HERPETOLOGICAL JOURNAL, Vol. 13, pp. 1-20 (2003)

THE AUSTRALIAN ELAPID GENUS CA COPHIS: MORPHOLOGY AND PHYLOGENY OF RAINFOREST CROWNED SNAKES

JOHN D. SCANLON

Department of Environmental Biology, University of Adelaide, Australia and

Department of Palaeontology, South Australian Museum, No rth Terrace, Adelaide, SA 5000, Australia

The genus Cacophis, comprising four species endemic to eastern Australia, is uniquely derived among terrestrial Australasian elapid snakes in the temporal scale pattern, presence of a relatively high and narrow dorsal crest ('choanal process') on the palatine bone, and presence of keeled supra-anal scales in adult males. Recent analyses based on morphology and genetics do not completely resolve relationships among Australasian elapids, but support relationships of Cacophis with the (Furino, Glyphodon) and (Aspidomorp hus, Demansia) clades, which are adopted here as outgroups for intrageneric analysis. Within Cacophis, morphoclines in size, head scalation, tooth numbers and colour patternsindic ate that C. squamulosus is the sister-group to the remaining three species; among the latter, there is conflictingevidence for both (harriettae, krejftii) and (churchi//i, krejftii) clades, but the latter alternative has greater support. Revised diagnoses are given for the genus and included clades, and a simple phylogeographic model proposed.

Key words: Hydrophiinae, morphology, skull, head scales, colour patterns,behaviour, phylogeography

INTRODUCTION level analysis (cf. Hutchinson, 1990). Such a 'global' analysis will not be presented here, as I concentrate on a Numerous studies have contributed to understanding particular genus and its putative close relatives. the phylogeny of Australian elapid snakes, providing Hutchinson (1990) considered diagnosis of the genus evidence for monophyly of a number of genera and of Cacophis problematic, and recognized it only 'tenta­ several suprageneric units (e.g. Schwaner et al. , 1985; tively' as distinct from Purina. Cacophis consists of Mengden, 1985; Shine, 1985; Hutchinson, 1990; Greer, four species of small nocturnal saurophagous (lizard­ 1997; Keogh, 1998, 1999; Keogh et al., 1998, 2000). eating) snakes, all restricted to rainforest or wet These probable clades include: the 'subfamily' sclerophyll habitats in coastal regions of eastern Aus­ Hydrophiinae comprising all terrestrial Australasian tralia (Queensland and New South Wales). Three of the elapids as well as marine forms (in either the sense of species have long been recognized, although they were McDowell, 1987, or that of Slowinski & Keogh, 2000, previously referred to as many as three separate genera which differ in whether Laticauda is included); the vi­ (Cacophis, Aspidomorphus and Glyp hodon in Worrell, viparous radiation (Shine, 1985); the true sea snakes 1963 ). McDowell ( 1967) suggested that these three spe­ (here regarded as a monophyletic 'tribe' Hydrophiini, cies (krejftii, Dwarf crowned snake· harriettae despite a recent analysis suggesting diphyly; White-crowned snake; and squamulo;us, Golden� Rasmussen, 2002); and a 'No techis lineage' comprising crowned snake) formed a single natural group distinct chromosome groups 4, 5 and 10 of Mengden ( 1985). from other genera; Cogger (1975) brought them to­ However, resolution remains poor because characters gether in Cacophis, and full synonymies are given in have often been inadequately defined or polarized, or Cogger, Cameron & Cogger (1983). insufficiently numerous to resolve the large number of The fourth species, found in the Wet Tropics of species (e.g. McDowell, 1967; Storr, 1985; Wallach, northernQueensland, was first recognized informally as 1985; Greer, 1997; Lee, 1997). I have studied external 'Glyphodon sp.' by Worrell (1963: 125 and plate 56), and skeletal morphology in the terrestrial Australasian and subsequently as Cacophis h. [harriettae]flavicollis elapid snakes, ·attempting to define and test additional [nomen nudum] (McDowell, 1967: 536) and Cacophis characters in order to improve phylogenetic resolution sp. (Wilson & Knowles, 1988: 332; Gow, 1989: 84; (e.g. Scanlon, 1985) and as a basis for interpretation of Ehmann, 1992: 392). Cogger (in Cogger et al., 1983: Miocene fossils (Scanlon, 1996). Animportant interme­ 219) includes the mentions by Worrell ( 1963) and diate goal is to establish the monophyly and internal McDowell ( 1967) in the synonymy of harriettae, but relationships of groups of species (e.g. genera) which C. notes that both refer to what is probably a distinct spe­ can conveniently be used as discrete units in a higher- cies. The name Cacophis churchilli Wells & Wellington, 1985 is available for this form, though it has Correspondence: J. D. Scanlon, Department of only recently come into wider use (Greer, 1997: 160, Palaeontology, South Australian Museum, North Terrace, Adelaide, SA 5000, Australia. 178; Shea & Sadlier, 1999; Queensland Museum, 2000: E-mail: Scanlon [email protected] 239; Cogger 2000: 771). Ehmann (1992) calls it the 2 J. D. SCANLON

