interorbital region of skull roof of Glaukerpeton avinoffi. A, holotype CMNH 8539; and B, referred specimen CMNH 11025, demonstrating their similar fineness of sculpture pattern.

10 42 annals of carnegie museum vol. 81 Fig. 9. Skull with right mandible of Glaukerpeton avinoffi, referred specimen CMNH 11025, in ventral view. A, photograph; and B, illustration. those in Onchiodon and Eryops, but in all three genera the jugal has a narrow, anterior process-like extension that extends between the lacrimal and maxilla to the posterior margin of the naris. The postorbital is triangular in outline, but slightly variable in width, depending on the medial encroachment of the postfrontal. The postfrontal and

11 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 43 prefrontal clearly contact one another in all three eryopids, but their dorsal orbital processes are proportionally slightly wider in Onchiodon and Eryops, with a slightly farther anterior contact. In Glaukerpeton and Onchiodon the parietals extend anteriorly to the level of the posterior margin of the orbits, whereas in Eryops they extend nearly to the midlength level of the orbits. Only in Onchiodon thuringiensis is the pineal foramen bordered by a band of smooth bone. The supratemporal in all three eryopids is broadly rectangular with a greater longitudinal dimension. The postparietals in Glaukerpeton, as in Onchiodon, are narrowly exposed along the occipital rim of the skull table and gradually narrow laterally to a very narrow contact with the triangular tabular occupying the posterolateral corner of the skull table. The skull-table exposure of the postparietals in Eryops is less expanded laterally and has a broader contact with the tabular. Together, both bones in all three genera form a deeply concave occipital margin of the skull table from which an occipital flange extends a short distance ventrally. The transverse widths of the squamosal, jugal, and in particular the quadratojugal in Glaukerpeton are proportionally much narrower than those in Onchiodon and Eryops, which accounts for the proportionally narrower width of the cheek region in the former. In Glaukerpeton the exposure of the quadrate on the occipital surface of the cheek (Figs. 3, 7) consists of a narrow, dorsally elongate process that is directed anteromedially between the squamosal and the quadrate ramus of the pterygoid. At the ventral margin of the dorsal process is a well-developed, boss-like protuberance. In Eryops there is an identical exposure of the quadrate, but the quadrate boss may be absent, whereas in Onchiodon the dorsal process of the quadrate is not exposed except for the ventral area that includes the boss, which is weakly developed. In Glaukerpeton a paraquadrate foramen near the posterolateral margin is visible in lateral, dorsal, and occipital views of the skull roof, whereas in Onchiodon and Eryops it is visible only in ventral view of the skull. An exception is Eryops sp., NMMNH P-46379, from the Late Pennsylvanian of New Mexico in which two small and a larger paraquadrate foramen at the posteroventral margin of the quadratojugal are visible in dorsal and lateral views of the skull roof (Werneburg 2010a). The presence of a paraquadrate foramen in Glaukerpeton, Onchiodon, and Eryops contradicts the statement by Yates and Warren (2000) that, whereas it is present in stereospondyls and dvinosaurians, it is absent in all eryopids. Furthermore, the presence of this character in eryopids contradicts its use by them as uniting a large clade of aquatic temnospondyls they termed Limnarchia. In contrast to Onchiodon and Eryops, only in Glaukerpeton are lateral line sulci present, but they are limited mainly to the circumorbital bones, where they are preserved as short, discontinuous segments. These are traceable variously in both specimens as parts of the supraoccipital sulcus on the prefrontal and postfrontal, parts of the infraorbital sulcus on the postorbital and suborbital area of Fig. 10. Ventral view of braincase with some associated elements of Glaukerpeton avinoffi, CMNH the jugal, and a short segment of the preopercular sulcus on the squamosal (Figs. 1 4). Possibly related to this feature is the thinness of the skull roofing bones in Glaukerpeton. Dermal Sculpturing The dermal sculpturing of the dorsal skull roof of Glaukerpeton is very characteristic of temnospondyls: a dense, reticulated pattern of small pits and valleys separated by narrow ridges (Figs. 1, 2, 4, 8). The dermal sculpture pattern of the skull roof in eryopid species, however, has been described alternately in subjective generalities, as either consisting of a dense, fine, or coarse sculpture pattern. In order to determine the utility of the sculpture pattern in diagnosing G. avinoffi, Vaughn (1958) compared the holotypic skull with an unquestionably Eryops skull, USNM 21860, of similar size. He recorded about 60 pits/in 2 in

