Journal of Vertebrate Paleontology 22(4):812829, December 2002 2002 by the Society of Vertebrate Paleontology

TESTING THE PHYLOGENETIC RELATIONSHIPS OF THE PLEISTOCENE SNAKE WONAMBI NARACOORTENSIS SMITH
OLIVIER RIEPPEL1, ARNOLD G. KLUGE2, and HUSSAM ZAHER3 Department of Geology, The Field Museum, 1400 S. Lake Shore Drive, Chicago, Illinois 60605-2496; 2 Division of Reptiles and Amphibians, Museum of Zoology, The University of Michigan, Ann Arbor, Michigan 48109-1079; 3 Universidade de Sao Paulo, Instituto de Biociencias, Departamento de Zoologia, Rua do Matao, Travessa 14, Cidade Universitaria, 05508-900, Sao Paulo, Sao Paulo, Brazil
1

ABSTRACTThe skeletal material of Wonambi naracoortensis has been re-examined, and used to test its phylogenetic relationships. The skeletal description, and the resulting data matrix, concentrates on characters that can be assessed by observation (on the associated material of Wonambi kept at the South Australian Museum). Macrostomatan afnities of Wonambi prove to be well supported. The taxon nests within Macrostomata, above Xenopeltis and Loxocemus.

INTRODUCTION Scanlon and Lee (2000:420) recently placed three groups of fossil snakes at the base of ophidian phylogeny in support of their claim that the snake body-form arose in a large, nonfossorial ancestor. The extinct taxa in question are Pachyophiidae, Madtsoiidae, and Dinilysia. Supplementary information presented in support of their hypothesis consists of a data matrix that comprises 234 characters (191 osteological, 43 soft anatomical) coded for 20 in-group taxa rooted on a reconstructed ancestor using varanoids and mosasauroids as outgroups. A detailed review of this evidence is beyond the scope of this paper (see Rieppel and Kearney, 2002, for further comments). Here, we briey identify some problems with the characters and analysis presented by Scanlon and Lee (2000) in support of our claim that the skeletal anatomy of Wonambi naracoortensis Smith, 1976, deserves a detailed description in order to provide a basis for the re-assessment of the phylogenetic relationships of basal snakes. According to Scanlon and Lee (2000) the vector of their ancestor represents the plesiomorphic character states inferred from terrestrial varanoids and mosasauroids. These were chosen as outgroups because snakes were previously found to be the sister-group of mosasauroids (Lee, 1998; Lee and Caldwell, 1998, 2000). Inference of character states for the ancestor on that basis remains problematic because support for the monophyly of Pythonomorpha, claimed to be provided by previous studies (Lee, 1998; Lee and Caldwell, 1998, 2000), was found to be questionable (Rieppel and Zaher, 2000a, 2001). The use of a reconstructed ancestor also precludes assessment of the globally most parsimonious relationships. There is furthermore no rational justication known to us for leaving some multistate characters non-additive on the grounds that the respective character states may not be homologous (Scanlon and Lee, 2000: supplementary information, chars. 76, 85, 178)! Scanlon and Lee (2000) coded all fossil snakes other than Dinilysia at the family level. The Pachyophiidae sensu Lee et al. (1999) includes Pachyrhachis Haas, 1979, and Pachyophis Nopcsa, 1923. A number of characters coded by Scanlon and Lee (2000) for Pachyophiidae are not preserved in Pachyophis or Pachyrhachis, suggesting the inclusion of other taxa (e.g., Podophis [Rage and Escuille, 2000], for the position of the hyoid cornua and the presence of chevrons [Scanlon and Lee, 2000:characters 165 and 186 respectively]) in the Pachyophiidae without demonstration of their relationships. Rieppel and Zaher (2000a) doubted the sister-group relationship of Pachy-

rhachis and Pachyophis as originally proposed, but in their most recent paper, Lee and Caldwell (2000:925) retrieve that sister-group relationship again, based on a new character added to the analysis, viz. very small head relative to trunk region. In the absence of a quantitative assessment of relative head size throughout basal snakes we continue to call into question the monophyly of Pachyophiidae. Scanlon and Lee (2000) based their codings for Madtsoiidae primarily on the South Australian Museum specimen of Wonambi naracoortensis from the Pleistocene of Victoria, Australia (SAM P30178: Barrie, 1990; see also Smith, 1976), and on a newly described species, Wonambi barrei from Riversleigh, Queensland, Australia. We consider the monophyly of Madtsoiidae to be poorly supported at this time. Madtsoiids were traditionally referred to boids, with which they share the short and broad, massively built vertebral centra and (with some boids) paracotylar foramina (Rage, 1984). Scanlon (1992) retained madtsoiids in the Alethinophidia recognizing derived characters (anterior dentigerous process of palatine; subdivided trigeminal foramen) described for the skull of Wonambi naracoortensis by Barrie (1990). However, the association of Wonambi with madtsoiids was considered tentative by Rage (1984; see also Smith, 1976), and it still is. Madtsoiids are characterized by their large size, and by the presence of large parazygantral and paracotylar foramina (Rage, 1984). The presence of parazygantral foramina was considered an autapomorphy of madtsoiids by Rage (1998) who noted, however, The presence of a rather large bilateral and well-dened parazygantral foramen is not constant in Wonambi (Rage, 1998:128). In the description of the holotype of Wonambi naracoortensis, Smith (1976) noted that the identication of parazygantral foramina is difcult due to the pitted bone surface. Madtsoiids are further characterized by the presence of two or three mental foramina in the dentary (Hoffstetter, 1959). Barrie (1990:g. 7) illustrated a single mental foramen in the dentary of Wonambi, whereas the reconstruction of the skull of Wonambi naracoortensis presented by Scanlon and Lee (2000:g. 1c) shows a large anterior (same as that drawn by Barrie, 1990), and a much smaller, posterior mental foramen (for which there is no evidence in specimen SAM P30178). By comparison to other madtsoiid dentaries (Hoffstetter, 1959; Scanlon, 1997; Rage, 1998), a difference in size of the anterior and posterior mental foramina is unusual. No explicit diagnosis of Madtsoiidae based on shared derived characters has so far been offered, and differences of vertebral structure, along with differences in the count of men-

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FIGURE 2. Tooth-bearing maxillary fragment of Wonambi naracoortensis showing a smooth enamel surface.

FIGURE 1. The left maxilla of Wonambi naracoortensis (SAM P30178). A, Left lateral view; B, Medial view of anterior end. Scale bar equals 20 mm.

tal foramina, indicate poor support for the monophyly of the Madtsoiidae, including both Madtsoia and Wonambi. Problems of character coding for madtsoiids, based primarily on Wonambi, are related to the referral of fully disarticulated material to this taxon, and to inferences about missing elements drawn from neighboring bones. The basis for inferences about missing elements may furthermore be prejudiced by a priori hypotheses of relationships. For example, Scanlon and Lee (2000) interpret a putative facet on the maxilla as having accommodated a jugal. This inference was justied by reference to the presence of a jugal in Pachyrhachis and, possibly (Scanlon and Lee, 2000:418), in Dinilysia. However, a jugal is absent in Pachyrhachis (Rieppel and Zaher, 2000a), whereas the putative jugal of Dinilysia was previously considered to most likely (Lee, 1997:68) represent the anterior end of the ectopterygoid. Lee (1998:442) further stated: a reexamination of Dinilysia conrms that the fragment is not a jugal (A. Albino and M. Caldwell, pers. comm.). The same point is reiterated by Caldwell (2000). To refer completely disarticulated material to a given taxon is always problematic, especially if these elements come from deposits that also yielded other reptilian elements (e.g., Wells et al., 1984; Molnar, 1991). An example is provided by chevrons, claimed to be present in the tail of Wonambi (madtsoiids). Scanlon and Lee (2000:g. 2g) attribute a caudal vertebra with an articulated chevron to the South Australian Museum specimen of Wonambi naracoortensis (SAM P30178). However, Barrie (1990:g. 11, 2nd from right) illustrated a caudal vertebra of that same specimen, which is about three times the size of that gured by Scanlon and Lee (2000), but lacks a chevron. Furthermore, the preserved part of the tail of Wonambi naracoortensis (SAM P30178) shows no caudal vertebra that would correspond to the one gured by Scanlon and Lee (2000: g. 2g), i.e., with most of the neural arch and with the right transverse process missing. The conclusion is that this vertebra does not belong to the Naracoorte specimen of Wonambi. Instead, it comes from the Riversleigh locality (M. Hutchinson, pers. comm.), i.e., from a different locality and age. Furthermore, other madtsoiids have been described as having caudal vertebrae with hemapophyses (Scanlon, 1993), and a caudal vertebra of Madtsoia camposi was described as having short

hemapophyses, a boid character that also matches the palatine morphology of that same species (Rage and Werner, 1999). The representation of the cranial material referred to Wonambi naracoortensis by Scanlon and Lee (2000) is similarly misleading. Figure 2a in Scanlon and Lee (2000) shows the braincase of specimen SAM P30178 in left lateral view. The dorsal and ventral anterior processes of the prootic meet each other in front of the trigeminal foramen, thus excluding the seemingly articulated parietal from the anterior margin. In fact, only the right parietal is partially preserved in specimen SAM P30178, and Scanlon and Lee (2000:g. 2a) fail to notify the reader that their rendition of the left lateral view of the braincase of Wonambi includes the graphic transposition of the right parietal to the left side of the skull. Such a transposition of the parietal is not trivial, as it creates the illusion of characters for which there is no empirical basis. Thus, Wonambi appears to show a laterally crested anterior process of the prootic, which abuts against the posterior end of a lateral crest on the parietal (Scanlon and Lee, 2000:g. 2a). If indeed present, this character would be uniquely shared by Wonambi and Acrochordus, and might hence be taken as an indication of possible phylogenetic relationships. At this point, we refrain from commenting further on the analysis presented by Scanlon and Lee (2000). We believe that the comments given above justify the re-assessment of the phylogenetic relationships of Wonambi naracoortensis on the basis of a thorough description of the only associated material that also yielded a partial skull (SAM P30178). MATERIAL AND METHODS Our redescription of the skeletal anatomy is based on the personal inspection of the South Australian Museum specimen of Wonambi naracoortensis (SAM P30178) by two of the authors (AGK and OR). According to information kindly provided by John and Julie Barrie (pers. comm.), specimen SAM P30178 was washed into a cavern of the Henschkes Quarry Cave System, which resulted in the complete disarticulation of the skeletal material. The vertebral sequence of the specimen was arranged according to relative size. The cloacal and caudal regions remain incomplete. Cranial elements such as the (right) parietal, prootic, basisphenoid and basioccipitalexoccipital-opisthotic complex were found separately, and were later glued together (the parietal is now separate from the remainder of the braincase). The compound bone of the lower jaw was reconstructed from numerous fragments found in association (the articular facet and retroarticular process were broken off when the specimen was temporarily lost).

