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Zoological Journal of the Linnean Society, 2010, 160, 118138.
With 32 gures
Cranial osteology and preliminary phylogenetic assessment of Plectrurus aureus Beddome, 1880 (Squamata: Serpentes: Uropeltidae)
REBECCA S. COMEAUX, JENNIFER C. OLORI* and CHRISTOPHER J. BELL
Department of Geological Sciences, Jackson School of Geosciences, 1 University Station C1100, The University of Texas at Austin, Austin, TX 78712, USA
Received 11 July 2008; accepted for publication 31 March 2009
Uropeltid snakes are among the most poorly understood clades within Alethinophidia. Their small size, limited geographic distribution, high incidence of endemism, and fossorial behaviour all contribute to the general paucity of systematic collections of these snakes, especially of adequate skeletal preparations, in most museum collections. Their hypothesized position within the higher-order phylogeny of alethinophidian snakes calls attention to the need for additional morphological work on the group. Hypotheses of uropeltid phylogenetic relationships based on morphological analyses are few, and continue to be hampered by limited taxon-sampling and character matrices that rely predominantly on features that are visible on articulated skulls. We utilized high-resolution X-ray computed tomography (HRCT) to investigate the cranial osteology of Plectrurus aureus Beddome, 1880, a species for which no osteological data were previously available. We provide a detailed description of the skull and mandible, and comment on morphological characters and potential phylogenetic relationships. Clarity in character descriptions is of paramount importance, and additional morphological characters are desirable. HRCT provides a nondestructive way to identify new systematically informative morphological characters from digitally disarticulated specimens. The small size of many uropeltid species, including P. aureus, will help to frame a greater appreciation of the limitations of traditional HRCT protocols for revealing detailed anatomical features of small vertebrates. 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 118138. doi: 10.1111/j.1096-3642.2009.00595.x
ADDITIONAL KEYWORDS: computed tomography (CT) plectrurus skull anatomy snake phylogeny.
INTRODUCTION
Uropeltids are a diverse clade of fossorial snakes that inhabit tropical montane forests, agricultural elds, and occasionally wet lowlands in southern India and Sri Lanka (Rajendran, 1970, 1977, 1985; Gans, 1976). Little is known of their evolutionary history and no fossil specimens are yet recognized. They range in body size from 20 to 80 cm in total length, and are up to 2 cm in diameter (Gans, 1973). Their heads are small and pointed, but their tails are often thick and reinforced with a robust skeletal structure, including
*Corresponding author. E-mail: jen_olori@mail.utexas.edu
a uniquely formed bony caudal plate, lending an overall shape that is easily mistaken for the head, and from which the name uropeltid, meaning roughtailed is derived (Gans, 1976). Higher-order snake phylogeny remains an active and controversial area of research (Caldwell, 2007). Recent phylogenetic hypotheses consistently place uropeltids near the base of the evolutionary tree of alethinophidian snakes, but their exact relationships remain obscure. Morphological analyses consistently yield hypotheses in which uropeltids are closely related to Anomochilus Berg, 1901, Cylindrophis Wagler, 1828, and Anilius Oken, 1816, either with a stepwise succession of basal alethinophidians
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2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 118138
CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS (Anomochilus, uropeltids, Cylindrophis, and Anilius; e.g. Cundall, Wallach & Rossman, 1993; Lee & Scanlon, 2002; Lee et al., 2007) or with uropeltids as a sister group to Anomochilus, with that clade having variable hypothesized relationships with Cylindrophis and Anilius (e.g. Scanlon & Lee, 2000; Tchernov et al., 2000; Lee & Scanlon, 2002; Lee et al., 2007). An exception is a hypothesis based in part on morphology and presented by White, Kelly-Smith & Crother (2005), in which Uropeltis Cuvier, 1829 was sometimes recovered as sister to Anilius, but Anomochilus and Cylindrophis were not included in the analysis. Molecular estimates of the position of uropeltids vary considerably, depending on the data set analysed and the taxon sampling. Uropeltids (variably represented by Rhinophis Hemprich, 1820, and/or Uropeltis) were placed as sister group to the African Calabaria reinhardtii (Schlegel, 1848) by Heise et al. (1995), to (Tropidophis + Casarea) by Slowinski & Lawson (2002; based on c-mos), to Caenophidia by Vidal & Hedges (2004), to the anomalepidid Liotyphlops Peters, 1881 by White et al. (2005; but Cylindrophis was not included in the analysis), to Cylindrophis (Slowinski & Lawson, 2002; Vidal & Hedges, 2002; Wilcox et al., 2002; Lee et al., 2007), and to (Cylindrophis + Anomochilus) (Gower et al., 2005). Eight genera and 47 species of uropeltids are currently recognized (McDiarmid, Campbell & Tour, 1999), but their interrelationships are unresolved (Dessauer, Gartside & Gans, 1976; Dessauer, Cadle & Gans, 1987; Cadle et al., 1990; Rieppel & Zaher, 2002), and alternative hypotheses of relationships cannot meaningfully be compared. The two most recent hypotheses of relationships among uropeltids were presented by Cadle et al. (1990; based primarily on protein data) and Rieppel & Zaher (2002; based on skull morphology). In the aggregate, only 19 species were included in the phylogenetic hypotheses presented by Cadle et al. (1990) and Rieppel & Zaher (2002), and only three species were common to both analyses. The small size of uropeltids, their limited geographic distribution, high incidence of endemism, fossorial behaviour, and local superstitions regarding their handling all contribute to the general paucity of systematic collections of these snakes (Rajendran, 1990), especially of adequate skeletal preparations, in most museum collections. The skull morphology of uropeltid snakes is not well studied. The most comprehensive taxonomic surveys are those by Cundall & Irish (2008) and Rieppel & Zaher (2002). Early discussions and illustrations of the skull and mandibles of uropeltids were provided by Dumril (1853), Peters (1861), Jan & Sordelli (1865), and Boulenger (1893). The rst detailed description was that by Baumeister (1908) for Rhinophis philippinus (Cuvier, 1829) and Rhino-
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phis homolepis (Hemprich, 1820) (see McDiarmid et al., 1999, for a discussion of taxonomic synonymy). Subsequent authors in the early half of the 20th century included uropeltids as part of larger surveys of squamate or snake anatomy and evolution (Radovanovic, 1937; Mahendra, 1938; Bellairs, 1949; Bellairs & Underwood, 1951), and the same was true in the reviews by Underwood (1967) and Bellairs & Kamal (1981). A few other authors dedicated their attention to the specic anatomical features of uropeltids, notably the mandible (Rieppel & Zaher, 2000), cranialvertebral joint (Hoffstetter, 1939; Williams, 1959), or the conguration of the skull (Gans, 1973), and Smith (1943) briey reviewed cranial features of the group and provided illustrations of the skull of Uropeltis smithi (Gans, 1966) (reported as Uropeltis grandis by Smith; see McDiarmid et al., 1999 for a discussion of taxonomic synonymy). A resurgence of interest in anatomical features of basal snakes (including uropeltids) and their importance for elucidating phylogeny is reected in a series of papers from the last quarter of the 20th century, all of which include descriptions of uropeltid anatomy (Rieppel, 1977, 1978, 1979, 1980a, b, 1983; Groombridge, 1979a, b, c; Cundall & Rossman, 1993; Cundall et al., 1993; Zaher & Rieppel, 1999). These papers provide the foundation upon which our modern understanding of uropeltid skulls is being constructed (Cundall & Irish, 2008). The analysis by Rieppel & Zaher (2002) remains the only in-group phylogeny of uropeltids based entirely on morphology. As a result of the framework provided by that landmark paper we can place basic anatomy in a phylogenetic context, and begin to test uropeltid morphological characters. In this paper we fully describe the osteology of the skull of Plectrurus aureus Beddome, 1880, and use this information to evaluate existing characters and comment on potential phylogenetic relationships.