'NorthernDwarf Crowned Snake', but 'dwarf' is not es­ (Swofford, 2002), in some cases using batch commands pecially appropriate since it attains body sizes similar to generated using TreeRot version 2 (Sorenson, 1999). C. harriettae (see below). While detailed studies of geographic and genetic OUTGROUP RELATIONSHIPS OF CA COPHIS variation remain to be done (J. Sumner in prep.), I re­ In order to assess the polarity of morphological char­ gard the the identity and boundaries of these species as acters contributing to the diagnosis of Ca cophis and now being stable, and a formal revision is not given resolution of relationships among the included species, here. Rather, this paper reviews evidence for relation­ relevant outgroups must be identified. Ideally, these ships between Ca cophis and other genera, reports should include the two clades most closely related to the observations of some unusual morphological features ingroup to allow the outgroup comparison procedure of contributingto the diagnosis ofthe genus, and uses read­ Maddison et al. ( 1984) . Previous analyses of Australa­ ily available data to derive an explicit phylogenetic sian elapid relationships support the basal position of hypothesis for the four included species. Laticauda and the Solomon Island genera (McDowell, One motive for investigation of this genus is the dis­ 1970; Keogh et al., 1998), and the monophyly ofa large covery of fossil material of small elapid snakes fromthe viviparous lineage which includes mainly Australian Miocene ofnortbernAustral ia, including a maxilla with terrestrial elapids and hydrophiine sea snakes (Shine, fe atures resembling those of Cacophis species (Scanlon, 1985; Keogh et al., 1998, 2000). These results imply 1995, 1996). However, as variation in skeletal features that the remaining Australo-Papuan oviparous genera (apart from tooth counts) within the genus is dominated form either one or several clades along the stem lineage by ontogenetic change in proportions (pers. obs.), the of the viviparous group. This intervening part of the tree emphasis here is on externalmor phology. (including Ca cophis) has been poorly resolved by prior work, which is attributable mainly to insufficient sam­ METHODS pling of characters and (especially Melanesian) taxa, but All data are drawn either frompublished sources or perhaps also to the rapidity of the adaptive ra diation ( cf. examination of specimens - including those in the col­ Schwaner et al., 1985; Wallach, 1985; Mengden, 1985; lections of the Australian Museum, Sydney (AMS); Lee, 1997; Greer, 1997). The selection of outgroups Queensland Museum, Brisbane (QM); South Australian must therefore be provisional at this stage. Museum, Adelaide (SAM); and Western Australian Classifications up to that of Worrell ( 1963, 1970) re­ Museum, Perth (W AM). Some additional specimens at ferred at least some Cacophis species to Aspidomorphus the American Museum of Natural History, New York (see Mengden, 1983, for review), but it has since been (AMNH) and Museum of Comparative Zoology, considered that Aspidomorphus is closest to Demansia Harvard (MCZ) were examined on my behalfby M. Lee. (McDowell, 1967; Keogh et al., 1998). Also, a consen­ A large number of externaland skeletal morphologi­ sus has developed that Ca cophis is closely related to cal features havebeen investigated fortheir potential to Purina and Glyp hodon (McDowell, 1967; Wallach, contribute phylogenetic information fo r Australian 1985; Hutchinson, 1990; Greer, 1997; Keogh et al, elapids, and many of them show overlapping variation 1998; Keogh, 1999). A recent analysis of DNA se­ of continuous or discrete characters across species or quence data (Keogh et al., 1998) has found support for a more inclusive groups (Wallach, 1985; Scanlon, 1985; clade compnsmg Cacophis, Demansia, Lee, 1997). This is consistent with the uncontroversial Aspidomorphus, Purina and Glyphodon. While the de­ hypotheses that novel characters (genetic, morphologi­ tailed results varied with different methods of data cal, or behavioural) must pass through a stage of analysis, they 'consistently grouped these four genera in polymorphic coexistence with their alternative, various combinations' (p. 77), with Demansia and plesiomorphic states before being fixed in one or more Aspidomorphus most strongly linked. As shown in Fig. descendant populations, and that such polymorphisms may be retained for evolutionarily significant periods. Cacop his In many cases, I recognize polymorphic coexistence of Glyphodon alternate conditions as a single, separate, intermediate Furina state, and such characters are treated as ordered morphoclines. In other instances, polymorphic taxa are Neelaps assigned to several states on the basis of relative fre­ ---c Simose/aps quency of alternate conditions observed in samples. This approach to definition and ordering of character Aspidomorphus states corresponds to a simplified version of the 'fre­ quency bins' method, and has been shown to perform Demansia well in simulations (Wiens, 1998). Data matricesand constraint treeswere edited using FIG. I. Relationships assumed between Cacophis and other genera used in outgroup comparisons. The two outgroups MacClade version 4.0 for Power PC (Maddison & Aspidomorphus + Demansia, and Glyphodon + Furina, Maddison, 2000) and phylogenetic analyses carried out contribute equally to the estimation of plesiomorphic on a Macintosh G4 using PAUP* version 4.0b lO characte states for Cacophis. CA COPHIS MORPHOLOGY AND PHYLOGENY 3