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Damage was inicted on the specimen not only during transport, but also by termites. For example, the left maxilla attributed to specimen SAM P30178 was found attached to a marsupial bone damaged by termites. Poorly preserved vertebrae were found inside extensions of termite nests. Fossil bone excavated from the Naracoorte cave systems is generally very fragile (Smith, 1971). In the case of the Naracoorte specimens of Wonambi, removal of the matrix from the fossil bones was found to be relatively easy (Barrie, 1990). However, the bones were found to be covered by a tough calcite patina. The most severe threat to inict damage on bone during preparation resulted from attempts to ake off this patina (J. Barrie, pers. comm.). It will be noted that some of the braincase elements, most notably the prootic, but also the posterior part of the dorsal surface of the basisphenoid, are completely devoid of the calcite patina. Comparative material used in this study is listed in Appendix 1. Abbreviations Anatomical ao.vc, anterior opening of Vidian canal; bo, basioccipital; bpt.p, basipterygoid process; cid, foramen for constrictor internus dorsalis nerve branch; cr.in, crista interfenestralis; cr.tu, crista tuberalis; eo, exoccipital; f.jug, jugular foramen; f.vest, vestibular fenestra; f.p, facet for parietal; f.st, facet for supratemporal; la.rsct, lateral opening of recessus scalae tympani; ls, laterosphenoid; opeo, opisthoticexoccipital complex; p, parietal; po.vc, posterior opening of Vidian canal; poz, postzygapophysis; pro, prootic; pro., prootic ange underlapping (posterior) trigeminal foramen; prz, prezygapophysis; psbs, parabasisphenoid; stp, stapes; s.trc, sella turcica; trf.ch, trigeminofacialis chamber (medial opening); zsp, zygosphenal tectum; V2, foramen for the maxillary branch of the trigeminal nerve; V3, foramen for the mandibular branch of the trigeminal nerve; VIIhy, foramen for hyomandibular branch of facial nerve. Institutional AMNH, American Museum of Natural History, New York; FMNH, Field Museum of Natural History, Chicago; IB, Instituto Butantan, Sao Paulo; LSUMZ, Museum of Natural Science, Louisiana State University, Baton Rouge; MNHN, Museum National dHistoire Naturelle, Paris; UMMZ, University of Michigan Museum of Zoology, Ann Arbor. THE SKELETAL ANATOMY OF WONAMBI NARACOORTENSIS The left maxilla is well preserved (Fig. 1; length: 81 mm as preserved). A total of 23 tooth positions can be identied, with 13 teeth preserved in ankylosis. The teeth are needle shaped with an expanded base, sharply recurved and pointed, comparable to those of Boa and Python. Tooth implantation is of the alethinophidian type (Zaher and Rieppel, 1999). The teeth show a smooth enamel surface (Fig. 2), and an anterolaterally as well as a posteromedially positioned cutting edge. The anterior tip of the maxilla is blunt, rounded, and only weakly turned medially. This is more closely comparable to the general macrostomatan condition than to the anilioid condition (Anilius, Cylindrophis), characterized by a maxilla that is much more distinctly turned medially in its anterior part. Accordingly, the anterior tip of the anilioid maxilla very closely approaches, or even contacts (sutured in uropeltines) the transverse process of the premaxilla, which contrasts with the macrostomatan condition (except Xenopeltis and Loxocemus), where a distinct gap persists between premaxilla and maxilla. Cundall (1995) reported limited independent mobility of the upper jaw arches in Cylindrophis. Contrary to Scanlon and Lee (2000:char. 11), Wonambi might therefore be assumed to have had a greater mobility of the maxilla relative to the premaxilla than is typical of ani-

FIGURE 3. The right palatine of Wonambi naracoortensis (SAM P30178). A, Dorsal view; B, Ventral view. Scale bar equals 10 mm.

lioids (Cundall, 1995). However, different degrees of mobility cannot be reliably inferred from fossils (Greene, 1994). Between the rst and second third of its length, the maxilla of Wonambi carries a prominent ascending process as is typical of madtsoiids (Fig. 1). The ascending process of Wonambi is developed to a somewhat lesser degree than was described for Nanowana (Scanlon, 1997). This is conrmed by the (only partially preserved) right maxilla (length: 45.6 mm as preserved). The rugose posterodorsal dorsal margin of the ascending process was taken as an indication of a tight, relatively immobile sutural contact between the prefrontal and the maxilla by Scanlon and Lee (2000), but transport of the bones as well as the calcite patina (left maxilla) or its removal (right maxilla) may well be responsible for such rugosity, which indeed indicates a damaged bone surface. Among extant snakes, a comparable ascending process is present in anilioids (most prominently developed in Cylindrophis), but is generally absent in macrostomatans. Posterolateral to the ascending process, the maxilla shows a shallow dorsolateral shoulder (Fig. 1A), which was interpreted as a facet receiving a jugal by Scanlon and Lee (2000; but see comments above). The medial maxillary process, establishing a contact with the palatine, is broken in the left maxilla (Fig. 1B), but it is well preserved in the right maxilla, where it is pierced by a canal that transmitted the superior alveolar nerve. The position of the medial maxillary process relative to the ascending process indicates that the rst was located below, not in front, of the orbit. The posterior end of the (left) maxilla gently tapers to a blunt tip. A series of alveolar foramina can be observed on the lateral surface of the maxilla. The palatine of Wonambi naracoortensis is characterized by a broad medial (vomerine) process arching over the choanal tube (Fig. 3). The anterior tip of the anterior dentigerous process (an alethinophidian feature) is missing (broken). As preserved, the palatine carries 11 tooth positions. The anteriormost (preserved) palatine tooth is split open along its length, which creates the (mistaken) impression of a striated enamel surface. The medial process originates from the posterior part of the

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FIGURE 4. The right parietal of Wonambi naracoortensis (SAMP 30178). A, Lateral view; B, Dorsal view. Scale bar equals 10 mm.

palatine. Its posterior margin is broad and transversely oriented (and probably incomplete), its medial margin is slightly angulated, and anteriorly the medial process terminates in a blunt tip which turns dorsally, enclosing the choanal tube both dorsally and medially. A closely similar palatine morphology is observed in Tropidophis among booids, but a broad medial process of the palatine is also characteristic of acrochordids (Hoffstetter and Gayrard, 1965). As is documented by representatives of these two genera, the presence of a broad medial process of the palatine does not in itself suggest a strong, abutting palatinevomer contact in Wonambi as was reconstructed by Scanlon and Lee (2000). At its posterior end, the body of the palatine carries a distinct groove on its dorsomedial surface for the reception of an anterior process of the pterygoid. Wonambi therefore shares the tongue-in-groove contact between pterygoid and palatine characteristic of some alethinophidians. The reconstruction of the skull of Wonambi presented by Scanlon and Lee (2000:g. 1) shows a pterygoid that closely resembles the macrostomatan condition, but no tooth bearing bone fragment referred to the specimen SAM P30178 can in our view be unequivocally identied as part of the pterygoid. The most revealing structure preserved in the skull of Wonambi is the braincase. As noted above, its components were found separately. It is incomplete dorsally, as the supraoccipital and most of the parietal are missing. Only part of the right parietal is preserved (Fig. 4). The parietal of Wonambi is expanded at its anterior end to form an anterior lateral wing (Barrie, 1990), which may have extended below a postorbital as is characteristic of basal (i.e., non-colubroid) macrostomatans above Xenopeltis and Loxocemus. However, to judge from the rounded lateral contours of that anterior expansion (Fig. 4B), the latter must be assumed to have been abraded to some degree. In Boa and Python, for example, the lateral expansion of the anterior end of the parietal is much more distinctly developed and terminates in a laterally pointed tip. Abrasion of the lateral wing of the parietal is indicated by its rounded edge, covered by the calcite patina. That the parietal did incur some damage (perhaps induced by termites: J. Barrie, pers. comm.) is furthermore indicated by the severe abrasion of a middle section of the sagittal crest (Fig. 4A). Scanlon and Lee (2000:g. 1) use a triangular pedicel located on the dorsal surface of the parietal as indicating the midline of the skull. If the ventrally descending ange of the parietal is arranged so as to