MATERIAL AND METHODS

A single specimen of P. aureus [California Academy of Sciences (CAS) 17177] was scanned at The University of Texas High Resolution X-ray Computed Tomography Facility (UTCT Facility). The specimen was collected by R.H. Beddome from Wynad, Kerala State, India, and is preserved in alcohol (Fig. 1). The date of collection is unknown. The total skull length is 9.4 mm from the tip of the rostrum to the distal edge of the occipital condyle. A wet specimen was utilized for scanning because in dry skeletal preparations elements may pull together creating false contacts. The use of an alcohol-preserved specimen produces CT scans that more accurately represent the gaps
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R. S. COMEAUX ET AL. scoring characters that are not visible in articulation, especially in cases where specimens may be too small, fragile, or rare for traditional methods of study, such as histological sectioning. For example, Rieppel & Zaher (2002) proposed a number of potentially informative characters (e.g. 27, 28, 31) that can only be scored on partially or fully disarticulated material. This is exactly a case where CT data have huge advantages. In sum, CT data reveal details of anatomy that are not generally accessible to many researchers.
Figure 1. Plectrurus aureus (CAS 17177), whole animal. Scale bar: 1 cm.
RESULTS
GENERAL
DESCRIPTION OF THE SKULL AND MANDIBLE
between bones, and render the intervening soft tissue clearly visible in unmodied CT slices. High-resolution CT scans were taken in the coronal (axial) plane, and resulted in 527, 16-bit TIFF images with an image size of 1024 1024 pixels. The original slices had an interslice spacing (slice thickness) of 0.01811 mm, with a eld of reconstruction of 7 mm. The completed scan of P. aureus yielded a data set of reasonably high quality, but the somewhat grainy nature of the renderings of the tiny, individual elements indicate that this uropeltid was near the size limit for the standard scanning and image-processing protocols used by UTCT at the time the specimen was scanned. The 3D graphics volume software VGStudioMax 1.21 was used to obtain slices in the other two orthogonal planes, producing a total of 744 sagittal slices and 503 frontal slices. 3D models of the skull were rendered using the same software package. Soft tissue was digitally removed by optimizing the density histogram for the greyscales representing bone. Where extremely thin bone exists in the skull (e.g. the notch in the nasal bone) this optimization can be challenging, because thin bone and some soft tissues may be rendered with similar greyscales. Thus, when the soft tissue is removed or rendered completely transparent, regions of extremely thin bone may also disappear. Individual cranial elements were digitally disarticulated using the manual segmentation tool within VGStudioMax. The resulting amplied images can be rotated and digitally manipulated to provide novel anatomical views, allowing a detailed analysis of each element. The entire, fully segmented skull is depicted in Figures 2 and 3 in dorsal, ventral, rightlateral, and anterior views. A major advantage of this technology is that it is non-invasive, permitting detailed data on skeletal anatomy to be gathered in a nondestructive manner. Digital disarticulation of the skull is particularly useful for identifying or
Premaxilla The edentulous premaxilla (Fig. 4) has a distinctly triangular form. The nasal process of the premaxilla is posterodorsally directed, and is exposed in dorsal view as a narrow wedge separating the nasals anteriorly. The vomerine process is a midline structure projecting posteriorly between the vomers. The medial edge of each vomer contacts the corresponding lateral edge of the vomerine process of the premaxilla. Ventrally, the premaxilla slots into a shallow notch on the ventral surface of the vomers; the vomer extends dorsal to the premaxilla in a horizontal, overlapping contact. The nasal process also extends above the medial portion of each septomaxilla, although actual bone-to-bone contact does not occur. A single premaxillary foramen is present, appearing as a slight circular indentation centered on the ventral surface of the bone. Distinct transverse processes (lateral processes of Rieppel, 1977) form a at, weakly buttressing contact with the anterior tip of each maxilla (= shizarthrotic contact of Cundall et al., 1993; Rieppel & Zaher, 2002; Fig. 5). There are grooves on the dorsal and ventral sides of the transverse process, the ventral side of the vomerine process, and on the rostrum. The anteromedial surface of the premaxilla is emarginated to form a distinctly bipartite rostrum (Rieppel & Zaher, 2002) that projects anteriorly a short distance beyond the transverse processes. Septomaxilla The posterolateral edge of the septomaxilla (Fig. 6) is directed dorsomedially and contacts the prefrontal. The nasal process of the premaxilla is positioned between the medial ange of each septomaxilla, and the vomerine process is ventral to, but not in contact with, the anteroventral edge. The lateral edge of the septomaxilla is medially inected, and is separated from lateral contact with the maxilla by a gap lled
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Figure 2. Articulated skull, anterior to the right. A, dorsal view. B, ventral view, lower jaws digitally removed. Abbreviations: cb, compound bone; co, coronoid; dt, dentary; ept, ectopterygoid; f, frontal; mx, maxilla; n, nasal; ot-occ, otico-occipital complex; pa, parietal; pf, prefrontal; pl, palatine; pm, premaxilla; pt, pterygoid; q, quadrate; sm, septomaxilla; vo, vomer. Scale bar: 1 mm.
