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The Skull of the Rare Malaysian Snake Anomochilus Leonardi Smith, Based Blackwell Publishing LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin- nean Society of London, 2007? 2007 1494 671685 Original Articles SKULL OF THE MALAYSIAN SNAKE ANOMOCHILUS LEONARDIO. RIEPPEL and J. A. MAISANO Zoological Journal of the Linnean Society, 2007, 149, 671–685. With 5 figures The skull of the rare Malaysian snake Anomochilus leonardi Smith, based on high-resolution X-ray computed tomography Downloaded from https://academic.oup.com/zoolinnean/article/149/4/671/2630933 by guest on 30 September 2021 O. RIEPPEL1* and J. A. MAISANO2 1Department of Geology, The Field Museum, 1400 S Lake Shore Drive, Chicago, Illinois 60605-2496, USA 2Jackson School of Geosciences, The University of Texas at Austin, Austin, Texas 78712, USA Received January 2006; accepted for publication June 2006 The skull of the rare Malaysian snake Anomochilus leonardi is described in detail on the basis of a high-resolution X-ray computed tomographic scan of a mature specimen. Its skull anatomy is compared with that of Anomochilus weberi, as well as with that of scolecophidians and basal alethinophidians such as Anilius, Cylindrophis, uropeltines and selected Booidea. Anomochilus leonardi is found to be more paedomorphic than Anomochilus weberi. The genus Anomochilus most closely resembles uropeltines in skull anatomy. Both Anomochilus and uropeltines develop a ‘central rod design’ of skull morphology, which requires the presence of medial frontal pillars, in adaptation to bur- rowing habits. These pillars are an alethinophidian characteristic, absent in the skull of scolecophidians, which develop an ‘outer shell design’ in adaptation to burrowing. These results are discussed in the light of the hypothesis that scolecophidians and basal (i.e. non-macrostomatan) alethinophidians are ‘regressed macrostomatans’. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 671–685. ADDITIONAL KEYWORDS: anatomy – paedomorphosis – phylogeny. INTRODUCTION East have ignited a vigorous debate on snake interre- lationships (e.g. Coates & Ruta, 2000; Rieppel & The snake genus Anomochilus comprises two species. Kearney, 2001; for a full discussion of the debate and Anomochilus weberi (Lidth de Jeude, 1890) is known references see Rieppel et al., 2003). Some analyses from three specimens collected in Sumatra and (e.g. Lee & Caldwell, 1998; Scanlon & Lee, 2000; Lee, Borneo (Lidth de Jeude, 1890, 1922; Brongersma & 2005; Scanlon, 2005) challenge the traditional view Helle, 1951), and A. leonardi Smith (1940) is known that the burrowing or fossorial (secretive) scolecophid- from five specimens collected in Malaysia (Lim & ians (blindsnakes and threadsnakes) and ‘anilioids’ Mohd. Sharef, 1975; Stuebing & Goh, 1993; Yaakob, (pipesnakes and shieldtails) are basal to macrostoma- 2003). Due to the scarcity of these Indonesian snakes, tan snakes (boas, pythons and colubroids). They do so little is known about their anatomy and biology. How- by placing the fossil snakes with well-developed hind ever, in light of the genus’ putative relationship to limbs at the bottom of the snake tree. But because other basal snakes such as scolecophidians or ‘anili- these fossils combine well-developed hind limbs with a oids’ (Lidth de Jeude, 1890; Boulenger, 1893; Under- macrostomatan skull structure (e.g. Zaher, 1998; Rage wood, 1967; McDowell, 1975, 1987; Rieppel, 1977, & Escuillié, 2000, 2003; Polcyn, Jacobs & Haber, 1979a; Groombridge, 1979a), Anomochilus has long 2005), their placement basal to all extant snakes been an important taxon in the analysis of snake renders scolecophidians and ‘anilioids’ ‘regressed mac- phylogeny. rostomatans’ (Rage & Escuillié, 2000, 2003; G. Under- More recently, fossil snakes with well-developed wood, pers. comm.). These recent controversies hind limbs from the mid-Cretaceous of the Middle highlight the importance of an improved understand- ing of the phylogenetic interrelationships of basal *Corresponding author. E-mail: orieppel@fieldmuseum.org snakes, including Anomochilus. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 671–685 671 672 O. RIEPPEL and J. A. MAISANO In two landmark papers, Cundall and collaborators The specimen was scanned using the following (Cundall & Rossman, 1993; Cundall, Wallach & parameters. A FeinFocus microfocal X-ray source oper- Rossman, 1993) presented a detailed description of ating at 180 kV and 0.088 mA with no X-ray prefilter the cranial anatomy of Anomochilus weberi, which was employed. An empty container wedge was used. formed the basis of a cladistic analysis of the phyloge- Slice thickness corresponded to two lines in a CCD netic relationships of the species. Anomochilus was image intensifier imaging system, with a source-to- found not to be a scolecophidian, as indicated by vis- object distance of 21 mm. For each slice, 1600 views ceral and integumental features (V. Wallach, cited in were taken with four samples per view. The field of Cundall & Rossman, 1993: 236), but instead to be a image reconstruction was 5.5 mm, and an image non-macrostomatan alethinophidian snake. Its inclu- reconstruction offset of 10 000 was used with a recon- sion in that analysis resulted in rejection of the puta- struction scale of 1500. Downloaded from https://academic.oup.com/zoolinnean/article/149/4/671/2630933 by guest on 30 September 2021 tive monophyly of Anilioidea (Rieppel, 1977, 1979a), The data set consists of 553 HRXCT slices taken however, as Anomochilus was found to be the sister- along the transverse (= coronal) axis of the skull from taxon of all other alethinophidian snakes. This is an the tip of the snout to the occiput. The scan axis was important result, as it supports the hypothesis that offset from orthogonal approximately 9° on the trans- ‘Anomochilus might be an intermediate between verse axis and 10° on the frontal axis due to a fixed tilt living scolecophidian and alethinophidian snakes’ of the specimen's head relative to its body. Each slice (Cundall & Rossman, 1993: 236). If correct, this image was gathered at 1024 × 1024-pixel resolution, hypothesis has major implications either for under- resulting in an in-plane resolution of 5.4 µm per pixel. standing the interrelationships of basal snakes, or for Each slice represents a thickness of 14 µm, with understanding the ‘regression’ of the macrostomatan 14-µm interslice spacing. skull structure. The data set was rendered in three dimensions The interrelationships of Anomochilus can be fur- using VGStudio MAX 1.2 (Volume Graphics, Heidel- ther clarified by the inclusion of the second species, berg, Germany). The data volume was rotated to posi- A. leonardi. In order to prepare the ground for such tion the skull orthogonally, then three-dimensional future analysis we describe and illustrate the skull of (3D) cutaway views were generated along its true this species, and compare and contrast its anatomy orthogonal axes. Noise in the HRXCT data set, due to with that of A. weberi, scolecophidians, Anilius, the very small size of the specimen, causes the 3D ren- cylindrophines and uropeltines, as well as some basal derings to appear grainy. macrostomatans. The slices spanned by each element in the original This description of the skull of Anomochilus HRXCT data set are noted at the beginning of the leonardi is based on a high-resolution X-ray computed description of that element, and selected labelled tomographic (HRXCT) scan of a mature specimen HRXCT cutaway views (Figs 3–5) illustrate the inter- (FRIM 0026, Forest Research Institute Malaysia). nal anatomy of the skull in transverse (Tra), frontal HRXCT permits the non-destructive visualization of (Fro) and sagittal (Sag) planes. Original data set slice hard tissues in preserved specimens, and is particu- numbers are referenced throughout the description. larly useful when dealing with small, rare taxa such An interactive, web-deliverable version of the HRXCT as Anomochilus. data set, as well as animations of 3D reconstructions, can be viewed at http://www.digimorph.org/specimens/ Anomochilus_leonardi, and the original full resolution MATERIAL AND METHODS HRXCT data are available from the authors. The head of a whole preserved specimen (FRIM 0026) At certain points in the description below, the Ano- was scanned at the High-Resolution X-ray CT Facility mochilus leonardi data set is compared with HRXCT at The University of Texas at Austin. The specimen data sets for Typhlops jamaicensis (Typhlopidae; was collected on the campus of the Forest Research USNM 12378, National Museum of Natural History), Institute Malaysia, Kepong, Selangor State, Malaysia. Leptotyphlops dulcis (Leptotyphlopidae; TNHC 60638, The skull measures 6.8 mm in length and 2.9 mm Texas Natural History Collection, Texas Memorial across the otic capsules. The snout–vent length of the Museum), Anilius scytale (Aniliidae; USNM 204078), specimen is 251 mm and its total length is 257 mm Loxocemus bicolor (Loxocemidae; FMNH 104800, (Yaakob, 2003). The scanned specimen is thus compa- Field Museum of Natural History), Xenopeltis unicolor rable in size (length) with the other specimens from (Xenopeltidae; FMNH 148900), Python molurus Peninsular Malaysia with total lengths ranging from (Pythonidae; TNHC 62769), Aspidites melanocephalus 220 mm (Lim & Mohd. Sharef, 1975) to 228 mm and (Pythonidae; FMNH 97055), Casarea dussumieri 227 mm, respectively (Stuebing & Goh, 1993), but is (Bolyeriidae; UMMZ 190285, University of Michigan smaller than the Sabah specimen
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