match the lateral margin of the parabasisphenoid, this triangular pedicel is seen to be located anterolaterally, most probably supporting a postorbital. In Wonambi, the anterior lateral expansion of the parietal extends posteriorly into a distinct, laterally projecting ridge that runs all along the (laterally) descending ange of the parietal (Fig. 4A). Scanlon and Lee (2000:g. 1a) reconstruct this ridge as abutting against a similar, laterally projecting ridge on the prootic (their alar process of the prootic: Scanlon and Lee, 2000:supplementary information, char. 116). This morphology represents an artifact of graphic reconstruction, however. In fact, the most posterior part of this lateral ridge on the parietal is, again, abraded and hence missing. In basal macrostomatans such as Xenopeltis and Loxocemus, a comparable ridge is absent. It is also absent in relatively small booids such as Casarea, Lichanura, Tropidophis, and Trachyboa. This may be a function of absolute size, since in small animals the brain is relatively larger, and the braincase therefore more inated. In Python, a distinct ridge with a sharp lateral edge is developed along the posterior margin of the anterolateral process of the parietal, but it becomes rather indistinct on the lateral surface of the posterior part of the descensus parietalis. However, a lateral parietal ridge, closely similar to the one observed in Wonambi, but with a sharp lateral edge, extends along the entire length of the descensus parietalis in Boa. In this taxon, the ridge extends posteroventrally to meet the anteroventral corner of the prootic below the trigeminal foramen. The dorsal view of the braincase exposes the dorsal surface of the basisphenoid (Fig. 5), which was illustrated both by Barrie (1990), and Scanlon and Lee (2000:g. 2b). An unusual feature of Wonambi, not encountered in any other squamate, relates to the sella turcica (Fig. 5B), which shows a more pronounced anterior margin as opposed to an indistinct posterior margin (Barrie, 1990:g. 9; Scanlon and Lee, 2000:supplementary information, char. 111). In snakes, the sella turcica is either very shallow with similarly pronounced anterior and posterior margins as in anilioids, or else the posterior margin is much more prominently developed, forming a crista sellaris (dorsum sellae) which may overhang the posterior part of the sella turcica to a variable degree (Rieppel, 1979a). This is in accordance with the fact that the crista sellaris ossies within the acrochordal cartilage, whereas the more anterior part of the sella turcica ossies in the membrane which closes the fossa hypophyseos ventrally. The condition of the crista sellaris in Wonambi might have resulted from some loss of bone around its posterior margin and behind it, perhaps as a consequence of the removal of the calcite patina. The same is suggested by grooves that were indicated by Barrie (1990) and that might be interpreted as partial exposure of the abducent nerve canals in the posterior part of the basisphenoid (dorsal view). However, these grooves might just as well represent simple cracks in the bone surface. However, at least part of the posterior margin of the sella turcica is preserved as smooth and shiny, nished bone surface, which indicates a low posterior margin. The Vidian canal does not open intracranially in Wonambi (Fig. 5B; Barrie, 1990; Scanlon and Lee, 2000:supplementary information, char. 109). The posterior opening of the Vidian canal can be located close to the laterodorsal margin of the basisphenoidparasphenoid complex at a level below the posterior margin of the trigeminal foramen, but well in front of the basisphenoidbasioccipital suture (Fig. 6A; Barrie, 1990). The anterior opening of the Vidian canal lies immediately in front of the base of the basipterygoid process (Fig. 6A). This conguration is closely comparable to that in Boa, where the Vidian canal again does not open intracranially. Indeed, the Vidian canal does not open intracranially in a variety of basal macrostomatans (Rieppel, 1979a). In others, the intracranial opening (primary anterior opening sensu Rieppel, 1979a) of the Vidian

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FIGURE 5. 10 mm.

The braincase of Wonambi naracoortensis (SAM P30178). A, Ventral view; B, The basisphenoid in dorsal view. Scale bar equals

canal closely approaches the anterior lateral margin of the basisphenoid, such that the intracranial opening of the Vidian canal is very short. In their rendition of the medial view of the braincase of Wonambi, Scanlon and Lee (2000:g. 2b) indicate what appear to be two openings in front of the (broken) right basipterygoid process, the anteroventral one of which they label as the anterior opening of the (right) Vidian canal. Indeed, the anterior opening of the Vidian canal is bipartite in Wonambi. The left side of the basisphenoid shows a single opening only in lateral view (Fig. 6A), but its internal subdivision is revealed in anterior view. The right side of the basisphenoid shows two juxtaposed anterior openings of the Vidian canal in lateral view as drawn by Scanlon and Lee (2000:g. 2b), indicating that the left side was subject to minor abrasion. A similar differentiation of the (extracranial) anterior opening of the Vidian canal is seen in most specimens of Boa (e.g., FMNH 22435), where the palatine nerve and artery appear to exit separately from the Vidian canal. Both Barrie (1990), and Scanlon and Lee (2000:g. 2b), illustrate rather distinct longitudinal grooves on the dorsal sur-

face of the basisphenoid (identied as grooves for the arteria ophthalmica by Barrie, 1990). These are deepest in their posterior part (more distinct on the left side of the basisphenoid), located lateral to the sella turcica and hence lateral to the openings of the cerebral carotids, but they become indistinct more anteriorly (Fig. 5B). As the cerebral carotids enter the sella turcica, they split into three branches. The anterior branches of either side run anteriorly on the dorsal surface of the basisphenoid, where they eventually meet each other to close the circle of Willis anteriorly (ODonoghue, 1912). Although shallow grooves generally outline this course of the anterior branches of the cerebral carotids on the dorsal surface of the basisphenoid of snakes (Rieppel, 1979a), the grooves observed on the dorsal surface of the basisphenoid of Wonambi cannot be interpreted as such, because they do not relate to the cerebral carotid foramina in their posterior part, and they do not converge anteriorly. In snakes, the abducent nerve generally enters the basisphenoid at a level somewhat behind the sella turcica. It continues anteriorly within a canal that pierces the laterally ascending wings of the basisphenoid (McDowell, 1967), and that opens on the dorsal surface of the basisphenoid at a level

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FIGURE 6. The skull of Wonambi naracoortensis (SAM P30178). A, The braincase in left lateral view; B, The braincase in ventral view; C, The braincase in posterior view. Scale bar equals 5 mm.

in front of the sella turcica, from where the nerve continues to travel intracranially until it leaves the skull through the optic foramen. Especially on the left side of the skull of Wonambi, it appears that the deep and distinctly delineated groove represents the abducent nerve canal piercing the laterally ascending wing of the basisphenoid, exposed through abrasion of supercial bone. The possibility also remains, however, that these grooves relate to the original sutural contact of the basisphenoid with the parietal (J. Barrie, pers. comm.), especially since the medial margin along the posterior part of the groove on the left side of the basisphenoid is lined by a sharp, clearly broken bony edge. The lateral wings of the basisphenoid are present in Wonambi (Fig. 5; contra Scanlon and Lee, 2000:supplementary in-

formation, char. 105), and developed to a similar degree as in Boa or Python. As is true of other alethinophidian snakes, the presence of lateral wings of the basisphenoid in Wonambi is best assessed in the medial view of the braincase (Scanlon and Lee, 2000:g. 2b). Comparing the disarticulated basisphenoid of Typhlops (Rieppel, 1979b:g. 3) with that of alethinophidians (Rieppel, 1979a) and with Wonambi is the best way to ascertain the presence of lateral wings of the basisphenoid in the latter genus. The presence of lateral wings of the basisphenoid is an alethinophidian character (McDowell, 1967). Scanlon and Lee (2000:supplementary information, char. 105) codied the presence or absence of lateral wings in terms of the outline of the sutural contact between the basisphenoid and prootic as seen in lateral view.

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fuse in front of the maxillary foramen, as in Python. In other basal macrostomatans, including Xenopeltis, Loxocemus, and Lichanura (Rieppel, 1979a), the parietal enters the anterior margin of the anterior (maxillary) trigeminal foramen, as is also the case in anilioids. In the disarticulated prootic of Boa, the anterior semicircular canal extends into the anterior dorsal process of the prootic, which is thus not directly comparable to the membranous alar process on the prootic of nonophidian squamates (Rieppel and Zaher, 2000b). Inspected in lateral view, the anterior end of the dorsal process of the (disarticulated) prootic of Boa appears to carry a lateral ridge slanting in a posteroventral direction (Fig. 8A), closely comparable to the same ridge seen in Wonambi (Figs. 6A, 7). However, this putative ridge on the prootic of Boa corresponds to the laterally exposed anterior edge of the element. From that edge, the anterior dorsal process of the prootic recedes inward as it underlaps the posterior margin of the descensus parietalis (in Boa, the supposed crista alaris of the prootic has a certain thickness as it houses the anterior semicircular canal; in nonophidian squamates, the alar process is a thin lamina of membrane bone added to the cupola anterior of the otic capsule and loosely overlapping the parietal). The close similarity of the disarticulated prootic of Boa with that of Wonambi suggests that the dorsal and ventral anterior processes of the prootic underlapped the descensus parietalis in an identical pattern in the two taxa. The absence of a laterosphenoid in the skull of Wonambi is best explained as a preservational artifact. Indeed, the trigeminal foramen identied by Scanlon and Lee (2000:supplementary information, char. 115) represents the medial (internal) aperture of the trigeminofacialis chamber (Rieppel, 1979a), which is an alethinophidian feature, probably correlated with the presence of a laterosphenoid. Such an interpretation implies, however, that the laterosphenoid was broken off or abraded in the skull of Wonambi. Closer inspection of the braincase does, indeed, indicate such damage to have occurred (note that the prootic in particular is completely devoid of the calcite patina). The trigeminal foramen of Wonambi is lined along its ventral margin by a laterally projecting, horizontally oriented ledge of bone that shows a deeply undercut posterior end (Figs. 5A, 6A, B, 7, 9A, B). This ledge of bone is particularly easily identied in the ventral view of the skull (Figs. 5A, 6B, 9B), and it is also distinct in the drawing presented by Scanlon and Lee (2000:g. 2a). Above this laterally projecting ledge is located the trigeminal foramen, the posterior margin of which is distinctly recessed medially relative to the lateral surface of more posterior parts of the prootic (Figs. 6A, 9A). The foramen for the exit of the hyomandibular branch of the facial nerve is exposed in lateral view within the recessed area behind the trigeminal foramen. Unless it is assumed to have had a separate exit, the palatine branch of the facial nerve would have emerged from the same foramen, turned ventrally and passed below the undercut posterior edge of the laterally projecting prootic ange on its way into the posterior opening of the Vidian canal. Wonambi is thus closely comparable to Boa, which also shows a laterally projecting ange of the prootic underlapping the posterior trigeminal (mandibular) nerve foramen. In Boa this ledge of bone is much more extensively developed, and it turns slightly upwards at its posterior end, thus concealing the facialis foramen in lateral view. Furthermore, in Boa, the palatine branch of the facial nerve does not pass behind the laterally projecting lappet of bone, but rather pierces it on its way to the posterior opening of the Vidian canal. A closely similar morphology is observed in Python (where the bony ledge underlapping the trigeminal foramen is less developed), and may well have existed in Wonambi if it is assumed that the laterally projecting ange underlapping the (posterior) trigeminal foramen was once more extensively developed, but now is incomplete. Indeed, the ledge underlapping the trigeminal fo-