I, the two outgroups to Ca cophis adopted in this paper strongly pigmented; scales less glossy; eyes pale; pos­ are (Furina, Glyphodon) and (Aspidomorphus, torbital lacks anteroposterior kinesis; no contact of Demansia), which is thus consistent with Keogh et al. preocular and frontal scales; occipital and dorsal ground (I 998) and most previous classifications and analyses. colour similar; pale spots on dorsal scales single-col­ oured), and can thus be excluded from the( Glyphodon FUR/NA AND GLYPHODON (Furina (Neelaps, Simoselaps))) clade. Thus Glyphodon and Furina, either alone or together McDowell (1967) and Wallach (1985) concluded withSimoselaps andNeelaps (Scanlon, 1985, 1988: Fig. from morphological analyses that Cacophis was most I), form a close outgroup to Cacophis (Hutchinson, closely related to Furina and Glyphodon. The latt�r gen­ 1990; Keogh, 1999). Either way, the species of era were synonymized as Furina by Hutchinson (1990) Glyp hodon and Furina are the most appropriate taxa to and, as noted above, regarded only tentatively as distinct estimate the ancestralstates of this outgroup clade. from Ca cophis. Keogh (1999) found strong similarities among the hemipenes of Cacophis and Furina (sensu DEMANSIA AND ASPIDOMORPHUS lato), including both in his 'Group 4' which he regards as a monophyletic group. These two genera, suggested by McDowell (1967) to Synonymy of Glyphodon and Furina has been be closely related, share several probable adopted by Hutchinson ( 1990) and others, in part to deal synapomorphies, including uniquely derived features of wi th the apparent problem of classifying Glyphodon the maxilla: tooth numbers are the highest of any elapids barnardi Kinghorn, I 939. Cogger's (1975) key to gen­ (Bogert, 1943; McDowell, 1967), the medial era purports to distinguish the genera on the criterion of (ectopterygoid) process is elongate, and in most species 'nasal undivided' (Furina) vs. 'nasal divided' the suborbital region is dorsoventrally extremelythin (Glyphodon), but in fact G. barnardihas the nasal undi­ and (in fresh or wet-preserved specimens) flexible (pers. vided and would be assigned to Furina by this criterion. ohs.). Mengden ( 1985) found that Demansia 'possesses Polarity of this character is ambiguous since both states a unique karyomorph not easily associated wi th any occur in related genera (see below), but other cranial other Australian elapid', while the karyotype of and external morphological evidence suggests that G. Aspidomorphus has not been reported. The relatively tristis and G. dunma/Ii are sister taxa (e.g. in both spe­ high genetic distances found by Cadle & Gonnan (198 I) cies the parasphenoid is excluded from the optic and Schwaner et al. (1985) between Demansia and all fenestra, an uncommon derived character not observed other genera - including Aspidomorphus - conflictwith in G. barnardior other Furina spp.; pers. ohs.), while G. evidence from morphology (the skeletal characters just barnardi, Furina diadema and F. ornata are more given, and others in McDowell, I 967) and DNA se­ closely related to the fossorial radiation of Neelaps and quences (Keogh et al., 1998), but can be explained by, Simoselaps (Scanlon, I 985, I 988, unpublished data). I for example, accelerated genetic change therefore retain Glyphodon as a distinct genus for G. (autapomorphy) in Demansia, as already suggested by tristis and G. dunmalli, and refer G. barnardito Furina. Cadle & Gorman (1981). I provisionally recognize a ( Glyphodon (Furina Demansia and Aspidomorphus, provisionally ac­ (Neelaps, Simoselaps))) clade which can be diagnosed cepted as sister taxa forming a single clade, are used as as follows: nasal and second supralabial separated from one outgroup in comparisons below. Skulls of all three the preocular (reversing twice in fossorial lineages); species ofAspid omorphus have been examined (see also ventral surface white; dorsal scales highly glossed; eyes McDowell, 1967), and a preliminary analysis suggests dark (Scanlon, 1985; Hutchinson, 1990; characters dis­ that A. schlegelii is basally related to the other two spe­ cussed below). Glyphodon spp. lack additional derived cies, although A. muelleri is the least derived in states shared by Furina with Simoselaps and Neelaps morphology (unpublished data). The larger number of spp.: postorbital bones with kinetic attachment to pari­ species (approximately 15 [Shea & Scanlon, unpub­ etal (involved in mechanism for maxillary erection and lished data]) and morphological diversity wi thin retraction; McDowell, I 969a; Scanlon, I 985); frontal Demansia present a greater problem, but it seems likely may contact preocular scales (rare to common variant, from external characters (e.g. number of ventral scales, [Storr, I 968, 198 I], never observed in Glyp hodon or occurrence of posterior scale-row reduction) that D. any other elapid genera, pers. ohs.); black head and nape simplex is basally related to all the other, larger and blotches contrasting with the dorsal ground colour and more elongate species (see Table 1 and characters dis­ separated by a distinct pale spot or bar; and a reticulate cussed below). While cranial data have been obtained dorsal pattern where each scale may have a black edge, for only a few species of Demansia, D. simplex is also yellow basal spot and red intermediate zone (three dis­ plesiomorphic relative to the others examined (D. tinct pigments; [Storr, I 968]). Ca cophis spp. lack most psammophis, D. vestigiata and D. sp. cf. olivacea) in of these derived features and retain the alternate states having a relatively broader frontal, less constrictedpari­ common to most other Australasiantaxa (preocular con­ etal, and less developed 'interorbital septum' tacts second labial and frequently nasal; ventrals (Underwood, 1967) of the parasphenoid. 4 J. D. SCANLON