with soft tissue. However, the anteroventral edge of the septomaxilla overlies and directly contacts the anteromedial process of the maxilla (Fig. 7). The dorsal surface of the septomaxilla is concave ventrolaterally, with a deep lateral edge curving upwards and medially, and a medial ange located on the anterodorsal medial edge. Dorsomedially, the septomaxilla is overlain by the ventromedial edge of the nasal. In the articulated skull this contact obscures a relatively large foramen for the vomeronasal nerve that is located on the posterodorsal medial tip of the septomaxilla (Cundall & Irish, 2008). The ventral surface of the septomaxilla forms a deep concave pocket for the vomeronasal organ. A strongly curved ange of bone from the lateral edge of the septomaxilla (lateral wall of Rieppel, 1977) forms the oor of the lateral side of the vomeronasal capsule, and contacts the dorsal side of the vomer. Maxilla The transverse process of the premaxilla abuts the anterior end of the maxilla (Fig. 8) in a weakly buttressing (but not fully buttressed, or shizarthrotic)
contact (Rieppel & Zaher, 2002), where the edges of the two bones are in articulation (Baumeister, 1908), just anterior to the anteromedial process. The dorsal surface of the anteromedial process of the maxilla underlaps the ventral side of the medially curved lateral edge of the septomaxilla, and closely approaches, but does not meet, the anterolateral vomerine process of the premaxilla. The palatine (or medial) process of the maxilla is posterior to the anteromedial process, and is in contact with the ventral anterolateral curvature of the choanal process of the palatine. The maxilla tapers gradually as it projects posteriorly to meet the lateral side of the maxillary process of the ectopterygoid in a mediolaterally overlapping contact. The ascending process (Rieppel, 1977; Rieppel & Zaher, 2002) (= prefrontal process of Baumeister, 1908; dorsal process of Cundall & Irish, 2008) of the maxilla contacts the prefrontal in a strong, interlocking articulation. The open superior alveolar nerve canal is exposed dorsally, just medial to the ascending process (Rieppel & Zaher, 2002: 124). Seven tooth positions are located ventrally: ve functional teeth are in place
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Figure 3. Articulated skull. A, right-lateral view, anterior to the right. B, anterior view. Abbreviations: ang, angular; cb, compound bone; co, coronoid; dt, dentary; ept, ectopterygoid; f, frontal; mx, maxilla; n, nasal; ot-occ, otico-occipital complex; pa, parietal; pl, palatine; pf, prefrontal; pm, premaxilla; q, quadrate; sm, septomaxilla; V2, opening for the maxillary branch of the trigeminal nerve; V3, opening for the mandibular branch of the trigeminal nerve. Scale bar: 1 mm.
Figure 4. Isolated premaxilla, anterior to the right except in (C). A, dorsal view. B, ventral view. C, anterior view. D, right-lateral view. Abbreviations: np, nasal process; pmf, premaxillary foramen; ro, rostrum; tp, transverse process; vop, vomerine process. Scale bar: 1 mm.
Figure 5. Dorsal view of the articulated maxillae and premaxilla with other skull elements digitally removed. Abbreviations: mx, maxilla; pmx, premaxilla. Scale bar: 1 mm.
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Figure 6. Isolated septomaxilla. A, posterior view, medial to the left. B, dorsal view, anterior to the right. C, ventral view, anterior to the right. D, right-lateral view, anterior to the right. E, right-medial view, anterior to the left. Abbreviations: mf, medial ange; vnc, vomeronasal capsule; vnf, vomeronasal nerve foramen. Scale bar: 1 mm.
Figure 7. Articulated septomaxillae and right maxilla with other skull elements digitally removed, anterior view. Abbreviations: mx, maxilla; sm, septomaxilla. Scale bar: 1 mm.
on the right side and appear to be fully ankylosed. The teeth are relatively large and distinctly recurved. A well-developed replacement tooth is visible in the second tooth position, but is not yet ankylosed (Fig. 8B, C); additional replacement teeth are visible for some of the distal tooth positions (Fig. 8C). Two foramina are present: the rst is ovoid in shape and located at the anterior end of the maxilla; the second, smaller opening is below the ascending process. Nasal The paired nasals (Fig. 9) are separated anteriorly by the premaxilla for approximately one-third of their length. A distinct medial process (not visible in the articulated skull) is located on the posteromedial portion of each nasal: it forms a weak vertical contact with the other nasal along the midline, but declines in
Figure 8. Isolated maxilla. A, dorsal view, anterior to the right. B, right-lateral view, anterior to the right. C, rightmedial view, anterior to the left. Abbreviations: amp, anteromedial process; asp, ascending process; ep, ectopterygoid process; plp, palatine process; sanc, superior alveolar nerve canal. Scale bar: 1 mm.
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R. S. COMEAUX ET AL. anterior margin of the orbit. The prefrontal wraps around a protrusion, the preorbital ridge (Rieppel, 1978), located on the anterolateral edge of the frontal. The dorsomedial edge of the prefrontal is positioned next to the posterolateral edge of the nasal; the lateral curvature of the septomaxilla contacts the anteromedial edge of the prefrontal. A broad, bipartite maxillary process extends from a lateral foot process (Rieppel, 1977) anteromedially to a medial foot process (Fig. 10A); the notch between these two accommodates the ascending process of the maxilla. The lateral foot process is nger-like, and extends posterolaterally to form a slight overlapping contact with the posterior edge of the ascending process of the maxilla (Fig. 3A). Medial to that articulation, the prefrontal is notched to form the dorsal portion of a lacrimal duct. In the articulated skull, the lateral wall of that duct is formed by the prefrontal, and is oored by the maxilla, whereas the palatine contributes to the medial margin of the duct (Fig. 11). However, a narrow line of soft tissue prevents the prefrontal from directly contacting the palatine. Vomer The dorsal surface of the anterolateral process of the vomer (Fig. 12) is situated beneath the ventral surface of the septomaxilla, and projects laterally towards the anteromedial process of the maxilla (although no actual contact occurs; Fig. 13). The dorsally concave pocket of the ventral surface of the septomaxilla overlies the dorsal surface of the vomer, and completes the vomeronasal capsule. Anteromedially, a premaxillary process meets the vomerine process of the premaxilla in a complex articulation. The vomerine process of the premaxilla is situated within a recess between the two vomers, but dorsally each vomer overlaps the caudal tip of the vomerine process of the premaxilla.
prominence anteriorly, and is completely diminished at the level of the posteriormost extent of the nasal process of the premaxilla. A small notch or emargination is present on the anterolateral edge of the nasal, a short distance behind the anteriormost tip (Fig. 9A, B). Although this region is notched in some uropeltids (Rieppel & Zaher, 2002), in the CT data set of P. aureus this probably represents an area of thin bone and not an actual notch. The nature of the notch mentioned by Rieppel & Zaher (2002) for their character 2 is unclear, but presumably refers to the nasal margin of the external naris(?). We interpret the notch in P. aureus as an artifact of digital image processing, resulting from the loss of bone when soft tissues were digitally removed from the original data set to reveal the skull. The nasal overlies the posterodorsal tip of the septomaxilla, with the lateral edge of the nasal curving ventrally inward towards the dorsally curving lateral edge of the septomaxilla. The lateral edge of the posteroventral surface of the nasal contacts the ventromedial curvature of the prefrontal. The posterior surface of the nasal directly meets the anterior face of the medial ange and the anterodorsal surface of the frontal in an overlapping contact. Prefrontal The frontal process (supraorbital process of Cundall & Irish, 2008) of the prefrontal (Fig. 10) is located posterodorsally, and articulates with the frontal via a shallow notch on the anterolateral surface of the frontal. The supraorbital process of the parietal does not meet the frontal process of the prefrontal, a feature that was reported as polymorphic for Plectrurus perroteti Dumril & Bibron, 1854 (Rieppel & Zaher, 2002: character 7). The posteroventral surface of the prefrontal, ventral to the frontal process, abuts the anteroventral surface of the frontal and forms the
Figure 9. Isolated nasal, anterior to the right. A, ventral view. B, dorsal view. C, right-lateral view. Abbreviation: mp, medial process. Scale bar: 1 mm.