In ventral view, the basicranium is distinctly keeled (Fig. 5A), another macrostomatan feature of Wonambi. Interesting with respect to this character is the ventral cresting of the basisphenoidbasioccipital suture, with paired spines projecting posteroventrally from the anterior margin of the basioccipital (Figs. 5A, 6B), again a condition closely comparable to the basicranium of an adult Boa or Python. The relief on the ventral surface of the basicranium tends to be much more weakly expressed if at all in booids of small size, and it is absent in anilioids and scolecophidians. A peculiarity of the basicranium of Wonambi is the ventral keel on the para-basisphenoid, which tapers off at about the level of the basipterygoid processes in the form of two low and diverging ridges that become indistinct anteriorly (Figs. 5A, 6B). A prominent feature on the basicranium of Wonambi is the basipterygoid processes which, in this snake, are rather longitudinally oriented (Figs. 5A, 6A, B). Similarly well developed basipterygoid processes are observed in basal macrostomatans, where they are more vertically (Boa) or transversely (Python) oriented. The lateral view of the braincase (Figs. 6A, 7) again exposes the prominently developed basipterygoid processes. In left lateral view, a distinct and elongate crista trabecularis runs along the basisphenoidal rostrum, with a straight, vertical, anterior margin against which the posterior end of the trabecula cranii abutted in this platybasic skull (Fig. 6A). Such well developed, elongate cristae trabeculares extending along the lateral margin of the parabasisphenoid complex is an alethinophidian feature, absent in scolecophidians (Rieppel, 1979b), as well as in the (tropibasic) nonophidian squamates. Among alethinophidians, the degree of development of cristae trabeculares, and their exposure in lateral view, is variable: they may be elongate or short, and they may or may not be concealed in lateral view by a ventrally projecting parietal crest. In anilioids, Boa, and Wonambi, the crista trabecularis is exposed in lateral view with a straight anterior termination. In other macrostomatans, such as Python, the contact of the cartilaginous trabecula cranii with the ossied crista trabecularis is recessed in a notch formed by the para-basisphenoid. Due to the presence of well-developed basipterygoid processes, the morphology of Wonambi, as exposed in lateral view, most closely resembles the condition seen in Boa, but Wonambi differs from genera such as Boa and Python in that it shows a relatively broad para-basisphenoidal rostrum (Figs. 5, 6B). The anterior end of the para-basisphenoidal rostrum is not preserved in Wonambi. The prootic is, arguably, the element most difcult to interpret in the skull of Wonambi. As shown by Scanlon and Lee (2000:g. 2a, and supplementary information, char. 115), the prootic appears to enclose a single trigeminal foramen with what can be described as a ventral and a dorsal anterior process (Figs. 6A, 7). This interpretation of the lateral braincase wall of Wonambi implies the absence of a laterosphenoid (Rieppel, 1976). According to Scanlon and Lee (2000:g. 2a) the dorsal anterior process of the prootic (their alar process) carries a distinct, posteroventrally sloping lateral crest, which they believe to have lined up with the lateral crest on the parietal. Furthermore, the anterior tip of this alar process forms a ventrally projecting spur, which contacts the dorsal margin of the anterior tip of the inferior anterior process of the prootic, thus closing the trigeminal foramen anteriorly and excluding the parietal from the anterior margin of the latter. A disarticulated skull of Boa (Fig. 8; FMNH 22435) suggests a different interpretation of the braincase in Wonambi. In Boa, the superior and inferior anterior processes of the prootic also establish an anterior contact with one another, thus excluding the parietal from the anterior margin of the anterior trigeminal (maxillary) foramen. Whereas the sutural contact of the prootic in front of the anterior trigeminal foramen remains distinct in Boa, the dorsal and ventral processes of the prootic may fully

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FIGURE 7. The braincase of Wonambi naracoortensis (SAM P30178). Top, Left lateral view; Bottom, The otic complex in left lateral view. Scale bar equals 2 mm.

ramen is so deeply undercut posteriorly that it might perhaps have been pierced by a separate (but now posteriorly incomplete) palatine nerve foramen. A distinct, posteriorly open groove connects the posterior recess in the bony ledge underlapping the trigeminal foramen with the posterior opening of the Vidian canal (Figs. 6A, B, 9A, B). In Boa and Python, the ange of bone underlapping the posterior trigeminal foramen constitutes the ventral part of the laterosphenoid, the more dorsal part of which is missing in Wonambi. But there is, in Wonambi, another distinct and laterally projecting bony rim, which extends backwards from the posterior dorsal corner of the putative trigeminal foramen (Figs. 6B, 7, 9A). This rim, again clearly delineated in the drawing presented by Scanlon and Lee (2000:g. 2a), corresponds exactly to the area of the prootic from which emerges the dorsal part of the laterosphenoid in Boa (Fig. 8). This level of the emergence of the dorsal part of the laterosphenoid from the prootic also demarcates the ventral margin of the facet which receives the supratemporal, both in Wonambi and in Boa. We

therefore conclude that Wonambi originally had a laterosphenoid and hence a bipartite trigeminal foramen, but that this element, an alethinophidian character, was lost in the fossil due to breakage. As a consequence, the medial aperture of the trigeminofacialis chamber appears as the lateral opening of an undivided trigeminal foramen in Wonambi. The morphology of the otic region of the skull of Wonambi is easily reproducible by abrasion of the laterosphenoid in an extant macrostomatan snake such as Boa (Fig. 8B), or Python. The facet on the prootic and opisthotic for the reception of the supratemporal is characteristically sculptured in Wonambi (Figs. 6A, 7), as it also is in Boa or Python. In these taxa, as well as in Wonambi, the supratemporal facet extends anterodorsally as it tapers to a pointed anterior tip which lies above, yet at the level of, the anterior margin of the (anterior, maxillary) trigeminal foramen. The reconstruction of the relations of the anterior end of the supratemporal proposed for Wonambi by Scanlon and Lee (2000:g. 1) is thus erroneous. Posteriorly, the facet for the supratemporal becomes broader both in Won-

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be developed, albeit to a lesser degree, posteroventral to the jugular foramen in macrostomatans. The crista circumfenestralis was originally described to be absent in Dinilysia (Estes et al., 1970), but Rage (1984:68) conrmed its presence in this fossil. The anterior circumference of the stapedial footplate is recessed below and behind a distinct crest, yet the juxtastapedial recess remains widely open posteriorly due to the poor development of the crista tuberalis in Dinilysia (Estes et al., 1970:g. 6). Indeed, the crista tuberalis does not close the juxtastapedial recess posteriorly in scolecophidians, Dinilysia, anilioids, Xenopeltis and Loxocemus. This contrasts with the condition observed in other macrostomatans, where the crista tuberalis forms a distinct, laterally projecting ange closing the juxtastapedial recess posteriorly. As is seen in the posterior view of the skull (Fig. 6C), the juxtastapedial recess is also closed posteriorly in Wonambi, even though the jagged lateral edge of the crista tuberalis indicates some degree of damage. This more extended degree of differentiation of the crista tuberalis conceals the jugular foramen in lateral view in Wonambi and macrostomatans. By contrast, the jugular foramen is always exposed in the lateral view of the skull in scolecophidians and anilioids, but it may also open laterally in booids of small size (Rieppel, 1979a). The crista circumfenestralis strongly converges upon the stapedial footplate in some macrostomatans such as boids or pythons, but much of the stapedial footplate remains exposed in lateral view in basal macrostomatans, as is also the case in Wonambi (hypothesized to represent the plesiomorphic state by Scanlon and Lee, 2000). In nonophidian squamates, the crista interfenestralis separates the oval from the round window. In scolecophidians and anilioids, the crista interfenestralis does not participate as a separate element in the formation of the ventral rim of the crista circumfenestralis, but remains recessed behind it instead. In macrostomatans, the crista interfenestralis does appear as an individualized component in the ventral rim of the crista circumfenestralis (Rieppel, 1979a), and such is also the case in Wonambi (Figs. 6A, 7; Barrie, 1990:g. 8, labeled as opisthotic; Scanlon and Lee, 2000:g. 2a, labeled as cri cir). In some macrostomatans (e.g., Boa, Fig. 8), this ventral foot process of the crista circumfenestralis may not ossify, which leaves a recess in the ventral margin of the crista circumfenestralis in the macerated skull. According to Scanlon and Lee (2000:supplementary information, char. 125), Wonambi shares with Dinilysia a derived character, viz. a proportionately large stapedial footplate, with a diameter at least subequal to that of the foramen magnum. Comparison of gure 2a (for the size of the fenestra vestibuli) with gure 2b (for the vertical diameter of the foramen magnum) in Scanlon and Lee (2000) indicates that the stapedial footplate of Wonambi is smaller than the foramen magnum, which differs from the size relations shown for these structures for Dinilysia by Estes et al. (1970:g. 6). The fenestra vestibuli is deeply recessed within the juxtastapedial recess, and its diameter hence difcult to measure accurately without risking to damage the fossil (Fig. 7). However, its diameter can be estimated to be 4.8 mm, while the diameter of the foramen magnum is 7.1 mm. The relative size of the stapedial footplate of Wonambi therefore rather closely matches that of the anilioids Anilius and Cylindrophis. In general, however, the relative size of the stapedial footplate on the one hand, and of the foramen magnum on the other, are variable in squamates, and are furthermore size dependent (the relative size of the vestibular fenestra becomes larger as the skull becomes absolutely smaller). Behind the fenestra ovalis, Scanlon and Lee (2000:g. 2a) indicate the presence of a fenestra pseudorotunda. The latter opening was found to be an anilioid synapomorphy instead (Rieppel, 1979a; Tchernov et al., 2000). In fact, the opening designated fenestra pseudorotunda by Scanlon and Lee

FIGURE 8. The braincase of Boa constrictor (FMNH 22435) in left lateral view. A, The laterosphenoid is retained; B, The laterosphenoid is removed, exposing the trigeminofacialis chamber. Scale bar equals 5 mm.

ambi and in Boa or Python, as it extends on to the dorsolateral surface of the opisthoticexoccipital complex. The pattern of overlap of the prootic and opisthotic-exoccipital in this area is identical in Wonambi (Fig. 6A) and Boa (Fig. 8A), including the differentiation, on the opisthotic-exoccipital, of a more or less transversely oriented, posterodorsally projecting rim with a jagged dorsal margin. This rim distinctly overhangs the dorsal surface of the exoccipital above the foramen magnum, as is also shown by Scanlon and Lee (2000:g. 2b). The same differentiation of the overlap of the supratemporal with the prootic and opisthotic-exoccipital can be seen in Python, where the opisthotic-exoccipital is even more distinctly drawn out posteriorly than is the case in Boa. Wonambi can therefore be concluded to share a distinctly macrostomatan morphology of the oticooccipital complex. Again, the sculpturing and protuberances seen in the differentiation of the supratemporal facet in Wonambi, Boa and Python, are much less expressed or entirely absent in booids of smaller size. The differentiation of the crista circumfenestralis in Wonambi is macrostomatan, too (Fig. 7). The crista circumfenestralis is a bony rim surrounding the oval and round fenestrae as well as the footplate of the stapes, thus dening the juxtastapedial recess which houses the pericapsular sinus of the perilymphatic system (Rieppel and Zaher, 2000b). The crista circumfenestralis of snakes is derived from the crista prootica (located above and in front of the oval fenestra), crista interfenestralis (located between the oval and the round fenestra), and the crista tuberalis (located between the round fenestra and the jugular foramen) of nonophidian squamates. At its ventral end, the crista tuberalis forms a distinct posterior projection in Wonambi (Fig. 7), which is not present in scolecophidians and anilioids, but which may

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FIGURE 9. The braincase of Wonambi naracoortensis (SAM P30178). A, The trigeminofacialis chamber in left lateral view. The arrow points to the broken dorsal rim of the laterosphenoid; B, The braincase in ventrolateral view, showing the posteriorly undercut prootic ange which underlaps the (posterior) trigeminal foramen.

