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30-Johnson (Xenacanth) Lucas, S.G., et al. eds., 2013, The Carboniferous-Permian Transition. New Mexico Museum of Natural History and Science, Bulletin 60. 161 XENACANTH SHARKS AND OTHER VERTEBRATES FROM THE GERALDINE BONEBED, LOWER PERMIAN OF TEXAS GARY D. JOHNSON Shuler Museum of Paleontology, Institute for the Study of Earth and Man, Southern Methodist University, PO Box 750274, Dallas, Texas 75275-0274; email: [email protected] Abstract—The Geraldine Bonebed occurs in the Nocona Formation (Wichita Group, Sakmarian age) in Texas. It has historically yielded a large number of mostly complete skeletons of four tetrapod taxa, for which it is famous, and also a diverse flora as well as other vertebrates. Bulk samples of matrix were recently screen-washed and sorted to produce a variety of vertebrate microfossils, including sharks, especially xenacanths. The non-xenacanth sharks are rare and include a petalodont tooth (Janassa?), Helodus sp. (4 teeth) and one partial hybodont tooth. These are all considerably more common higher in the Wichita Group. Only the petalodont and possibly the hybodont and Helodus represent a marine component in the fauna, but the marine faunas are more extensive higher in the Wichita. Also new to the fauna are acanthodians, actinopterygians, Cardiocephalus and Ophiacodon. Among the xenacanths are two typically small Xenacanthus sp. occipital spine fragments, two Orthacanthus sp. occipital spine frag- ments (one small, one very small) and hundreds of Orthacanthus teeth. Orthacanthus texensis teeth are much more common than O. platypternus teeth. Teeth of O. texensis and O. platypternus are comparable in size distributions, as determined by statistical analyses of the tooth-base measurements, to those higher in the Wichita Group. With one possible exception (the exact locality cannot be confirmed), O. texensis and O. platypternus are not known to occur below the Nocona Formation in Texas, nor are they anywhere older than Sakmarian age. INTRODUCTION Group) in central Archer County (Fig. 2) and recognized four types of The purpose of this study is to add additional taxa to the previ- associated deposits. One of these, a catastrophic event bonebed, is rep- ously known fauna of the Geraldine Bonebed with emphasis on the resented by the Geraldine Bonebed, but Sander’s (1989) primary intent xenacanth sharks, obtained by bulk processing of matrix to yield a verte- was to describe four occurrences of floodplain pond bonebeds. All four brate microfossil component (Johnson et al., 1994). This bonebed, dis- of these probably contain a more diversified vertebrate fauna than does covered by A. S. Romer in 1932 in central Archer County, probably the Geraldine Bonebed (three of the faunal lists were updated by Johnson, represents the most prolific source of articulated tetrapod skeletons in 2007, 2012). The other two types of deposits recognized by Sander the Lower Permian of North America (Sander, 1987). These include 11 (1989) are isolated skeletons and lag bonebeds. mostly articulated skeletons of Archeria crassidisca, an embolomerous In his description of the pond bonebeds, Sander (1989) did not amphibian; 15 or more associated or articulated skeletons of the laby- recognize any evidence of marine incursions. Hentz (1988, figs. 11-12) rinthodont amphibian Eryops megacephalus; 14 or more partial to com- presented a broad overview of the paleogeography of north-central Texas plete skeletons of the herbivorous synapsid Edaphosaurus boanerges; during the time of deposition of the Archer City Formation (Asselian and three associated skeletons of a carnivorous synapsid, Dimetrodon age). Based on this, Sander’s (1989) pond bonebeds occurred in the natalis. Sander (1987) provided a history of collecting these specimens upper part of a lower coastal plain. This could reasonably explain the together with pertinent associated details. He also provided a detailed presence of marine taxa in these bonebeds under varying circumstances, study of the sedimentology, flora (some two dozen taxa) and taphonomy although their occurrence in the Geraldine Bonebed is more problematic. of the bonebed. It is of Sakmarian (Wolfcampian) age and occurs in the GERALDINE BONEBED VERTEBRATE FAUNA Nocona Formation, Wichita Group (Figs. 1-2). Sander (1987) noted the low diversity of the vertebrate fauna with only three amphibian taxa Taxa in addition to those listed by Sander (1987, table II), includ- (including Diadectes sp.) and three amniote taxa (including Bolosaurus ing indeterminate partial bones and teeth plus tooth and bone fragments striatus) from the bonebed proper, plus two more amphibians that were obtained by bulk processing of matrix from the bonebed, are (Trimerorachis insignis and Zatrachys sp.) and one additional synapsid cataloged