12 44 annals of carnegie museum vol. 81 Fig. 11. Reconstruction of skull and visceral skeleton of Glaukerpeton avinoffi, CMNH 11025, in ventral view. the same region of the frontal in both specimens, which convinced him that there is no basis for differentiation of the two genera using sculpture density. He also recorded 25 pits/in 2 in the same area of the frontal in a larger but average-sized specimen of E. megacephalus, USNM 6721, leading him to suggest that the relationship of greater sculpture fineness and smaller skull size may simply reflect immaturity. Murphy (1971) quantified the sculpturing pattern in G. avinoffi, CMNH 11025, as pits/in 2 on the right postfrontal at the midlength level of the orbit. Despite the fact that it is difficult to quantify the degree of fineness or coarseness of the dermal sculpturing and that using this character to distinguish between taxa may be compromised to some degree by growth stage, evolutionary trends, quality of preservation, or environmental factors, the measurements presented here suggest otherwise. In Appendix 2 the density of the sculpture pattern in Glaukerpeton, Onchiodon, and Eryops specimens is quantified as the number of pits/in 2 on the frontal and jugal, typically well-preserved bones in eryopid skulls, and as a proportion of those counts to skull length. What is immediately obvious in Eryops specimens with skull lengths ranging from 10.0 to 43.5 cm is an ontogenetic trend toward greater coarseness of the sculpturing. Most revealing and germane to this discussion, however, is if the same measurements are applied to the two Glaukerpeton specimens with skull lengths of about 23.0 cm, they exhibit a fineness of sculpturing closely approaching only juvenile Eryops specimens with skull lengths of 10.0, 13.0, and 18.5 cm. This strongly suggests that the fineness or coarseness of sculpturing is a valid character difference between Glaukerpeton and Eryops. Even if a greater ontogenetic skull-length series was available for Glaukerpeton and exhibited the same trend toward greater coarseness of sculpturing with size as in Eryops, the proportion of fineness or coarseness of sculpturing to skull length would still be sufficient to distinguish between the two genera. A coarser pattern is also present in the single large skull (length 28.5 cm) of a late adult stage of Onchiodon (Appendix 2) in which this feature can be measured (frontal only), suggesting that it falls within the same ontogenetic rate of coarsening with changes in growth/size as in Eryops. Possibly related to the finer or coarser pattern of sculpturing in eryopids, in Glaukerpeton there is an absence of large,

13 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 45 Fig. 12. Skull of Glaukerpeton avinoffi, referred specimen CMNH A, anterior; and B, posterior views. strut-like ridges of the skull roof seen in Onchiodon and Eryops. A proportionally large variability of dermal sculpture pattern also occurs in the stem-stereospondylomorphs Sclerocephalus and Glanochthon from the Early Permian of Germany (Werneburg 1992; Schoch and Witzmann 2009a, 2009b), but this has not been related to changes in growth. Palate and Braincase The palate and endocranium are nearly complete, little disturbed, and well exposed in ventral view in CMNH (Figs. 9, 10) and can be confidently reconstructed (Fig. 11). The interpterygoid vacuity in Glaukerpeton is much more expanded at its midlength than those in Onchiodon and Eryops. As in Onchiodon and Eryops, the ventral surfaces of the pterygoid and ectopterygoid exhibit a strutting pattern of well-developed ridges that presumably provided greater mechanical strength to the palate. In Glaukerpeton two ridges on the anterior palatal ramus of the pterygoid extend anteriorly and slightly laterally onto the ectopterygoid and end at the palatal tusks, and a third extends anteriorly along the anterior half of the medial margin of the palatal ramus. The palatal ridges in Eryops are greatly subdued and difficult to follow, but appear to exhibit a pattern very similar to those in Glaukerpeton, whereas in Onchiodon a slightly different pattern of the palatal ridges is exhibited. Two ridges extend posteriorly from the posterior margin of the ectopterygoid tusks, one nearly the entire length of the bone that lies just medial and parallel to the skull margin and the other posteromedially and a short distance onto the palatal ramus of the pterygoid. A third ridge extends along the posterior half of the medial margin of the palatal ramus of the pterygoid and after a short break anteriorly is continued a short distance onto the vomer to the posteromedial margin of the choana. In Glaukerpeton the medial margin of the choana posterior to the vomerine tusks is bordered by a pronounced, narrow ridge. A similar but wider ridge