(2000) in the skull of Wonambi corresponds to the lateral aperture of the recessus scalae tympani (Barrie, 1990). Kamal (1971) noted important differences in the developmental fate of the embryonic ssura metotica in nonophidian squamates and in snakes. The embryonic skull of most snakes retains an occipitocapsular ssure between the occipital arch and the otic capsule, which is closed in lizards by the fusion of the occipital arch with the posterior wall of the otic capsule above the posterior part of the ssura metotica. In all squamates, the recessus scalae tympani is located in the anteroventral part of the occipitocapsular recess (deBeer, 1937). Unlike nonophidian squamates, the recessus scalae tympani in embryonic snakes becomes separated from the jugular foramen by a strut of cartilage that grows downward from the medial wall of the otic capsule to meet the lateral margin of the basal plate (Kamal, 1971; Rieppel, 1988). In basal snakes such as scole-

cophidians (Rieppel, 1979b) and anilioids (Rieppel, 1979a), the vagus nerve passes through the ventral part of the occipitocapsular ssure, closely behind the recessus scalae tympani. Correspondingly, the jugular foramen is located in a ventral position in the ossied skull, and is always exposed in the lateral view of the skull. By contrast, the jugular foramen passes through the dorsal part of the occipitocapsular recess in macrostomatans other than Xenopeltis and Loxocemus, as is the case in boas, pythons, and in Wonambi (Fig. 6C). Variations exist with respect to this character among macrostomatans, however, and these may again be correlated with absolute size. The jugular foramen tends to open laterally in small booids with a relatively larger brain and a relatively larger otic capsule. It is interesting to note that snakes such as some anilioids (Rieppel, 1979a:g. 2), Boa and Python (Rieppel, 1979a:g. 6B) actually retain the occipitocapsular ssure in the ossied skull,

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FIGURE 10. The left dentary of Wonambi naracoortensis (SAM P30178). A, Left lateral view; B, Left lateral view of anterior end. The arrow points to the single mental foramen. Scale bar equals 10 mm.

which is then more appropriately termed an occipitocapsular recess. No such distinct recess is present in the skull of Wonambi. As indicated by the drawing presented by Scanlon and Lee (2000:g. 2b), the medial side of the oticooccipital complex again shows extensive damage to the medial wall of the otic capsule, which obscures the acoustic recess (sensu Oelrich, 1956; housing the ganglion of the acoustic nerve and transmitting the auditory nerve branches to the inner ear) as well as the relations of the perilymphatic foramen (located in the posterior wall of the cavum cochleare). The medial aperture of the recessus scalae tympani is located on the juncture of the exoccipital, basioccipital and opisthotic. It is connected with the dorsally positioned vagus foramen by a shallow groove (rather than by a deep occipitocapsular recess, as is present in some alethinophidians). However, this groove follows the suture between exoccipital and opisthotic, which hence remain separate on the medial side of the braincase of Wonambi, indicating the presence of an occipitocapsular ssure early in ontogeny. Three separate foramina for the hypoglossal nerve can be identied in the medial wall of the exoccipital of Wonambi. A similar number may obtain among extant snakes (Rieppel, 1979a), although the number of separate hypoglossal foramina varies extensively. The dorsal view of the braincase shows a characteristic notch in the posterior margin of the exoccipital of Wonambi, another similarity shared with boas and pythons. The medial margin of the exoccipital trends posteromedially in Wonambi, as it does to a somewhat lesser degree in Boa. Both Barrie (1990) and Scanlon and Lee (2000) reconstruct Wonambi with a narrow contact of the exoccipitals dorsal to the foramen magnum. It should be noted, however, that if inspected in dorsal view, the exoccipitals of Boa may establish a broad or a narrow contact behind the supraoccipital spine, or, especially in large individuals, may fail to contact each other. In ventral view, however, the exoccipitals can always be seen to meet in a broad medial contact below the supraoccipital spine, which was probably the case in Wonambi. The medial margin of the exoccipital seems to have been subject to some abrasion along its medial margin.

FIGURE 11. Medial view of the anterior end of the left dentary. The arrow points to the anteriormost tooth which shows the smooth internal (dentine) surface of the pulp cavity. Scale bar equals 10 mm.

The jugular foramen of Wonambi is exposed in the posterior view of the opisthotic-exoccipital (Fig. 6C). It is internally subdivided into a larger dorsal and a smaller ventral foramen. A single hypoglossal foramen can be identied in the lateral wall of the exoccipital next to the jugular foramen. Of the lower jaw elements, the left dentary is well preserved (Fig. 10), carrying 25 tooth positions (Barrie, 1990), with 11 teeth preserved in ankylosis. The (preserved) dentary teeth are strongly recurved, and diminish in size from front to back. Tooth implantation is again of the alethinophidian structure (Zaher and Rieppel, 1999). The enamel surface appears corrugated by the calcite patina, but the anteriormost tooth is broken near its base, documenting (unequivocally) the absence of plicidentine (Fig. 11). The lateral surface of the dentary reveals a single mental foramen (Fig. 10B). This contrasts with the presence of two or three mental foramina in Nanowana (Scanlon, 1997), as well as in South American madtsoiids (Hoffstetter, 1959; Rage, 1998). The posterior dentigerous process of the dentary is prominent and elongate, as is characteristic of macrostomatans. The posterior ventral process of the dentary appears to be incomplete. The splenial is preserved in its posterior part, but broken anteriorly. The posterior margin of the splenial is incomplete,

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FIGURE 12. The right compound bone of Wonambi naracoortensis (SAM P30178). A, The anterior portion as reconstructed by Barrie (1990); B, The articular portion in lateral view; C, the articular portion in dorsal view. Scale bar equals 10 mm.

suggesting a deeply concave conguration, but it does preserve the smooth anterior margin of the anterior mylohyoid foramen, which was located some distance in front of the original posterior margin of the element. The anterior tip of the splenial is broken, exposing Meckels canal in medial view. It is also possible, however, that splinters of bone now positioned on the medial surface of the anterior part of the dentary represent parts of the broken ventral margin of the dentary itself. The right compound bone of Wonambi is well preserved (Fig. 12), although its anterior end, entering between the two posterior prongs of the dentary, is incomplete. The articular facet is saddle-shaped in the transverse plane, and its anterior margin rises to a higher level than its posterior margin (Fig. 12B, C). Scanlon and Lee (2000:character 146) nd madtsoiids to share with Dinilysia a plesiomorphic trait, i.e., the incomplete fusion of the articular and surangular in the region of the articular facet. This character is difcult to evaluate, as it may reect nothing but the transition from the smooth, nished bone surface to the unnished bone underlying the cartilaginous articular surface, a transition which is distinct in Wonambi, as it is in all other snakes. No other separation of the (ossied) articular from neighboring dermal bones could be identied in Wonambi, but it should be noted that the compound bone is subject to numerous cracks (as it was reconstructed from numerous separate fragments: J. Barrie, pers. comm.). The retroarticular process is short, stubby, and shows a smoothly rounded posterior surface (Fig. 12B). The compound bone forms a very shallow coronoid process that rises from its lateral margin at a level just behind the point to which the posterior dentigerous process of the dentary would have extended. The medial facet formed by the compound bone and its shallow coronoid process was reconstructed to have received a coronoid bone by Scanlon and Lee (2000:g. 1). No remains of the coronoid bone are preserved, however. The ad-