Figure 10. Isolated prefrontal. A, right-lateral view, anterior to the right. B, posterior view, lateral to the right. Abbreviations: fp, frontal process; lc, notch for lacrimal canal; lfp, lateral foot process; mfp, medial foot process. Scale bar: 1 mm.
CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS The posterolateral process appears midway along the length of the vomer, curving laterally and dorsally towards the maxilla, although the two do not meet (Fig. 13). It is widest anteriorly, and is connected to the medial ridge via a low crest that marks the posterior surface of the vomeronasal capsule. A small, rounded foramen transmits the vomeronasal nerve, and penetrates the crest near where it meets the medial ridge (Figs 12D, 14). The posterolateral process tapers posteriorly to a pointed palatine process that is directed posteriorly and positioned inside the choanal process of the palatine; the palatine process of the vomer lls a narrow ventral gap running along the lateral edge of the choanal process of the palatine (Fig. 15). The medial edge of the vomer forms a dorsally projecting ridge that serves as a vertical contact
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between the two vomers anteriorly. The interchoanal process of the sphenoid extends anteriorly along the midline to the level of the posteromedial ends of each vomer, but does not contact either vomer (Fig. 16). Palatine The palatines (Fig. 17) are edentulous, as in all uropeltids studied so far except for Melanophidium punctatum Beddome, 1871 (Rieppel & Zaher, 2002:
Figure 11. Posterior view of the snout region with the otico-occipital, frontals, parietal, pterygoids, ectopterygoids, quadrates, and lower jaws digitally removed. Abbreviations: lc, lacrimal canal; mx, maxilla; n, nasal; pf, prefrontal; pl, palatine. Scale bar: 1 mm.
Figure 13. Ventral view of the articulated maxillae and vomers with the other skull elements digitally removed; anterior to the right. Abbreviations: mx, maxilla; vo, vomer. Scale bar: 1 mm.
Figure 12. Isolated vomer, anterior to the right except in (D). A, ventral view. B, dorsal view. C, right-lateral view. D, anterior view, medial to the right. Abbreviations: alp, anterolateral process; mr, medial ridge; plp, palatine process; pmp, premaxillary process; polp, posterolateral process; vnc, vomeronasal capsule; vnf, vomeronasal nerve foramen. Scale bar: 1 mm.
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Figure 14. Posterior view of the septomaxillae (top) and vomers (bottom) in life position, with other skull elements digitally removed. Abbreviations: crest, vertical crest along dorsal surface of posterolateral process of the vomer; mr, medial ridge of vomer; vnf, vomeronasal nerve foramen. Scale bar: 1 mm.
Figure 15. Anterolateral view of the complex articulation between the palatines and vomers; anterior to the frontright. Other skull elements have been digitally removed; Abbreviations: pl, palatine; vo, vomer. Scale bar: 1 mm.
Figure 16. Dorsal view of the otico-occipital complex and palate with the frontals, nasals, parietal, pterygoids, ectopterygoids, quadrates, and lower jaws digitally removed; anterior to the right. Abbreviations: mx, maxilla; ot-occ, otico-occipital complex; pf, prefrontal; pl, palatine; pmx, premaxilla; sm, septomaxilla; vo, vomer. Scale bar: 1 mm.
Figure 17. Isolated palatine, anterior to the right, except in (D). A, ventral view. B, dorsal view. C, right-lateral view. D, anterior view, medial to the right. Abbreviations: chp, choanal process; ich, internal choana; lp, lateral process of Rieppel & Zaher (2002); mxf, facet for maxilla; ptp, pterygoid process; V2f, foramen for the maxillary branch of the trigeminal nerve; vop, vomerine process. Scale bar: 1 mm.
CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS character 3), and are separated from one another along their entire midline length. The choanal process is broad, arching in a medial curvature to form the internal choana. Medially the ventral edge of the process is elongated into distinct anterior and posterior extensions. The anterior end of the palatine is broadened, and underlaps the posterior portion of the vomer along its medial edge. On its lateral side the anterior surface of the palatine overlaps the maxilla. A distinct facet on the ventrolateral surface of the anterior palatine marks the articulation with the palatine process of the maxilla. A large foramen for the maxillary branch of the trigeminal nerve is present on the anterolateral surface of the palatine. Its ventral margin is formed by what Rieppel (1977) called the lateral process of the palatine. In an articulated skull, the complex articulation of the palatine and maxilla create the appearance of a distinct process in ventral view, but in the disarticulated skull of P. aureus, it is clear that the process is merely the ventral portion of the palatine ossication surrounding the foramen for the nerve (Rieppel, 1977; Figs 2B, 17). The anterior tip of the pterygoid inserts into a posterior groove on the pterygoid process of the palatine (Figs 2B, 17A). The palatine closely approaches, but does not contact, the frontal dorsally: the space separating the elements is small, presumably lled with soft connective tissue, and would probably be seen as a clear contact in a dried skull. Similarly, the dorsolateral edge of the palatine is positioned ventral to the prefrontal, but the two elements do not make direct contact. Pterygoid The anterior end of the pterygoid (Fig. 18) forms a palatine process that articulates with the grooved surface of the pterygoid process of the palatine. The quadrate ramus extends posteriorly to the level of the mandibular articulation, beneath the crista circumfenestralis of the otico-occipital complex. The ectopterygoid process is an anterolateral projection that extends beneath the ectopterygoid, overlapping it in a
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horizontal contact (anteriorly, the pterygoid underlies the ventral side of the ectopterygoid). No teeth or foramina are present. Ectopterygoid The anterior maxillary process of the ectopterygoid (Fig. 19) meets the maxilla in an overlapping contact along the posteromedial surface of the maxilla. The pterygoid process is located on the posterior end of ectopterygoid, and overlaps the pterygoid in a horizontal contact. No foramina are present. Frontal The frontal (Fig. 20) closely approaches, and in dried skull may contact, the palatine anteroventrally: in our digital renderings, there is a narrow gap between them that is lled with connective tissue, as in other alethinophidian snakes. In medial view the frontal forms an open, anteriorly tapering chamber that accommodates the olfactory bulbs of the brain. Anteriorly, an olfactory tract canal is formed lateral to the open medial contact between the dorsal and ventral medial edges of each frontal (Rieppel, 1977). A ridge located ventrolaterally and directed towards the palatine forms the palatal process of the frontal. It contacts the parasphenoid region of the sphenoid bone medially: a at surface formed between the sphenoid and the palatal process presumably overlies the trabecula, extending from the base of the ossied crista trabecularis, although the cartilage is not visible in the scans.