14 46 annals of carnegie museum vol. 81 Fig. 13. Right stapes of Glaukerpeton avinoffi, CMNH A, photograph in posterior view; and B, C, illustrations in ventral and posterior views. is present in Onchiodon and Eryops, although in the latter it is continued as a much narrower ridge along the lateral margin of the alveoli of the vomerine tusk. In all three eryopids the vomerine ridge medially bordering the choanae is continued abruptly medially across the palate at the level of the vomerine tusks, creating shallow, dorsally depressed areas anterior and posterior to the ridge. The dorsally depressed area anterior to the transverse ridge on the vomers is divided by a midline ridge into paired anterior palatal fossae that are bordered anteriorly by the dental shelf of the premaxilla. Whereas in Glaukerpeton the anterior palatal fossae are restricted to the premaxillae, in Onchiodon they are expanded a short distance farther posteriorly onto the vomers. In Eryops their posterior extent onto the vomers is even much greater, so that their size is approximately triple and doubles those in Glaukerpeton and Onchiodon, respectively. The process-like posterolateral corner of the vomer in Glaukerpeton and Onchiodon extends a short distance between the pterygoid and palatine, though much narrower in the latter, whereas in Eryops the process is much longer and vermiform. In Onchiodon and Eryops the choanae are subrectangular with the widths of the anterior and posterior margins being essentially equal, whereas in Glaukerpeton the width of the posterior margin is about twice that of the anterior margin. The anterior margin of the choana in all three eryopids exhibits a narrow, short, V-shaped notch of variable development lateral to the alveolus of the vomerine tusks. This feature is also seen in the Eryops sp., NMMN-P46379, from the Upper Pennsylvanian of New Mexico (Werneburg et al. 2010b). The free lateral margin of the basipterygoid region of the pterygoid in Glaukerpeton and Eryops exhibits a low, angular, medial expansion adjacent to its contact with the ectopterygoid that projects into the adductor chamber. In Onchiodon, on the other hand, the entire, free, lateral margin of the pterygoid is greatly expanded into a right-angled projection. The articular condyle of the quadrate in all three eryopids is transversely expanded and divided into a pair of condylar facets in which a smaller, lateral, suboval condyle is separated narrowly by a narrow channel from a larger, medial, anteromedially elongated oval condyle. A narrow, notchlike channel that separates the quadrate condyle from the posterior end of the ventral margin of the quadratojugal in Glaukerpeton and Onchiodon is absent in Eryops. In Glaukerpeton and Onchiodon the basicranial union is formed by the internal or basal process of the pterygoid suturally overlapping the ventral surface of the anterolateral corner of the parasphenoidal plate, whereas in Eryops a short, stout, laterally projecting basipterygoid process of the braincase unites with the internal process of the pterygoid in a near vertical interdigitating suture. The narrow cultriform process of the parasphenoid in Glaukerpeton CMNH is essentially complete. It is narrowest just anterior to its broad triangular base and gradually expands to twice its width distally, but then quickly narrows at its distal end to a pointed contact with the posterior margin of the midline union of the vomers. The process in Onchiodon is identical to that in Glaukerpeton except for terminating a short distance between the midline union of the vomers. On the other hand, the process in Eryops is proportionally twice the width of those in Glaukerpeton and Onchiodon. Furthermore, a short, proximal section of the process in Eryops exhibits slightly convex, bilateral expansions, but then continues anteriorly with a constant width until its distal end, where it narrows to a short, angular terminus that extends a short distance between the midline union of the vomers. In Eryops the cultriform process supports a wide, equal-sided, diamond-outlined sphenethmoid, whereas in Glaukerpeton and Onchiodon the anterolateral margins of the sphenethmoid are much shorter, giving it a lance-like outline. The main body of the parasphenoid, the parasphenoidal plate, in all three eryopids is indistinguishably fused with the basisphenoid, strongly flared posterolaterally posterior to the basipterygoid articulation, with the lateral margins curving dorsally to contact the lateral wall of the otic complex. However, in Glaukerpeton and Eryops the width of the parasphenoidal plate is greater than the length, whereas in Onchiodon the opposite is true. The basioccipital and exoccipitals in all three eryopids extend a short distance beyond the posterior