ductor fossa is shallow and elongate. The adductor fossa is also shallow yet reduced in acrochordids, and it may be so in colubroids. In more basal snakes, the adductor fossa is generally well delineated, and in basal macrostomatans in particular the compound bone forms a dorsally projecting ange that distinctly delineates the adductor fossa medially. In its reduced condition (Acrochordus), the adductor fossa is much shorter than that of Wonambi. In this character, therefore, Wonambi most closely resembles Cylindrophis (Rieppel and Zaher, 2000a:g. 7), as well as the condition seen in basal macrostomatans such as Xenopeltis and Loxocemus. Scolecophidians lack an adductor fossa in the compound bone (Rieppel and Zaher, 2000a). Middle or posterior trunk vertebrae of Wonambi are short, wide and massive as is characteristic of the Boidae (Rage, 1984:15). The hypapophyses, present on the anterior trunk vertebrae, are reduced to shallow and ventrally rounded hemal ridges in the middle and posterior trunk region. The presence of hypapophyses on the anterior trunk vertebrae only is another character shared with boids (except Candoiini and Bolyerinae: Rage, 1984). The centrum is short and wide, and shows relatively small subcentral foramina on either side of the hemal ridge. The neural spine is distinct, massive, and sloping backwards. The paradiapophysis is distinctly separated into a dorsal, laterally convex diapophyseal part, and a ventral, slightly concave, parapophyseal part. The separation of a diapophyseal and parapophyseal part on the paradiapophysis is an alethinophidian character which is absent in scolecophidians (Rage, 1984:8). There is no hint of a prezygapophyseal process in Wonambi. As in all snakes (Hoffstetter and Gasc, 1969), the prezygapophyseal and zygantral articular facets are separated by a non-articular area in Wonambi. The zygosphenal tectum has a straight, or even slightly convex anterior margin in vertebrae assumed to be part of the anterior trunk region in Wonambi (Fig. 13B). Vertebrae attributed to the posterior trunk region show a distinctly notched zygosphenal tectum (Fig. 13A). A straight zygosphenal tectum is frequently taken as diagnostic of Serpentes, as it differs from the deeply concave zygosphenal tectum seen in those nonophidian squamates that have zygosphenes. However, a lot of confusion appears to surround this character. Caldwell and Cooper (1999: 443) indicate the presence of a relatively weakly concave zygosphenal tectum in Coniasaurus (by reference to other nonophidian squamates), and note the variability of this character among snakes in general. An isolated vertebra from the Early Cretaceous of Spain, with a distinctly notched, i.e., V-shaped zygosphenal tectum was identied as the earliest snake by Rage and Richter (1994:563), who also noted that the depth of indentation is magnied by distortion. Later, Rage and Werner (1999:85) claimed that new data on mid-Cretaceous varanoid squamates cast doubt on the referral of this fossil [vertebra] to snakes. Still later, Rage and Escuillie (2000:15) considered the distinctly notched zygosphenal tectum of Podophis a primitive feature of fossil snakes. But of the fossil snakes included in their phylogenetic analysis by Scanlon and Lee (2000), the zygosphenal tectum is unknown in Pachyrhachis (Lee and Caldwell, 1998; also unknown in Haasiophis: Tchernov et al., 2000), and whereas Rage and Albino (1989:437f) note that the indentation of the zygosphenal tectum is distinct in Dinilysia, this statement is contradicted by their illustrations of the vertebrae of Dinilysia in dorsal view (Rage and Albino, 1989:gs. 3C, 4A; see also Hecht, 1982:g. 1). According to these illustrations, the anterior margin of the zygosphenal tectum is straight (Hecht, 1982:g. 1; Rage and Albino, 1989:g. 4A), or even slightly convex (Rage and Albino, 1989:g. 3C). Variability of this character appears extensive, and is documented along the vertebral column of Wonambi (SAM P30178), which renders its usefulness for phylogenetic analysis questionable. In addition to subcentral foramina, the vertebrae of Wonambi

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g. 2f), shows short, button-like projections near the posterior end of the ventral surface of the centrum. These were interpreted as facets for the reception of an articulated chevron by Scanlon and Lee (2000:supplementary information, chars. 185, 186), but these short projections might just as well be identied as modied hemapophyses. Finally, it should be noted that typical hemapophyses have been described for putative madtsoiids by Scanlon (1993), and by Rage and Werner (1999). The only articulated fossil snake reported so far with putative chevrons is Podophis from the Cenomanian of Lebanon (Rage and Escuillie, 2000). Some confusion surrounds these elements as well, because unlike Scanlon and Lee (2000), Rage (in Rage and Escuillie, 2000) considered the lack of fusion of hemapo physes to the centrum as the dening character of chevrons. This is why Rage (in Rage and Werner, 1999) considered mosasaurs with fused chevrons to have hemapophyses. Thus, it seems that chevrons are a plesiomorphic trait at the level of Serpentes, and hence cannot be used in an argument supporting the basal position of Wonambi. Ribs of snakes have been characterized as holocephalous, with a posterodorsal protuberance of their proximal end identied as tuberculiform process (Rage, 1984). Scanlon and Lee (2000:g. 2h) identify a dorsal tubercle (presumably the tuberculiform process) on the ribs of Wonambi, as is conrmed here. DISCUSSION AND CONCLUSIONS In the absence of support for the monophyly of Madtsoiidae, we limit our discussion of phylogenetic relationships to the genus Wonambi. Two species have been assigned to this genus, Wonambi naracoortensis Smith, 1976, and Wonambi barrei Scanlon and Lee, 2000. Of these two species, only Wonambi naracoortensis is known from associated material, most notably the South Australian Museum specimen SAM P30178, which is also the only specimen with preserved parts of the skull. The second species, Wonambi barrei, is known only from disarticulated and dissociated material which comes from the OligoMiocene sites at Riversleigh, Queensland, deposits which have yielded an abundance of other reptilian faunal remains (Molnar, 1991; see also Archer et al., 1991). The referral of isolated elements to a given species is always problematic (see the discussion of the occurrence of chevrons above), for which reason we prefer to base our phylogenetic analyses on articulated or associated specimens wherever possible. In essence, therefore, we use the closely associated material of Wonambi naracoortensis (SAM P30178) as basis for the evaluation of the phylogenetic relationships of this genus. The description of the braincase of Wonambi presented above suggests an alethinophidian, indeed a macrostomatan status of this genus. The same is indicated by associated postcranial remains. In order to test the phylogenetic relationships of Wonambi, we composed a data matrix for the skeletal variation discussed in this paper. We restrict the coding of characters to those that are actually preserved in Wonambi. We code these characters for the following terminal taxa: Dinilysia, Haasiophis, Pachyrhachis, Podophis, Wonambi, Scolecophidia (based on Leptotyphlops, Liotyphlops, and Typhlops), Anilioidea (based on Anilius, Cylindrophis, and uropeltines), Xenopeltis, Loxocemus, Boa, Python, Exiliboa, Trachyboa, Tropidophis, Ungaliophis, and Acrochordus. The inclusion of Podophis in this analysis is based strictly on the description given by Rage and Escuille (2000); a proper assessment of its taxonomic status must await its monographic description. We rooted the analysis on varanoids (Varanus, Lanthanotus, and Mosasauroidae), and on Dibamidae plus Amphisbaenia. The characters used in the analysis, as well as the corresponding data matrix, are given in Appendices 2 and 3.

FIGURE 13. Vertebrae of Wonambi naracoortensis (SAM P30178). A, Posterior presacral vertebra with notched zygosphenal tectum; B, Anterior presacral vertebra showing a zygosphenal tectum with a straight anterior margin. Scale equals 10 mm.

also share paracotylar and parazygantral foramina. Large paracotylar foramina also occur in some boids, in acrochordids (Hoffstetter and Gayrard, 1965), and in a variety of colubroids (Hoffstetter and Gasc, 1969). By contrast, the presence of parazygantral foramina is considered an autapomorphy of Madtsoiidae (Rage, 1984; Rage and Werner, 1999). In this respect, however, Wonambi differs from madtsoiids such as Gigantophis, Madtsoia (Rage, 1984:g. 18), Nanowana (Scanlon, 1997), and Yurlunggur (Scanlon, 1992), in that the presence of parazygantral foramina is variable, and the parazygantral foramina may be paired (Smith, 1976). In the holotype of Wonambi naracoortensis, the region in which the parazygantral foramina would be located is pitted, rendering the unequivocal identication of these foramina difcult (Smith, 1976:g. 2). Accordingly, Rage (1984:30; see also Rage and Werner, 1999) expressed doubts as to the putative homology of the parazygantral foramina in Wonambi and in other genera referred to madtsoiids. The presumed cloacal and anterior caudal vertebrae of Wonambi share distally bifurcated lymphapophyses, the posterior ones of which are fused to the centrum, as is typical of snakes (Barrie, 1990:g. 11). As was already discussed above, Scanlon and Lee (2000:g. 2g) describe and illustrate an isolated caudal vertebra with an articulated chevron from the Riversleigh deposits, which they erroneously attribute to specimen SAM P30178 (M. Hutchinson, pers. comm.). A caudal vertebra referred to Wonambi barrei, gured by Scanlon and Lee (2000:

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FIGURE 14. The phylogenetic relationships of the fossil snake Wonambi. Wonambi groups with other fossil snakes such as Haasiophis, Pachyrhachys and Podophis within Macrostomata. Bootstrap percentages (1000 replications) and Bremer support indices are given for relevant nodes. See text for further discussion.

Using the Ratchet in WinClada (Nixon, 1999; 21 taxa, 45 non-additive characters; ratchet parameters as follows: 500 was the number of iterations per replication, 50 trees were held per iteration, 12 characters were sampled) yielded 50 equally most parsimonious trees (all multistate characters non-additive), with a tree length of 98 steps, an ensemble consistency index of 0.57 and a retention index of 0.81. The strict consensus tree (10 nodes collapsed; Fig. 14) shows Haasiophis, Pachyrhachis, Podophis, and Wonambi, to form a monophyletic clade with Boa, Python, Exiliboa, Trachyboa, Tropidophis, Ungaliophis, and Acrochordus (Bremer support index [Bremer, 1988]:2). Within that clade, relationships remain unresolved except for a grouping of Exiliboa with Ungaliophis. Macrostomatans other than Loxocemus and Xenopeltis share with Wonambi, Haasiophis, Pachyrhachis, and Podophis nine unambiguously optimized (Fitch optimized:Fitch, 1971) synapomorphies. These are: maxilla with an anterior end that is only weakly bent inwards (6[0]), absence of a small horizontal shelf on the medial surface of the anterior end of the maxilla (7[1]), location of the medial maxillary process below the orbit (9[1]), presence of the lateral wing of the parietal (14[1]), parietal excluded from (anterior) trigeminal foramen (27[1]), presence of the short and reduced opisthotic paroccipital process (33[1]), the posteriorly closed juxtastapedial recess (unknown in Haasiophis, and Pachyrhachis) (34[2]), crista tuberalis conceals the jugular foramen in lateral view (37[1]), presence of a distinctly projecting medial margin of the adductor fossa (39[1]). The relative relationships of Xenopeltis and Loxocemus remain unresolved, but the two taxa are found to be included with the clade discussed above in the monophyletic Macrostomata (Bremer support index:2). The monophyly of Macrostomata is supported by seven unambiguous synapomorphies: marginal teeth elongate, needle-shaped, distinctly recurved (4[2]), ascending process of maxilla absent (5[1]), maxilla projects distinctly beyond posterior margin of orbit with a broad, at surface (8[2]), basicranium keeled (22[1]), presence (assumed for Wonambi) of a free ending posterior process of the supratemporal (32[1]), crista interfenestralis forms an individualized component of the ventral rim of the crista circumfenestalis (36[1]), elongate posterior dentigerous process of the dentary present (38[2]). The monophyly of Alethinophidia (which includes Anilioidea and Macrostomata; Bremer support index: 4) is supported