Figure 18. Isolated pterygoid in dorsal view, anterior to the right. Abbreviations: ep, ectopterygoid process; plp, palatine process; qr, quadrate ramus. Scale bar: 1 mm.
Figure 19. Isolated ectopterygoid, anterior to the right. A, ventral view. B, dorsal view. Abbreviations: mxp, maxillary process; ptp, pterygoid process. Scale bar: 1 mm.
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Figure 20. Isolated frontal. A, anterior view. B, rightlateral view, anterior to the right. C, right-medial view, anterior to the left. Abbreviations: opf, optic foramen; otc, olfactory tract canal; palp, palatal process; por, preorbital ridge; sog, groove for the supraorbital process of the parietal. Scale bar: 1 mm.
The open posterior edge of each frontal contacts the open anterior surface of the parietal. The supraorbital process of the parietal projects anteriorly into a groove on the dorsolateral surface of the frontal. A large optic foramen is enclosed entirely within the frontal, anterior to the frontalparietal contact. Anterolaterally, a slight dorsal groove marks the articulation with the frontal process of the prefrontal. Ventral to that groove, the preorbital ridge forms an additional contact surface with the prefrontal. The ridge extends anteriorly past the margin of the dorsal exposure of the frontal, a feature that is shared with other uropeltids and related taxa (Rieppel & Zaher, 2002: character 27). Although there is a narrow gap between the dorsal surfaces of the nasal and frontal (lled with soft tissue), the anteromedial and anterodorsal edges of the frontal directly contact the nasal. Postorbital and supraorbital The postorbital and supraorbital are absent as discrete ossications in P. aureus and other uropeltids. Parietal The parietal (Figs 21, 22) is an unpaired element with pronounced descending anges that form much of the body of the bone and contact the sphenoid region of the otico-occipital complex. The posterodorsal surface extends farther posteriorly from the level of the descending anges, and overlaps the otic region of the otico-occipital complex. The paired supraorbital processes (possibly homologous with the postfrontal; Rieppel, 1977; Cundall & Irish, 2008) are located on the lateral sides of the parietal and project anteriorly,
Figure 21. Isolated parietal, anterior to the right. A, dorsal view. B, ventral view. Abbreviations: aff, articulation facet for the frontal; sag, sagittal crest; sop, supraorbital process; V2f, fenestra for the V2 branch of the trigeminal nerve; vlf, ventrolateral ange. Scale bar: 1 mm.
articulating with the frontal via a shallow groove on the lateral surfaces of the frontal, dorsal to the optic foramen (Figs 3A, 22B). The supraorbital processes do not contact the prefrontals (Fig. 3A). The anterior surface of the main body of the parietal contacts the open posterior edge of each frontal, extending the open cavity and enclosing the central portion of the brain. A narrow shelf of bone between the supraorbital processes forms the articulation facet for the frontals. A faint sagittal crest is located on the posterodorsal surface of the parietal along the sagittal midline (Fig. 21A). The parietal does not contribute to the margin of the optic foramen in P. aureus. Posterolaterally, there is a shallow notch in the wall of the parietal that marks the passage of the V2 branch of the trigeminal nerve (Figs 21, 22); in the articulated skull, the parietal thus forms the anterior margin of the opening for the passage of the V2 branch (Fig. 3A). Otico-occipital complex The otico-occipital complex (Figs 2325) is a single element in P. aureus, presumably composed of parasphenoid, basisphenoid, basioccipital, laterosphenoid, prootic, opisthotic, exoccipital, and possibly supraoc-
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Figure 22. Isolated parietal. A, anterior view. B, rightlateral view, anterior to the right. Abbreviations: sag, sagittal crest; sop, supraorbital process; V2f, fenestra for the V2 branch of the trigeminal nerve; vlf, ventrolateral ange. Scale bar: 1 mm.
cipital ossications, all in a state of complete fusion with one another (no sutures can be seen). The occipital condyle, otic region, and sphenoid region are distinct regions of the element. In dorsal view, the parietal articulation facet is extensive, extending back to the position of the common crus between the anterior and posterior semicircular canals. A low, midline sagittal crest extends from the posterior end of the synotic tectum, anteriorly to the midline emargination of the roof. On either side of the sagittal crest, a distinct foramen opens into a canal that traverses the bone posteroventrally. These foramina were illustrated by several authors (e.g. Smith, 1943; Gans, 1973; Rieppel, 1977; Rieppel & Zaher, 2002; Cundall & Irish, 2008), but remain unnamed, and their function is unknown. We here name these as Rieppels canal, in honor of Olivier Rieppel, who has done so much to advance our understanding of uropeltid snakes. In some uropeltid taxa, these canals are incompletely formed (Rieppel & Zaher, 2002: g. 5C), and are instead developed as a posteriorly positioned notch (Rieppels notch); in some species this may be asymmetrical within an individual [e.g. Uropeltis ocellata (Beddome, 1863); Cundall & Irish, 2008]. The exoccipitals and basioccipital are fused to form the occipital condyle (Baumeister, 1908; Rieppel & Zaher, 2002). The occipital condyle forms a hemispherical knob that is positioned on an elongated neck of bone (Baumeister, 1908; Williams, 1959).
There is no indentation (fovea dentis of Williams, 1959) on the dorsal surface of the condyle, although a trough for the brainstem is visible in dorsal view, similar to the situation in Uropeltis, Rhinophis, and P. perroteti (Rieppel & Zaher, 2002: character 17). The sphenoid region of the otico-occipital complex, consisting of fused parasphenoid and basisphenoid elements, makes up approximately one-half of the complex in length, and is open dorsally. Its lateral edges taper anteriorly in a stepwise fashion from the anterior portion of the otic region. The rst (most posterior) stepwise reduction in lateral extent happens at the position of the secondary anterior opening of the vidian canal (secondary anterior foramen of Underwood, 1967). The next lateral reduction happens at the position of the ossied base of the crista trabecularis, which ends behind the (lateral) frontoparietal suture (Rieppel & Zaher, 2002: character 6). Continuing anteriorly from this point, the sphenoid region tapers smoothly to terminate in a ventrally positioned, narrow interchoanal (or intervomerine) process. That process extends to sit between the posterior ends of the vomers in dorsal view, and is positioned medially between the dorsal surfaces of the choanal processes of the palatines posteriorly. A low keel extends posteriorly from the interchoanal process (Fig. 23B). The junction between the otic and sphenoid regions is roughly marked by the anterior opening of the vidian canal, and the anterior opening of the sixth cranial nerve (CN VI). The anterior opening for CN VI probably also transmits the internal carotid artery (Rieppel, 1979). The passage for CN VI follows a posterolateral course, merging with the vidian canal and emptying posteriorly into the prootic canal. The passage of the second (maxillary) branch of the trigeminal nerve (V2) is visible in lateral view, where the parietal meets the otico-occipital complex (Fig. 3A). In the isolated otico-occipital element, this opening is marked by a shallow notch along the anterior surface of the otic region (Fig. 25). The foramen for the third (mandibular) branch (V3) is posterior to V2, and is separated from it by a fused, broad laterosphenoid ossication (Rieppel, 1976; Rieppel & Zaher, 2002). Posterior and slightly ventral to the V3 foramen, the prootic canal is clearly a separate opening. The facial foramen (transmitting CN VII) opens within the prootic canal and traverses its length. The posterior opening of the vidian canal opens into the prootic canal internally along the lateral edge of the anteroventral portion of the prootic canal. The conguration of the foramina follows that reported by Rieppel & Zaher (2002: character 12) for Uropeltis, P. perroteti, and Rhinophis drummondhayi Wall, 1921 (but not Rhinophis sanguineus Beddome, 1863).