as SMU 76693-76753 (Shuler Museum of Paleontology, South- (Ophiacodon uniformis) from the same vicinity. Among the fishes, only ern Methodist University Locality 161). Additional surface-collected one shark (Orthacanthus texensis) and one crossopterygian fossils (SMU 69461-69472, 69499) did not add any taxa to those listed (Ectosteorachis nitidus) were recorded from the bonebed, plus one lung- by Sander (1987). The screen-washed bulk samples (two sites several fish (Sagenodus sp.) from nearby. meters apart within the bonebed) produced the following taxa (catalog Sander (1987) concluded that the Geraldine Bonebed and related numbers in parentheses; the xenacanths are treated separately below; sediments and flora constituted a floodbasin of a small meandering river * taxa not listed in the bonebed proper by Sander, 1987): system. The vertebrate-bearing facies contain only a minor fine-grained sandstone with ripple bedding in what otherwise is mudstone (Sander, Class Chondrichthyes 1987, fig. 3), which he interpreted to represent a freshwater pond in an Subclass Elasmobranchii overall swamp environment. Although he presumed the presence of ox- *hybodontid indet. (partial tooth, 76713) bow lakes in the region, he did not specify such an occurrence for the Orthacanthus texensis bonebed, presumably because of the geometry of the facies distribution. *O. platypternus Sander (1989) provided an analysis of the sedimentology of a portion of *Xenacanthus sp. the Nocona Formation and subjacent Archer City Formation (Bowie Subclass Holocephali *Helodus sp. (4 teeth, 76714) 162 FIGURE 1. Stratigraphic section of western North-Central Texas; from Johnson (2011), based on Hentz and Brown (1987). Abbreviations: Pcj, Coleman Junction Formation; Psb, Santa Ana Branch Shale; Pse, Sedwick Formation; Pmo, Moran Formation; Ppb, Pueblo Formation; lPP, Pennsylvanian- Permian; lPPh, Harpersville Formation. 163 Subclass Incertae Sedis In addition, there are a variety of fish teeth including *Janassa? (single incomplete petalodont tooth, actinopterygians (SMU 76621, 76723, 76724), amphibian teeth (76732) 76717) and a reptile caudal? vertebra (76737) and claw (76738). Three small Class Incertae Sedis Acanthodii coprolites (76739) are present; the smallest (6 mm) has a spiral structure *Acanthodes sp. (partial fin spines and scales, not and the other two contain palaeoniscoid scales. A variety of partial bones, common, 76715, 76716) isolated teeth and fragments are present (76740-76745), some of which Class Osteichthyes are probably identifiable. Subclass Actinopterygii *palaeoniscids indet. (scales and teeth common, XENACANTH SHARKS IN THE GERALDINE BONEBED 76718, 76719) Three species of xenacanths (Xenacanthiformes Berg, 1937, 1940; *Platysomus? (single “button tooth,” 76720; see Xenacanthodii Olson, 1946?; Xenacanthida Glikman, 1964) occur in the Johnson and Zidek, 1981) bonebed. Orthacanthus texensis is represented by teeth (SMU 76693- Subclass Sarcopterygii 76702) as is O. platypternus (SMU 76703-76707). Two Orthacanthus crossopterygian indet. (skull fragments, scales sp. small to very small occipital spine fragments (SMU 76709) were common, 76722) recovered. Also, two small spine fragments of Xenacanthus sp. (SMU Sagenodus sp. (5 partial tooth plates, 76725) 76710) occur in the fauna, but Xenacanthus teeth were not recovered, Class Amphibia similar to the faunas in the Archer City Bonebed 3 and Conner Ranch Order Temnospondyli Bonebed (Fig. 2; Johnson, 2012). Xenacanths are also represented by Trimerorhachis sp. (teeth and partial jaws denticles and prismatic cartilage (SMU 76711, 76712). uncommon, 76729) Orthacanthus texensis teeth are very common (total of 1808) in Eryops sp. (skull and jaw fragments common, 76728) the Geraldine fauna. Besides normal teeth, 13 teeth are germinal (under- Order Anthracosauria developed; Johnson, 2005) and three are deformed (Johnson, 1987) (SMU Archeria sp. (6 vertebrae, 76726) 76697, 76698). Of the remaining teeth, 141 were measured (Fig. 3A). A Order Microsauria statistical analysis is summarized in Table 1. The anteromedial-postero- *Cardiocephalus sp. (3 teeth, 76731) lateral (length of tooth base) dimension is taken as the independent Class Reptilia variable because it is usually easier to measure in Orthacanthus teeth. Order Parareptilia The measured population may be skewed toward the lower range (Fig. Bolosaurus sp. (14 teeth, partial jaw, 76730) 3A) because nearly all of the larger teeth were probably removed by Class Synapsida earlier surface collecting. Forty-nine teeth were surface-collected (SMU *Ophiacodon sp. (teeth common, 76735) 69461), but they are incomplete with some badly worn or weathered. Dimetrodon sp. (neural spine fragments and teeth Sander (1987, p. 228) noted that the teeth are smaller
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