by four unambiguous synapomorphies: anterior dentigerous process of the palatine present (10[1]), tongue-and-groove joint between pterygoid and palatine (12[1]), lateral wings of the basisphenoid present (21[1]), laterosphenoid present (28[1]). The monophyly of a clade that includes Dinilysia and Alethinophidia (Bremer support index: 4) is supported by six unambiguous synapomorphies: alethinophidian type of tooth implantation (1[1]); maxillary teeth robust and recurved (4[1]), pterygoid teeth present (13[1]), posterior dentigerous process of the dentary present (38[1]), hemapophyses present (42[1]) and paradiapophysis separated into two facets (43[1]). In summary, Wonambi is here shown to be a macrostomatan snake, as are the other fossil snake taxa Haasiophis, Pachyrhachis, and Podophis. As found previously (Tchernov et al., 2000), the potential exists for these snakes to nest within Macrostomata, i.e., above Xenopeltis and Loxocemus. This refutes Scanlon and Lees (2000) hypothesis of the phylogenetic relationships of Wonambi, as well as of the other fossil snakes with relatively well-developed hind limbs. While this result was already obtained by the revision (Rieppel and Zaher, 2000a) of the data set used by Lee (1998), we here add to the severity of the test of the data set presented by Scanlon and Lee (2000; for further comments see Rieppel and Kearney, 2002) by dening new characters. On the basis of our analysis, Wonambi furthermore adds no support to the conclusion reached by Scanlon and Lee (2000:420) that the snake body form arose in a large, non-fossorial ancestor. ACKNOWLEDGMENTS Our thanks go to Mark N. Hutchinson, who allowed us to investigate the South Australian Museum specimen of Wonambi. We are very grateful to John and Julie Barrie for their engagement in interesting discussions of the taphonomy of Wonambi and the fossils of the Henschkes Quarry Cave System in general. The following colleagues allowed the examination of specimens in their care: Darrell Frost, American Museum of Natural History, New York; Harold Voris and Alan Resetar, Field Museum of Natural History, Chicago; Guiseppe Puorto, Instituto Butantan, Sao Paulo; Douglas A. Rossman, Museum of Natural Science, Louisiana State University, Baton Rouge; and Alain Dubois, Museum National dHistoire Naturelle, Paris. We thank David Cundall and Garth Underwood for their reviews of this paper. Part of this paper was written at the Cladistics Institute, Harbor Springs, Michigan. LITERATURE CITED
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APPENDIX 1
Comparative material used in this study included the following specimens: Acrochordus granulatus: FMNH 232788, 234129, 234135, 244290. Acrochordus javanicus: AMNH 140813; FMNH 11081. Anilius scytale: AMNH 85980; FMNH 11175, 35683, 35688; UMMZ 183253. Aspidites melanocephala: AMNH 69302. Boa constrictor:

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FMNH 22435, 22438; IB 10769. Boa madagascariensis: AMNH 60728; MNHN 1983.484. Bothrochilus boa: AMNH 44002. Candoia carinata: AMNH 76701. Casarea dussumieri: MCZ 49135. Charina bottae: LSUMZ 36901, 15361. Charina reinhardtii: AMNH 10092; LSUMZ 34130; MNHN 84. Charina trivirgata: LSUMZ 15373, 14088. Corallus caninus: AMNH 63587; IB 2201. Corallus cropanii: AMNH 92997. Cylindrophis ruffus: AMNH 85647; MNHN 1970.411; FMNH 13100, 131780, 179033; LSUMZ 14221, 14075. Epicrates cenchria: AMNH 75796; IB 14624. Epicrates striatus: AMNH 77633. Eryx johnii: AMNH 43341; LSUMZ 48221, 16823. Eryx conicus: AMNH 2259; LSUMZ 48005, 48050. Eryx jaculus: AMNH 38168. Eunectes deschauenseei: IB 14619. Eunectes murinus: AMNH 62560; IB 15204. Exiliboa placata: AMNH 102892; UMMZ 183573. Leiopython albertisii: AMNH 104062. Leptotyphlops dulcis: UMMZ 149633, 149634. Leptotyphlops emini: FMNH 56374. Liasis mackloti: AMNH 69334. Liotyphlops albirostris: UMMZ 173967. Liotyphlops beui: IB 25344, 42519. Loxocemus bicolor: AMNH 19393; LSUMZ 49634; UMMZ 128027, 149657. Morelia amethistina: AMNH 117804. Morelia spilota: AMNH 79043; MNHN 1991.4048. Python molurus: AMNH 36573; FMNH 223198. Python regius: AMNH 75263; LSUMZ 13983. Trachyboa boulengeri: UMMZ 183143. Trachyboa gularis: AMNH 28982. Tropidophis melanurus: UMMZ 149664. Tropidophis haetiana: LSUMZ 28022, 28023. Tropidophis pardalis: UMMZ 149665, 150788, 150789, 150790. Typhlops sp.: FMNH 98952. Typhlops diari: UMMZ 190756. Typhlops platycephalus: UMMZ 149635. Typhlops rostellatus: UMMZ 149636. Ungaliophis continentalis: LSUMZ 55454; UMMZ149666, 190698, 190699, 190700. Xenopeltis unicolor: MNHN 1991.4446, 1973.138; LSUMZ 14074, 16821; FMNH 11524; UMMZ 131952, 225466, 227055, 227056, 227057.

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APPENDIX 2
List of characters, and data matrix, used in the phylogenetic analysis. 1. Tooth implantation pleurodont (0), or alethinophidian (as dened by Zaher and Rieppel, 1999) (1). Mosasauroids are coded pleurodont rather than thecodont (Zaher and Rieppel, 1999). 2. Plicidentine absent (0), or present (1). 3. Enamel surface smooth (other than presence of an anterior and/or posterior cutting edge) (0), or distinctly ridged (striated) (1). The enamel surface is smooth in basal mosasauroids (Polcyn et al., 1999). 4. Maxillary and dentary teeth relatively short, conical, upright (0), robust and recurved (1), or elongate, needle-shaped, and distinctly recurved (2). 5. Maxilla ascending process present (0), or absent (1). 6. Anterior end of maxilla straight or only weakly bent inwards (0), or distinctly curved inward (1). 7. Small horizontal shelf on medial surface of anterior end of maxilla present (0), or absent (1). This character was offered by Scanlon and Lee (2000:character 14). 8. Posterior end of maxilla does not project beyond posterior margin of orbit (0), projects moderately beyond posterior margin of orbit (1), or projects distinctly beyond posterior margin of orbit with broad, at surface (2). Character state 2 is correlated with a broad, smooth and loose overlap of ectopterygoid and maxilla. Wonambi shows the appropriate morphology of the maxilla for character state 2, but the ectopterygoid is unknown. The ectopterygoid as reconstructed by Scanlon and Lee (2000:g. 1) for Wonambi might be referable to a scincid (Tiliqua), which are also known from these cave deposits (Molnar, 1991). Ungaliophis and Tropidophis were wrongly coded for this character by Tchernov et al. (2000). 9. Medial (palatine) process of maxilla located in front of orbit (0), or below orbit (1). 10. Anterior dentigerous process of palatine absent (0), or present (1). According to Caldwell (2000:188), this is a two part character, because The anterior process of the palatine must be present in order for it to be toothed. By comparison to Dinilysia (Estes et al., 1970), it is clear that the lateral (maxillary) process of the palatine of alethinophidians corresponds to the part of the palatine located lateral to the choanal groove and contacting the maxilla in Dinilysia. The medial (choanal) process of the alethinophidian palatine corresponds to the part of the palatine which carries the choanal groove and contacts the vomer in Dinilysia. The toothed anterior process of the alethinophidian palatine is therefore a neomorph, and as such a well-dened character. The presence of teeth is part of its description. 11. Medial (choanal) process of palatine forms extensive concave

surface dorsal to ductus nasopharyngeus (0), narrows abruptly to form curved nger-like process (1), or forms short horizontal lamina that does not reach vomer (2). 12. Pterygoid contacts palatine in more or less complex pattern with clasping projections (0), in tongue-in-groove joint (1), or palatine-pterygoid contact reduced to at-overlap (2), 13. Pterygoid teeth absent (0), or present (1). Although the pterygoid as reconstructed by Scanlon and Lee (2000:g. 1) for Wonambi naracoortensis is very macrostomatan-like, it is not part of the material available to us. 14. Parietal without (0), or with (1) lateral wings meeting postorbital bones (where present). We here code Dinilysia and Pachyrhachis for the presence of such lateral parietal wings (contra Tchernov et al., 2000). The coding for Dinilysia is based on the gures in Estes et al. (1970). The coding for Pachyrhachis takes into account the general pythonomorph morphology of the parietal as exposed in dorsal view (Rieppel and Zaher, 2000a:g. 15A, B), even though the contact of the parietal and postorbital is not exposed in ventral view. 15. Parietal without (0), or with (1) distinct lateral ridge extending posteriorly from anterior lateral wing all along laterally descending parietal ange up to prootic. Many basal snakes show a more or less well developed ridge on the lateral surface of the anterior part of the descending ange of the parietal behind the orbit. We code the derived condition only for those taxa where a distinctly developed lateral ridge runs all along the descending ange of the parietal up to the prootic. 16. Parietal without sagittal crest (0), with low sagittal crest in its posterior part that merges into weakly expressed, anteriorly diverging ridges at about middle of parietal (1), or with prominent sagittal crest that merges into anterolaterally curving ridges at anterior end of parietal (2) (in adult). 17. Vidian canal does not (0), or does (1) open intracranially. This character refers to a condition where the Vidian canal forms a groove on the dorsal (intracranial) surface of the basisphenoid. 18. Anterior opening of Vidian canal (secondary anterior opening sensu Rieppel, 1979a, where present) single (0), or divided (1). We coded Dinilysia unknown (?) for this character. Estes et al. (1970) show a single anterior opening of the Vidian canal, above which they identify an unknown foramen. 19. Sella turcica located far in front of posterior end of basisphenoid (0), or close to posterior end of basisphenoid (1). 20. Sella turcica bordered posteriorly by well developed dorsum sellae (0), dorsum sellae low (1), or dorsum sella not developed, sella turcica with shallow posterior margin (2). 21. Lateral wings of basisphenoid (sensu McDowell, 1967) absent (0), or present (1). 22. Ventral surface of basisphenoid smooth (0), with weakly developed sagittal crest from which protractor pterygoidei originates (1), or with strongly projecting sagittal crest (2) (in the adult). This character denition represents a ner subdivision of character 77 in Tchernov et al. (2000) 23. Basisphenoidbasioccipital suture smooth (0), or transversely crested (1). 24. Basipterygoid processes present (0), or absent (1). We do not consider the basipterygoid processes of nonophidian squamates and macrostomatans to be homologous based on developmental evidence (Kluge, 1991; Tchernov et al., 2000). Because development cannot be assessed for fossils, and because the orientation of the basipterygoid processes in Wonambi is longitudinal rather than transverse as in Python, we here code the presence of basipterygoid processes in Dinilysia, Haasiophis, and Wonambi as comparable to outgroups as well as to macrostomatans in order to one more time test their homology by congruence. However, as also argued by Lee and Caldwell (2000) in their revision of their data matrix, we refrain from using details of shape and orientation of the basipterygoid processes as separate characters, as these vary widely among squamates. Although the basipterygoidpterygoid contact in Loxocemus is more highly differentiated than in most other basal macrostomatans included in this analysis, we do recognize that the basipterygoid processes are less well dened than they are in Boa, Python, and Wonambi, and hence code Loxocemus as lacking basipterygoid processes for this analysis (see comments on the putative homology of basipterygoid processes in snakes above). 25. Cristae trabeculares short and/or indistinct (0), or elongate and distinct in lateral view of the basisphenoidal rostrum (1). 26. Parabasisphenoidal rostrum narrow (0), or broad (1). Dibamus