15 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 47 Fig. 14. Left mandible of Glaukerpeton avinoffi, CMNH A, B, C, photographs in lateral, medial, and dorsal views; D, E, F, illustrations of A C. margin of the parasphenoidal plate to contribute to a double, cotyliform facet of the occipital condyle, but, whereas the basioccipital and exoccipitals are fused into a single complex in Glaukerpeton and Onchiodon, a sutural separation persists in Eryops. The basioccipital-exoccipital complex is more expanded laterally in Onchiodon than in Glaukerpeton or Eryops, and the double, cotyliform facet of the condyle faces ventrally in Glaukerpeton and Onchiodon,

16 48 annals of carnegie museum vol. 81 Fig. 15. Axial elements of Glaukerpeton avinoffi, CMNH A, fourth and fifth neural arches in left lateral view and rib; B, isolated neural arch in left lateral view; and C, isolated pleurocentrum and intercentrum in anterior, posterior, and ventral views. more so in the former, whereas in Eryops it faces posteriorly. In Glaukerpeton a fine groove for the carotid artery closely parallels the medial margin of the facet for the internal process of the pterygoid, ending anteriorly near the foramen for cranial nerve VII. Visceral Skeleton Tentatively identified elements of the visceral skeleton in CMNH include the right ceratohyal and ceratobranchials 1 3 of the probable original four, whereas the basibranchial and right stapes can be confidently recognized (Figs. 9, 10, 12B, 13). A reconstruction of the visceral skeleton and its position relative to the skull is illustrated in Fig. 11. The nearly complete, anteroposteriorly oriented, rod-like basibranchial has been displaced slightly to the left of its midline position beneath the cultriform process of the parasphenoid. From a level slightly posterior to its midlength, it widens gradually anteriorly and posteriorly, much more so posteriorly. The basibranchial in Onchiodon labyrinthicus Geinitz, 1861, has been described as having a wider anterior portion at early stages of ontogenetic development, but the reverse in adult stages of development (Boy 1990). The reverse pattern of development has been described also in the stem stereospondylomorph Sclerocephalus haeuseri Goldfuss, 1847 (Boy 1988). With the exception of the stapes, no elements of the visceral skeleton are known in Eryops. In ventral view (Fig. 11) the ceratohyal and ceratobranchials 1 3 have much the same appearance as the basibranchial. All four elements exhibit a slight but gradual serial decrease in the length posteriorly and, with one possible exception, the width of the expanded ends; the exception being the probable narrow, proximal end of the ceratohyal, which is greatly expanded dorsoventrally. The ends of the first ceratobranchial are only slightly expanded dorsoventrally, whereas they are greatly expanded dorsoventrally in the second ceratobranchial; only remnants of the third ceratobranchial are preserved. The right stapes in CMNH (Fig. 13) is preserved nearly in its correct position (Figs. 9, 10). The footplate is only slightly widened and is pierced by the stapedial foramen, and the shaft is very thin and without a dorsal process. In these features the stapes is identical to those in Eryops (Sawin 1941: pl. 6) and the stem-stereospondylomorph Sclerocephalus (Schoch and Witzmann 2009a, 2009b). In Onchiodon, on the other hand, the stapes differs from that in Glaukerpeton only in the shaft being more widened anteriorly.