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Figure 23. Isolated otico-occipital complex, anterior to the right. A, dorsal view. B, ventral view. Abbreviations: cer, cerebral carotid; ct, anterior end of the ossied crista trabecularis; ef, endolymphatic foramen; ik, interchoanal keel; ip, interchoanal process; jsr, juxtastapedial recess; oc, occipital condyle; pavc, primary anterior opening of the vidian canal; Rc, Rieppels canal; savc, secondary anterior opening of the vidian canal; VI, cranial nerve six. Scale bar: 1 mm.
A conspicuous juxtastapedial recess is visible immediately posterior to the prootic canal. In P. aureus the recess is mostly open, although the anterior half of the recess is somewhat restricted by a dorsal incursion of the crista circumfenestralis. Along the medial portion of the recess, the large fenestra ovalis opens into the otic chamber. The stapedial footplate lls the fenestra ovalis (Fig. 26). At the posteroventral edge of the stapedial footplate, the foramen pseudorotunda is visible in lateral view (Fig. 25A). Just anterior and ventral to the foramen pseudorotunda, the lateral aperture of the recessus scalae tympani opens beneath the stapedial footplate. It is obscured in lateral view by the development of the crista circumfenestralis. The recessus scalae tympani traverses the otic region, and a prominent medial aperture opens internally near the oor of the braincase (Fig. 25B). The vagus foramen (transmitting CN X and the jugular) is located posterior to the fenestra ovalis, in a shallow lateral pocket of bone formed by a lateral extension of the crista circumfenestralis; it can be
interpreted to be inside the juxtastapedial recess, because the recess is open posteriorly, with no distinct posterior margin (Rieppel & Zaher, 2002: character 14). The vagus foramen is bifurcated internally, a feature also exhibited by Uropeltis, Rhinophis, and P. perroteti (Rieppel & Zaher, 2002: character 15). A single, small hypoglossal foramen (transmitting CN XII) perforates the otico-occipital complex lateral to the base of the foramen magnum and ventral to Rieppels canal. The medial surface of the otic region is also pierced by foramina (Fig. 25B). Just ventral to the trigeminal nerve branches, the vidian canal is directed posteriorly; medial to the vidian canal is an opening to transmit CN VI and the internal carotid artery. Beneath the ventral margin of the otic capsule, the bone is excavated into the internal auditory meatus (transmits CN VIII). A smaller opening for CN VII passes through the otic chamber anterodorsal to the internal auditory meatus. The endolymphatic foramen pierces the medial wall of the otic chamber (Fig. 27).
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complex. The statolith on the right side of the skull has been digitally removed to better display the inside of the bony vestibule. Each statolith is ovoid in form and is composed of very dense material. In other snakes, the statolithic mass is not ossied, but consists of a densely packed clump of crystals (C. Bell, pers. observ.). Quadrate The quadrate (Fig. 28) is suspended from the ventrolateral surface of the otic capsule. The suprastapedial process (Lee, 2005) projects posteriorly, tapering into a rounded tip that extends over the stapedial shaft. The suprastapedial process is longer than the mandibular condyle, a feature that is characteristic of uropeltids and Anomochilus (Rieppel & Zaher, 2002: character 22). A shallow groove is located on the anteroventral surface of the condyle where it articulates with the compound bone. There is a low crest curving laterally along the dorsal surface of the quadrate.
MANDIBLE
The mandible is delicately built and in the adult is composed of ve separate ossied elements (Fig. 29). In the articulated cranium, there is a distinct gap between the anteromedial tips of the mandibles, indicating the presence of extensive soft tissues at the symphyseal region. The mandible reaches its greatest height at the level of the coronoid. A short retroarticular process is formed posteriorly.
Figure 24. Isolated otico-occipital complex. A, anterior view. B, posterior view. Abbreviations: cer, cerebral carotid; ct, anterior end of the ossied crista trabecularis; ef, endolymphatic foramen; fpsr, foramen pseudorotunda; ik, interchoanal keel; ip, interchoanal process; jsr, juxtastapedial recess; ls, laterosphenoid; oc, occipital condyle; pavc, primary anterior opening of the vidian canal; Rc, Rieppels canal; savc, secondary anterior opening of the vidian canal; st, stapes; tfc, trigeminofacialis chamber; VI, cranial nerve six; XII, hypoglossal foramen. Scale bar: 1 mm.
Stapes The stapes (Fig. 26) has a large stapedial footplate that lls the fenestra ovalis within the juxtastapedial recess of the otico-occipital complex. There is a short but well-ossied stapedial shaft. Statolithic mass A large statolithic mass (Fig. 27) is located within each of the otic chambers of the otico-occipital
Dentary A pronounced posterodorsal process of the dentary (Fig. 30) contacts the compound and coronoid bones. A posteroventral groove on the medial surface of the dentary marks the articulation with the splenial and angular. There is no posteroventral process of the dentary, which differs from the situation reported for P. perroteti by Rieppel & Zaher (2002: character 18). There are eight tooth positions located on the dorsal surface. A few of the teeth do not appear to be ankylosed to the dentary. These are probably replacement teeth, and are apparently less dense near their bases. Meckels canal (Lee, 2005) is closed anteriorly (except for a tiny ventromedial opening at the extreme anterior end of the dentary), but is open posteriorly as a prominent groove just dorsal to the splenial and angular. A single mental foramen is present on the lateral surface, at the level of the third tooth position. Splenial The splenial (Fig. 31) is a small, sharply triangular bone positioned on the posteromedial surface of the dentary. The broad, at posterior end of the splenial
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Figure 25. Isolated otico-occipital complex, anterior to the right; A, right-lateral view. B, left-medial view. Abbreviations: apm, medial aperture for the recessus scalae tympani; cer, cerebral carotid; ct, anterior end of the ossied crista trabecularis; ef, endolymphatic foramen; fpsr, foramen pseudorotundum; iam, internal auditory meatus; ik, interchoanal keel; ip, interchoanal process; jug, jugular foramen; lsf, laterosphenoid foramen; oc, occipital condyle; pc, prootic canal (containing the posterior opening of the vidian canal and cranial nerves VI and VII); savc, secondary anterior opening of the vidian canal; st, stapes; V2, notch for trigeminal nerve branch; V3, opening for trigeminal nerve branch; vc, vidian canal; VI, cranial nerve six; VII, cranial nerve seven; X, vagus nerve. Scale bar: 1 mm.