DISTINCTIVE FEATURES OF CA COPHIS sence in Cacophis but did not discuss their occurrence in A revised diagnosis of Cacophis is given in a later any other taxa. The foramina are present in nearly all section, in which several classes of characters are in­ outgroup skulls examined (but not in D. simplex, NTM cluded: unambiguous autapomorphies, characterizing Rl8625; one specimen of F. diadema, SAM R 6703), Ca cophis but absent or uncommon in the outgroups and and absent in nearly all Cacophis (Figs 2, 3c; present other Australasian elapids; possible apomorphies, con­ unilaterally in one specimen of C. kreffiii, SAM ditions with a more restricted distribution including one R26974, Fig. 3a). In some other taxa (e.g. Pseudechis spp.), the foramina may be obliterated during adult life or more of the outgroup genera; and likely by forward extension of median contact between the plesiomorphies, conditions which are shared widely mandibular adductor muscles forming a sagittal crest, among Australasianelapids but lost or modifiedin vari­ but in Ca cophis they are typically absent even when the ous lineages from which Cacophis can thereby be muscles are still widely separated. excluded. Contrary to Hutchinson ( 1990), Ca cophis can readily be diagnosed on the basis of autapomorphic A2. Choanal process of palatine bone. McDowell ( 1970, 1987) diagnosed the subfamily Hydrophiinae of states of external as well as cranial characters. 'palatine draggers' on the basis of the palatine 's clasp­