17 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 49 Mandible The right mandible in CMNH is complete, but because it is firmly articulated to the cranium in essentially its correct position (Fig. 9) its description is greatly limited. The detached left mandible, on the other hand, is well preserved except for missing a small anterior portion (Fig. 14). It exhibits a morphology that with few exceptions is very similar to those of most other eryopids. The prearticular is elongated anteriorly along its contact with the dorsal margin of the presplenial and postsplenial to a level a short distance beyond their contact. In Eryops megacephalus (Sawin 1941), Eryops grandis Langston, 1953 (Langston 1953), and Onchiodon labyrinthicus (Boy 1990), however, the prearticular ends a short distance posterior to the splenial-postsplenial contact. A similar sutural configuration to that in Glaukerpeton is seen in O. thuringiensis (Werneburg 2007). This corrects the misinterpretation by Werneburg (2007: fig. 9C) of the partial mandible of O. thuringiensis in which the presplenial was mislabelled as the postsplenial and the postsplenial as the angular. The fine ornamentation on the lateral surface of the mandible, which matches that of the skull roof, also distinguishes Glaukerpeton from Eryops. Dentition The marginal dentitions of the premaxilla and maxilla in Glaukerpeton consist of 10 or 11 and 37 or 38 teeth or their empty alveoli, respectively (Figs. 4, 9, 11, 12A). Maximum premaxillary and maxillary tooth counts in Onchiodon and Eryops are 13 and 20, and 13 and 38, respectively. As in Eryops and Onchiodon, the teeth have the form of sharply pointed, slightly recurved cones with prominent labyrinthine enfolding. The premaxillary teeth increase serially in size slightly posteriorly and are slightly longer than those in the maxilla, and those in the canine region near the anterior end of the maxilla are not significantly larger than their neighboring teeth. Only in Eryops do the premaxillary and maxillary dentitions include significantly larger caniniform teeth. In Glaukerpeton small denticles are randomly scattered over the entire parasphenoidal plate, but are more concentrated centrally between the basipterygoid processes, the entire palatal ramus of the pterygoid, and all but the posterior margin of the vomer (Figs. 9 11). This denticle distribution pattern is reduced in Onchiodon to a small patch between the basipterygoid processes, the transverse flange, and the anterior end of the palatal ramus of the pterygoid and continued onto the posterior half of the palatine, and the lateral margin of the vomer posterior to the vomerine tusk. The ectopterygoid in Glaukerpeton is unique in bearing three large tusks (Figs. 9, 11), rather than the two possessed by Onchiodon and Eryops, as well as in all other eryopids. However, paired tusks borne by the vomer and palatine are present in all eryopids. Together, the mandibles of Glaukerpeton indicate about 39 marginal tooth positions (Fig. 14), and it is estimated Fig. 16. Right clavicle of Glaukerpeton avinoffi, CMNH A, photograph; and B, C, illustrations in anterior and ventral views. that the complete series included slightly more than 40 labyrinthine teeth. As in Onchiodon, the marginal teeth of the mandible are small compared to those of the maxilla, and there is no development of a caniniform region. In Eryops, on the other hand, the eighth and ninth teeth are noticeably enlarged compared to the rest of the series. All the coronoids in Glaukerpeton bear small denticles, which are visible also on a coronoid fragment in the holotype (Fig. 1). The pattern of coronoid denticulation in some specimens of Onchiodon, and Eryops is known to extend from the symphysis to the level of the posterior end of the marginal teeth (Broom 1913; Langston 1953; Werneburg 2007). Postcranial Elements A few axial and appendicular elements were found closely associated with the skull of CMNH (Murphy 1971). The axial elements are well preserved (Fig. 15) and include the articulated fourth and fifth neural arches with a few closely associated ribs exposed on a block of matrix, whereas a sacral neural arch, pleurocentrum, and intercentrum are completely freed of matrix. All these elements are essentially identical to those in Eryops based on Moulton s (1974) detailed description of its vertebral column. Identification of the articulated fourth and fifth neural arches is based on the anterior neural arch spine being unusual short and pointed, as in Eryops, although its base is slightly wider

18 50 annals of carnegie museum vol. 81 Fig. 17. Right scapulocoracoid of Glaukerpeton avinoffi, CMNH A, B, photographs in medial, and lateral views; and C, D, illustrations of views in A and B. than that in Eryops (Moulton 1974). The identification of the isolated neural arch as belonging to a sacral vertebra is based on its large, circular facet for the rib. The intercentrum is slightly shorter anteroposteriorly relative to those in larger specimens of Eryops, but Moulton (1974: fig. 14) also figured a proportionally small intercentrum in Eryops. The pleurocentrum does not deviate from those of Onchiodon and Eryops except in its proportionally smaller size to that of the latter. Preserved appendicular elements in CMNH 11025