Figure 26. Close up of the right-lateral occipital region; anterior to the right. The stapes is outlined, with the shaft located in the bottom left of the outline. Scale bar: 1 mm.
meets the anterior end of the angular in a buttressing vertical contact. The lateral side of the splenial lies at against the medial side of the dentary. A small foramen is positioned dorsally, just posterior to the position of the last tooth on the dentary. There is no dorsal process (Cundall & Irish, 2008).
Figure 27. Axial section through the otic region of the otico-occipital complex in anterior view. Abbreviations: ef, endolymphatic foramen; fm, foramen magnum; hcc, horizontal semi-circular canal; jsr, juxtastapedial recess; otic, otic capsule; pcc, posterior semi-circular canal; stat, statolithic mass. Scale bar: 1 mm.
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Figure 28. Isolated quadrate in right-lateral view; anterior to the right. Abbreviations: cr, crest; mc, mandibular condyle; ssp, suprastapedial process. Scale bar: 1 mm.
Figure 30. Isolated right dentary. A, right-lateral view, anterior to the right. B, right-medial view, anterior to the left. Abbreviations: antf, anterior opening of Meckels canal; Mklc, Meckels canal; mtlf, mental foramen; pdp, posterodorsal process. Scale bar: 1 mm.
to the compound process of the dentary, although no direct contact occurs between the two elements. There are no foramina.
Figure 29. Right lower jaw. A, lateral view. B, medial view. Abbreviations: ang, angular; cb, compound bone; co, coronoid; d, dentary; spl, splenial. Scale bar: 1 mm.
Angular The angular (Fig. 31) is a small, triangular bone situated along the anteroventral side of the compound bone and the posteroventral edge of the dentary. It contacts the splenial anteriorly in a at, vertical, buttressing contact. A small, nger-like splenial process extends anteriorly between the splenial and the dentary on the lateral side. The angular sits at against the ventral side of the compound bone. There are no foramina. Coronoid The anteromedial process of the coronoid (Fig. 31) articulates along the medial side of the anterodorsal portion of the coronoid process of the compound bone. A slight groove on the lateral surface of the coronoid facilitates additional articulation with the compound bone. The anteromedial process is positioned medial
Compound bone A short retroarticular process forms the posterior portion of the compound bone (Fig. 32), and immediately anterior to the process is a deep, crescentric notch for the mandibular condyle of the quadrate. At the anterior end of the medial surface of the retroarticular process, a small foramen for the chorda tympani nerve enters the compound bone. A pronounced coronoid process rises on the lateral side of the compound. The coronoid bone articulates along the medial surface of this process, and extends farther dorsally than the compound bone, so that the coronoid is visible in lateral view. Anterior to the coronoid process, the compound bone slopes anteroventrally, and forms an elongated slanting contact with the posterodorsal process of the dentary. Anteroventrally, a horizontal contact is formed with the angular. The anterior end of the compound bone is strongly bifurcated, with lateral and medial processes positioned on either side of a central mandibular canal. At about the midpoint of the element on the medial side, an elongate mandibular fossa opens ventrally to the interior of the bone. A small foramen is present anteriorly on the lateral surface.
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Figure 31. A, articulated splenial and angular in left-lateral view, anterior to the left. B, articulated splenial and angular in right-medial view, anterior to the left. C, articulated splenial and angular in dorsomedial view, anterior to the right. D, isolated coronoid in right-lateral view, anterior to the right. E, isolated coronoid in right-medial view, anterior to the left. Abbreviations: ang, angular; amp, anteromedial process; spl, splenial; splf, splenial foramen; splp, splenial process of the angular. Scale bar: 1 mm.
Figure 32. Isolated right compound bone. A, right-lateral view, anterior to the right. B, right-medial view, anterior to the left. Abbreviations: cop, coronoid process; ctf, foramen for the chorda tympani; ldp, lateral dentary process; mdbc, mandibular canal; mdbf, mandibular fossa; mdp, medial dentary process; mj, mandibular joint; rap, retroarticular process. Scale bar: 1 mm.
DISCUSSION
More complete knowledge of cranial morphology in P. aureus allows for phylogenetically meaningful comparison with P. perroteti and other uropeltids included by Rieppel & Zaher (2002). Their work provides a framework of potentially useful morphologic characters that we used in our preliminarily assessment of P. aureus. In order to summarize important features and facilitate comparison, scores for P. aureus were added to Rieppel & Zahers (2002) matrix, and this information is presented in Table 1.
Signicantly, P. aureus differs from P. perroteti in regard to three of the characters described by Rieppel & Zaher (2002). The rst is character 6, which refers to the position of the ossied base of the crista trabecularis. In P. perroteti the base ends behind the frontalparietal suture (Rieppel & Zaher, 2002), whereas in P. aureus it terminates at the suture. Plectrurus perroteti shares this condition with more basal uropeltids (according to tree of Rieppel & Zaher, 2002), whereas P. aureus is most similar to Brachyophidium rhodogaster Wall, 1921, and to some, but not all, Rhinophis. Secondly, P. aureus differs from P. perroteti in character 13 (Rieppel & Zaher, 2002), whether or not the juxtastapedial recess is wide open laterally. This character was difficult to apply because the condition in P. aureus is intermediate between the states described and illustrated by Rieppel & Zaher (2002). The incongruity between P. aureus and P. perroteti may be lessened if this character was coded in a different fashion. As it stands, P. aureus shares a less open recess with the more basal taxa included by Rieppel & Zaher (2002), whereas P. perroteti more closely resembles the derived Uropeltis and Rhinophis. The third character that demonstrates variation between P. aureus and P. perroteti is character 18 of Rieppel & Zaher (2002), which describes the development of the posteroventral process of the dentary. The process was scored as reduced for P. perroteti by Rieppel & Zaher (2002), but it is clearly absent in P. aureus. Again, P. aureus is most similar to B. rhodogaster and some Rhinophis species, as well as Uropeltis and Pseudotyphlops Schlegel, 1839. Plectrurus perroteti exhibits a morphology like that of Platyplectrurus Gnther, 1868. In several instances we had difficulty interpreting the telegraphic character descriptions provided by Rieppel & Zaher (2002) and applying them to our description of P. aureus. This is not a problem unique
Table 1. Data matrix modied from Rieppel & Zaher (2002), with the addition of Plectrurus aureus (top) 3 1 0 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 0 0 0 1 ? 0 1 0 0 1 0 0 0 2 0 0 0 1 0 1 1 2 0 1 1 1 1 1 1 1 1 1 2 1 1 1 1 0 0 1 1 1 1 ? 1 2 2 2 1 1 0 1 1 1 1 1 1 1 1 2 1 0 1 1 2 2 2 2 2 2 1 0 0 1 1 1 1 1 0 0 1 0 1 0 2 1 1 1 1 1 1 0 1 1 2 2 1 0 0 1 1 0 1 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 2 0 1 1 1 0 1 2 2 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 0 0 0 1 0 1 0 0 1 1 0 1 0 1 0 1 0 2 0 0 0 1 0 0 0 1 0 1 0 2 0 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 1 1 1 1 1 2 1 1 1 1 1 1 0 29 2 1 1 2 2 1 2 2 2 2 2 1 1 30 1 1 1 1 1 1 1 1 1 1 0 0 0 31 1 1 1 1 1 1 1 1 1 1 0 0 0 32 1 1 1 1 1 1 1 1 1 1 0 0 0 33 1 1 1 1 1