AUTAPOMORPHIES ing articulation with the pterygoid, and lack of choanal and perforate lateral processes. The dorsomedial edge A 1. Parietal fo ramina. Most Australasian elapids, of the palatine is smooth and nearly parallel to the tooth like many other colubroids, have a pair of small fo­ row (i.e. choanal process totally absent) in most ramina (or sometimes closed pits, not piercing the bone) hydrophiine taxa and their probable sister group near the centre of the dorsal surface of the parietal. laticauda (McDowell, 1970; pers. ohs.; State 0). How­ These are presumably not equivalent to the median pin­ ever, a number of species in the Australian radiation eal foramen which was lost in an ancestor of all snakes, have a low to moderate laminardorsal process similar to but there do not seem to have been any descriptions of the choanal process of such forms as Bungarus, but it is the detailed anatomy, function or phylogenetic value of usually nearly vertical rather than arching medially over the paired openings. Greer ( 1997: 178) noted their ab- the choana. Greer ( 1997) recognized this as diagnostic

TABLE I. Comparative morphological data for species of Cacophis and outgroup genera: maximum known snout-vent length (max. SYL); and observed ranges of the number of ventral and subcaudal scales; and number of alveoli fortooth attachment on the maxilla (exclud ing the two enlarged anterior fangs), palatine, pterygoid and dentary. Scalation and size data from Brongersma (1934 ), Cogger (1992, 2000), Greer ( 1997), McDowell ( 1967), Scanlon (1985, unpublished data), Shea and Scanlon (unpublished data), Shine ( l 980a,b, 1981), Shine and Keogh ( 1996), Storr ( 1978), Storr et al. ( 1986). Tooth counts fromskel etal material listed in Appendix, with additional data from Boulenger ( 1 896) for Glyphodon tristis, and McDowell ( 1967) for Cacophis harriettae, C. kre.fftii and Aspidomorphus spp.

Genus species Max. SVL Ventrals Subcaudals Maxilla Palatine Pterygoid Dentary

Ca cophis churchi/li 53.8 154-176 25-38 6 9-12 17-18 21-23 harriettae 48.8 168-200 25-45 3-5 9-12 12-18 16-19 krefftii 34.5 140-160 25-40 2-5 9- 11 11-16 14-16 squamu/osus 71.5 165-185 30-50 6-8 11-17 19-24 21-28 Aspidomorphus lineaticollis 48.0 139-174 24-40 11-18 13-16 31-34 31-32 muel/eri 62.0 160- 177 29-40 10-18 16-19 36-40 31-36 schlegeli 50.8 137-160 19-29 11-13 14 24 30 Demansia psammophis 83.5 172-205 63-91 8-13 12-15 26-38 19-25 simplex 43.6 140-158 49-66 9 12 33-34 30 other species 47.6-154.5 160-230 63-113 8-13 10-18 22-38 19-25 Glyphodon dunmalli 56.2 166- 189 37-46 8-9 13 20-23 24 tristis 77.8 171-181 44-5 1 6-8 9-15 16-23 16-20 Furina barnardi 54.8 157-221 35-58 6-7 10-12 15-23 19-2 1 diadem a 34.3 156-203 35-54 4-5 8-12 15-18 14-17 ornata 58.1 164-217 37-63 5 12 19-20 18-19 CA COPHIS MORPHOLOGY AND PHYLOGENY 5