19 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 51 Fig. 18. Humeri of Glaukerpeton avinoffi, CMNH A, B, C, D, photographs of right humerus in extensor, flexor, anterior, and posterior views; E, F, photographs of distal end of left humerus in flexor and extensor views; and G, H, illustrations of C, anterior, and B, flexor views. include the complete right clavicle, nearly complete right scapulocoracoid, and complete right and distal half of the left humerus. The ventral or external surface of the ventral blade of the clavicle (Fig. 16) is sculptured almost entirely by fine, mediolaterally oriented grooves. On the distal portion of the blade some of the grooves extend a short distance laterally before ending in a pit. The morphology of the clavicle is identical to that described in detail in Eryops by Pawley and Warren (2006). The elongation of the ventral blade is of interest, because it gives

20 52 annals of carnegie museum vol. 81 Fig. 19. Cladogram illustrating hypothesized eryopid relationships of Glaukerpeton, Onchiodon, and Eryops. The numeral 4 is the Bremer (1994) decay value for the Eryops and Onchiodon node. an indication of relative size of the missing interclavicle. On this basis the blade in Glaukerpeton is proportionally a little more elongated medially than that in Eryops (see Cope 1888; Williston 1899; Case 1911; Pawley and Warren 2006), thereby suggesting that the width of the interclavicle may have been proportionally greater than that in Eryops. Werneburg (2007) discussed the ontogenetic trend toward reduction of the mediolaterally width of the interclavicle in Eryops, and it is possible that the same trend may have occurred also in Glaukerpeton. The right scapulocoracoid (Fig. 17) is well preserved and complete except for missing dorsal portion of the blade. The angle between the supraglenoid buttress and the anterior margin of the scapular blade is less than 90, whereas in most Eryops specimens the angle is greater than 90 (Broili 1899; Case 1911; Langston 1953; Pawley and Warren 2006; pers. obs. RW). However, a few exceptions exist in Eryops in which the angle between these two structures is less than 90 (Williston 1899; FMNH UR 756), but this may be due to postmortem distortion. In a small juvenile scapulocoracoid of Eryops, CM 8535, described by Romer (1952) from the Lower Permian Dunkard Group of West Virginia, the angle is more than 90. This indicates that this feature in Glaukerpeton is not ontogenetically variable and does not account for the difference in angulation in the two genera. The dorsal blade in Glaukerpeton appears to be narrower than that in Eryops, but this cannot be determined in the absence of an ontogenetic series of the former. However, it does appear that the glenoid region differs from that in Eryops in being directed more posteriorly and the ventral margin of the coracoid region being longer. Otherwise the scapulocoracoid in Glaukerpeton appears to be identical to those in Eryops (Pawley and Warren 2006). The clavicle and scapulocoracoid are too poorly preserved in Onchiodon to compare with those of Glaukerpeton. The humerus in Glaukerpeton (Fig. 18) is very similar to that in Eryops (Miner 1925; Romer 1971; Pawley and Warren 2006), but some minor differences are evident. The supinator process in Glaukerpeton is thicker than but not as elongated as that in Eryops. In addition, the supinator process in Glaukerpeton is directed at a right angle to the long axis of the humeral shaft rather than slightly proximally as in Eryops. As a result, the ectepicondylar notch in Glaukerpeton is much narrower and shorter than in Eryops (Miner 1925; Pawley and Warren 2006). The entepicondyle and radial condyle are far less expanded distally than in Eryops, resulting in the distal margin of the humerus having a convex margin in extensor (dorsal) and flexor (ventral) views. On the other hand, in Eryops the distal margin of the humerus in the same views is slightly concave due to the more distally expanded entepicondyle. As in Eryops (Pawley and Warren 2006), the deltoid and pectoral crests in Glaukerpeton occupy a distal position on the humeral shaft and are narrowly separated from the supinator process. However, in contrast to Glaukerpeton and as figured by Miner (1925) and Pawley and Warren (2006), the notch between the supinator process and the deltoid and pectoral crests is much larger in Eryops. The latissimus dorsi process in Glaukerpeton (see also Murphy 1971: fig. 21C) is much more weakly developed than in Eryops. Yet, the scapulohumeral ridge in Glaukerpeton is much more strongly developed than in Eryops (Pawley and Warren 2006). Near the base of the scapulohumeral ridge on the extensor surface of the humerus in Glaukerpeton is a deep depression, possibly a foramen, which is not known in Eryops. The humerus is unknown in Onchiodon. RELATIONSHIPS Proposed diagnoses of Eryopidae express quite differing opinions (Sawin 1941; Milner 1989, 1990; Boy 1990; Werneburg 1993, 1997, 2007; Werneburg and Steyer 1999; Schoch and Hampe 2004; Pawley and Warren 2006; Schoch and Witzmann 2009a, 2009b). As a result, the genera recognized as eryopids have been inconsistent (Milner 1989; Boy 1990; Werneburg 2007; Schoch and Witzmann 2009a, 2009b). Until the recently proposed diagnosis of the family by Werneburg (2007), there has been an inability to identify cranial synapomorphies that would unite all of the genera recognized as eryopids, which collectively include Eryops, Glaukerpeton, Onchiodon, Actinodon, Osteophorus, and Clamorosaurus. However, comparisons of Glaukerpeton with other eryopids, using primarily skull characters, must be limited by necessity to the well-known, unequivocal eryopids Onchiodon from the Early Permian and Eryops from the Late Pennsylvanian and Early Permian, as descriptions of the skulls of the other three eryopid genera are currently being revised. Osteophorus includes a single species, O. roemeri Meyer, 1856 (Meyer 1856, 1860), which is based on a single specimen from the Lower Permian Rotliegend of the North Sudetic Basin of Poland that consists of the greater portion of the skull roof. Unfortunately, the holotype was lost during World War II, and our understanding of its anatomy has been dependent solely on a drawing by Meyer (1860). However, this impasse may be overcome soon by skulls from the Rotliegend of the Intra Sudetic Basin of the Czech Republic that were tentatively assigned