1 0
1 0
Plectrurus aureus Melanophidium punctatum Melanophidium wynaudense Platyplectrurus 0 1 0 1 0 1 1 0 1 1
Uropeltis Teretrurus
1 1
1 0
Rhinophis drummondhayi Rhinophis sanguineus Plectrurus perroteti
1 1 1 1 0 0 0
1 1
1 1
Pseudotyphlops Anomochilus Cylindrophis
1 0 0
1 0 0
Anilius
0 0 1 1 1 0 1 0 0 0 0 1 0
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R. S. COMEAUX ET AL. posed by Rieppel & Zaher (2002), signifying a need to better understand variation in uropeltid snakes. Overall it is unclear where P. aureus ts among the hypotheses of uropeltid relationships. The majority of the features shared by P. aureus and P. perroteti are common to all other uropeltids found by Rieppel & Zaher (2002) to be more derived than Melanophidium. In other words, at the level of our current understanding of uropeltid anatomy these features are not phylogenetically informative for many taxa. Looking closely at the characters in which P. aureus and P. perroteti differ, both species exhibit a combination of states shared with both the derived and basal taxa hypothesized by Rieppel & Zaher (2002). It is apparent that P. aureus and P. perroteti may not necessarily be sister taxa, and that broader taxonomic sampling of uropeltid species will in all likelihood result in new phylogenetic hypotheses not predicted by existing analyses. This conclusion may not be surprising considering that monophyly has not been established for any non-monotypic genus included in any previous uropeltid phylogenetic analyses.
to their paper, but rather is one that impacts many morphological studies. Space constraints in journals and the additional expense incurred from publishing numerous illustrations appear to be the primary forces contributing to a persistent problem with adequate character descriptions. Careful attention to the description, and especially illustration, of morphological character states will greatly reduce the potential for confusion or misunderstanding in subsequent analyses (Joyce & Bell, 2004). For example, character 4 of Rieppel & Zaher (2002) is problematic because of unclear wording and a lack of visual representation. The buttressing contact between the anteromedial process of the maxilla and the anterolateral process of the vomer described by those authors for many uropeltids appears to be somewhat misleading, because this region of contact includes contributions from the maxilla, vomer, and premaxilla; in other taxa (e.g. some Rhinophis and Uropeltis species), there may also be a ventral projection from the septomaxilla that is visible in palatal view (J. Olori, C. Bell, pers. observ.), although this is not the case in our specimen of P. aureus. A second major issue affecting character description and interpretation stems from incomplete taxon sampling, which can result in the discovery of new character states that are not encompassed by the original character description. This situation can present obstacles for the inclusion of new taxa in subsequent analyses. When intermediate or previously unknown states are identied, characters must be redescribed, which necessitates rescoring the taxa used in the original analysis. Demonstrating this point, our study of P. aureus reveals multiple states intermediate to those documented by Rieppel & Zaher (2002). Character 1, for example, does not appear to adequately represent the total range of variation present among uropeltid snakes. In P. aureus, a clear contact between the maxilla and premaxilla is formed (unlike the conguration in Melanophidium Gnther, 1864, depicted by Rieppel & Zaher 2002), but the bones do not form a fully straight and tightly buttressing articulation, as was depicted for R. sanguineus by Rieppel & Zaher (2002). Likewise, Rieppel & Zaher (2002) reported that all of the uropeltids that they surveyed exhibited contact between the premaxilla and vomer within a welldened recess, rather than as an overlapping articulation (their character 26). This description is unclear and decient because we nd that the contact is complex and intermediate between these two states in P. aureus. Character 13 of Rieppel & Zaher (2002), which refers to the openness of the juxtastapedial recess, is also subjective because the character denitions are ambiguous. We found that the morphology in P. aureus is again intermediate to the states pro-
CONCLUSION
Existing morphological and molecular data do not yield a pretty picture of the probable stability of current uropeltid taxonomy. Our study demonstrates a need for the clarication of existing characters, as well as a need for improved taxon sampling in anatomical studies and phylogenetic analyses. Furthermore, the acquisition of new material, the discovery of additional characters for analysis, and an improved understanding of patterns of variation in all characters will play an important role in helping to recover a more thorough understanding of the evolutionary dynamics of this interesting and enigmatic group of snakes. High-resolution X-ray CT provides a ready means of gathering additional data on skeletal morphology, and permits the nondestructive utilization of existing large collections of preserved uropeltids for that purpose. The small size of most uropeltid skulls appears to place them near the limit of traditional CT scanning protocols, but a new generation of micro-CT scanners provides a promising technological advance that could yield higher-resolution anatomical data for these tiny snakes. Our efforts to gather and interpret such additional data are now underway.
ACKNOWLEDGEMENTS
We offer special thanks to J. Vindum of the California Academy of Sciences for facilitating the loan of the specimen used in this study. R. Ketcham and M. Colbert conducted the CT scanning and digital data acquisition. We beneted greatly from discussions
CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS with D. Cundall, C. Gans, D. Gower, and O. Rieppel, all of whom freely shared their thoughts and opinions about uropeltids. Special thanks also go to D. Cundall and O. Rieppel for providing advance copies of their unpublished works. B.-A. S. Bhullar, K. Claeson, E. Ekdale, T. LaDuc, M. Maga, J. Rodgers, and T. Rowe provided encouragement, advice, and comments on various aspects of this project. Funding for this research was provided by a University COOP fellowship award from The University of Texas at Austin to RC, and from the Jackson School of Geosciences at The University of Texas at Austin.
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Zoological Journal of the Linnean Society, 2010, 160, 118138.

MATERIAL AND METHODS

CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS

CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS

CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS

CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS

CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS

Plectrurus aureus Melanophidium punctatum Melanophidium wynaudense Platyplectrurus 0 1 0 1 0 1 1 0 1 1

Rhinophis drummondhayi Rhinophis sanguineus Plectrurus perroteti

Pseudotyphlops Anomochilus Cylindrophis

CRANIAL OSTEOLOGY OF PLECTRURUS AUREUS

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