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cess, which is distinct in Pachyrhachis. Here we code it for its distinctiveness from the exoccipital. 34. Juxtastapedial recess, dened by crista circumfenestralis, absent (0), present but open posteriorly (1), or present and closed posteriorly (2). We here follow Rage (1984) and code Dinilysia for a posteriorly open juxtastapedial recess. On the basis of recent investigations of embryonic skulls of Acrochordus, we here code this character as inapplicable for that taxon (contra Rieppel, 1980). 35. Crista circumfenestralis exposes most of stapedial footplate (0), or converges upon stapedial footplate (1). 36. Crista interfenestralis does not (0), or does (1) form individualized component in ventral rim of crista circumfenestralis. 37. Jugular foramen exposed in lateral view (0), or concealed in lateral view by crista tuberalis (1). 38. Posterior dentigerous process of dentary absent (0), short (1), or long (2). 39. Medial margin of the adductor fossa is relatively low and smoothly rounded (0), or forms a distinct, dorsally projecting crest (1). The present character delimitation reverts the polarity of character 88 as used by Tchernov et al. (2000). 40. Coronoid process formed by surangular, or surangular portion of compound bone (with or without participation of the coronoid bone) absent (0), or present (1). 41. Chevrons present (0), or absent (1). 42. Hemapophyses absent (0), or present (1). 43. Para-diapophysis conuent (0), or separated into dorsal and ventral facet (1). 44. Prezygapophyseal processes absent (0), or present (1). 45. Subcentral foramina absent (0), present, consistently small (1), or present, of variable size (2).

has no ossied parabasisphenoidal rostrum (Rieppel, 1984); amphisbaenians have a short parabasisphenoidal rostrum, which is broad posteriorly, but rapidly tapers to a pointed tip lined laterally by the orbitosphenoids (Zangerl, 1944). 27. Prootic does not (0), or does (1) exclude parietal from (anterior) trigeminal foramen. 28. Laterosphenoid absent (0), present (1). The presence of a laterosphenoid is correlated with the presence of a trigeminofacialis chamber containing the Gasserian ganglion (absent in scolecophidians). 29. Posteriorly undercut prootic ledge underlapping posterior trigeminal foramen absent (0), or present (1). 30. Anterior end of supratemporal (facet) located behind or above posterior trigeminal (V3) foramen (0), or in front of posterior trigeminal (V3) foramen (1). This character delimitation differs from that of Tchernov et al. (2000:character 40) in an attempt to better capture the variation observed among the terminal taxa. Pachyrhachis is coded (?), because we believe that the supratemporals are dislocated anteriorly, while the supratemporal facet is indistinct. 31. Supratemporal facet on opisthotic-exoccipital at (0), or sculptured and delineated with transversely oriented, dorsally projecting posterior rim that overhangs exoccipital (1). 32. Free-ending posterior process of supratemporal absent (0), or present (1). We assume the presence of a free-ending process of the supratemporal for Wonambi on the basis of the proportion of the (preserved) mandibular ramus relative to the (preserved parts of the) skull (Barrie, 1990; Scanlon and Lee, 2000:g. 1). 33. Paroccipital process of opisthotic underlying supratemporal present (0), or reduced to short projection which is separated from exoccipital by deep notch (1), absent (2). Tchernov et al. (2000) coded this character mainly for the relative size of the opisthotic paroccipital pro-

APPENDIX 3
Data matrix for the analysis of the phylogenetic relationships of Wonambi. 1 Varanus Lanthanotus Mosasauroidea Dibamidae Amphisbaenia Pachyrhachis Hassiophis Dinilysia Wonambi Scolecophidia Anilioidea Xenopeltis Loxocemus Boa Python Trachyboa Exiliboa Ungaliophis Tropidophis Acrochordus Podophis 0 0 0 0 0 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 2 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 3 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 ? 4 0 0 0 0 0 2 2 1 2 0 1 2 2 2 2 2 2 2 2 2 ? 5 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 ? 6 0 0 0 0 0 0 0 ? 0 0 1 1 1 0 0 1 0 0 0 1 0 7 0 0 0 1 ? ? 1 1 ? 1 ? 0 0 0 1 1 1 1 1 1 1 ? 8 0 0 0 0 0 2 2 1 2 0 1 1 2 2 2 2 2 2 0 0 2 ? 9 0 0 0 ? ? ? 1 0 1 ? 0 0 0 1 1 1 1 1 1 1 ? 10 0 0 0 0 0 1 1 0 1 0 1 1 1 1 1 1 1 1 1 1 1 11 0 0 ? ? ? ? 2 0 0 0 1 0 1 1 0 1 2 1 1 1 0 0 2 12 0 0 0 0 0 1 1 0 1 2 1 1 1 1 1 1 1 1 1 1 1 13 0 1 1 0 0 1 1 1 ? 0 1 1 1 1 1 1 1 1 1 1 1 14 0 0 0 0 0 1 0 1 1 0 0 0 0 1 1 1 1 1 1 1 ? 15 0 0 0 0 0 ? ? 1 1 0 0 0 0 1 0 0 0 0 0 1 ? 16 0 0 0 0 0 2 1 2 2 0 1 1 1 2 2 1 1 1 1 0 ? 17 0 0 0 1 0 ? ? ? 0 ? 1 1 1 0 0 1 1 1 1 1 ? 18 0 0 0 0 0 ? ? ? 1 0 0 0 0 1 0 0 0 0 0 0 ? 19 0 0 0 0 0 ? ? ? 0 1 1 0 0 0 0 ? ? 0 0 0 ? 20 0 0 0 2 2 ? ? ? 2 1 2 1 1 1 1 1 ? ? 1 1 1 ? 21 0 0 0 0 0 ? ? 0 1 0 1 1 1 1 1 1 1 1 1 1 ? 22 0 0 0 0 0 ? 1 0 2 0 0 1 1 2 2 1 0 0 1 1 ? 23 0 0 0 0 0 ? 0 0 1 0 0 0 0 1 1 0 0 0 0 0 ? 24 0 0 0 0 0 ? 0 0 0 1 1 1 1 0 0 1 1 1 1 1 ? 25 0 0 0 0 0 ? ? ? 1 0 1 1 1 1 1 1 1 1 1 1 ?

RIEPPEL ET AL.RELATIONSHIPS OF SNAKE WONAMBI

APPENDIX 3. Continued.
26 Varanus Lanthanotus Mosasauroidea Dibamidae Amphisbaenia Pachyrhachis Haysiophis Dinilysia Wonambi Scolecophidia Anilioidea Xenopeltis Loxocemus Boa Python Trachyboa Exiliboa Ungaliophis Tropidophis Acrochordus Podophis 0 0 0 ? 1 0 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 ? 27 0 0 0 0 0 ? ? 0 1 0 0 1 0 0 1 1 1 1 0 0 1 1 ? 28 0 0 0 0 0 ? 1 0 1 0 1 1 1 1 1 1 1 1 1 1 ? 29 0 0 0 0 0 ? 1 0 1 0 0 0 0 1 0 0 0 0 0 0 ? ? ? 0 0 1 ? 0 0 1 1 1 1 0 0 1 1 ? 30 0 0 0 0 31 0 0 0 0 ? 0 0 0 1 0 0 0 0 1 1 1 0 0 1 0 ? 32 0 0 0 0 ? 1 1 0 1 0 0 1 1 1 1 1 1 1 1 1 1 33 0 0 0 2 2 1 ? 2 1 2 2 2 2 1 1 1 2 1 1 2 ? 34 0 0 0 0 0 ? ? 1 2 1 1 1 1 2 2 2 2 2 2 ? ? 35 ? ? ? ? ? ? ? 0 0 0 1 0 0 0 1 1 0 0 0 0 ? ? 36 0 0 0 0 0 ? ? 0 1 0 0 1 1 1 1 1 1 1 1 0 ? 37 ? ? 0 0 0 ? ? 0 1 0 0 0 0 1 1 1 0 0 1 0 ? 38 0 0 0 0 0 2 2 1 2 0 1 2 2 2 2 2 2 2 2 2 ? 39 0 0 0 0 0 1 1 0 0 ? 0 1 0 0 1 1 1 1 1 1 0 ? 40 0 0 0 0 1 0 0 ? 1 0 1 1 1 1 1 1 1 1 1 1 ? 41 0 0 0 0 0 ? 1 ? 0 1 1 1 1 1 1 1 1 1 1 1 0 42 0 0 0 0 1 0 1 ? 1 1 0 0 0 1 1 1 1 1 1 1 1 1 1 ? 43 0 0 0 0 0 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 ? 44 0 0 0 0 0 0 0 1 0 1 1 1 1 1 1 1 1 1 1 1 ?

829

45 0 0 0 1 1 0 1 1 1 0 1 1 2 0 1 1 1 1 1 1 1 1 1 1 ?