21 2012 WeRnebuRg and berman late PennSylvanian eryopid Glaukerpeton 53 Fig. 20. Whole body restoration of swimming pair of Glaukerpeton avinoffi. to Onchiodon by Werneburg (1993), but who now believes may pertain to O. roemeri based on a restudy in progress by the senior author. If this reassessment is correct, then the holotypic skull of O. roemeri exhibits anomalous features not seen in the Czech Republic specimens, such as the possession of an interfrontal bone and the absence of a right prefrontal-postfrontal contact (but present on the left side of skull). Clamorosaurus is known only by Gubin s (1983) original description of C. nocturnus Gubin, 1983, and C. borealis Gubin, 1983, is based on several skulls from the Middle Permian of Russia. However, an ongoing revision of Clamorosaurus by the senior author has identified several important differences (manuscript in progress) from the skull reconstruction of Clamorosaurus by Gubin (1983). Werneburg (1988; see citations in Werneburg 2007) published descriptions of several eryopid skulls and postcranial remains from the Lower Permian Rotliegend of Germany that he attributed to different species of Onchiodon, but who now believes pertain to the Actinodon-complex of Werneburg (2007), which is based on the holotype of Actinodon frossardi and other material from the Lower Permian of France, Czech Republic, and Germany, and should be considered a member of Eryopidae (see discussion in Werneburg 2007). A restudy of this material is presently in progress by Werneburg and Steyer. A cladistic analysis using the Willi Hennig Society edition of TNT (Tree Analysis Using New Technology; Goloboff et al. 2008) was performed to resolve the relationships of Glaukerpeton, Onchiodon, and Eryops. The characters, 19 cranial and two postcranial, and their states are given in Appendix 4 and the character-taxon matrix is given in Appendix 5. The polarities of the character states were determined using the well-known Early Permian Sclerocephalus as an outgroup comparison taxon. This follows the cladistic analyses by Schoch and Witzmann (2009a, 2009b), which resolved Sclerocephalus it as basalmost member of the closest sister taxon to Eryopidae, Stereospondylomorpha. The analysis produced only one most

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