c

FIG. 3. Skulls of Cacophis krefftii and C. squamulosus. C. krefftii (SAM R26974) in (a) dorsal view, and (b) left lateral view of the anterior part of the skull; the left prefrontal and palatomaxillary arch, postorbitals, quadrates and mandibles d are not shown. Braincase of Cacophis squamulosus (SAM R2263A) in (c) dorsal view, and (d) anterior view of the frontals and parasphenoid rostrum; the snout un it, prefrontals, palatomaxillary arches, postorbitals, quadrates and mandibles are not shown. Scale bar= 5.0 mm (for a and FIG. 2. Skull of Cacophis churchilli (QM 153282). Dorsal c only). (a) and ventral (b) views of skull, right mandible, and displaced left palatal elements; (c) right lateral view of skull sheet of connective tissue connecting to the and right mandible, and (d) medial view of left mandible. parasphenoid, prefrontal and palatine shaft, forming the While smaller than some of the C. squamulosus skulls examined, this relati vely large adult specimen (SVL 42 I mm) medioventral wall of the orbit, but even in Cacophis the exceeds them in the development of bony crests for muscle process remains completely free of other bones and does attachment and other fe atures associated with large size. not appear to form a functional articulation with the Scale bar = 5.0 mm. snout. A3. Anterior extent of the ectopterygoid. The anterior of Ca cophis, but a crest-like process is also present in extremity of the ectopterygoid, nearly always its Glyphodon, Demansia, and Aspidomorphus spp. (but anteromedial tip, lies close to the same horizontal plane not in any Furina spp. examined) as well as a number of as the palatine-pterygoid joint, and slightly lateral to it, other Australasian taxa. The crest may be either angular so these landmarks are readily comparable. The com­ or rounded dorsally, and is never as high as Jong in the mon and primitive condition appears to be a outgroups or other hydrophiines. The 'short' choanal longitudinal overlap with the palatine, i.e. the process is therefore considered plesiomorphic for ectopterygoid extends anteriorly somewhat past the Ca cophis (State 1 ). In all Ca cophis palatines examined joint (State 0). This occurs in some Purina, some there is a well-developed choanal process which is Aspidomorphus, and all Demansia examined apart from higher than long (i.e. 'tall'), and directed dorsally or D. simplex. An intermediate state can be recognized slightly anteromedially (Figs 2, 3; also figuredby Greer, where the ectopterygoid extends to approximately level 1997). This condition (State 2), while structurally ap­ with the joint (or within the region of overlap), which proaching that of Naja and more distant outgroups, is characterizes the remaining outgroup tax a (State I). The considered derived within Hydrophii'nae, and diagnostic most derived state, where the ectopterygoid fails to of Ca cophis. reach the palatine (State 2), is not found in the outgroups Loss or reduction of the choanal process in and is hence considered apomorphic in Cacophis, where hydrophiine elapids appears to be related to increased it is the only state observed. longitudinal mobility of the palatines relative to the A4. Supra-anal keels. A patch of keeled lateral scales vomer and snout complex, in contrast to the rotation of is present in the cloaca! region in males of all four the palatine about its contact with the vomer in many Ca cophis species (a series of each examined at the other snakes (McDowell, 1970; cf. Cundall, 1995; AMS). This secondary sexual character is sporadic but Cundall & Shardo, 1995). In incompletely cleaned quite wi despread among colubroids (e.g. Blanchard, skulls the palatine dorsal process is seen to lie within a 1931; Mertens, 1936; Gyi, 1970; Roze, 1996), but has 6 J. D. SCANLON

not been seen in the outgroups or any other Australasian the lower postocular, separating the upper postocular elapids examined. Extent of the keeling is variable when from thetem porals (State 1). Among the outgroups, this present, ranging from barely detectible (one or two state characterizes only Aspidomorphus spp. (it also oc­ keeled midlateral scales on each side in C. krefftiiAMS curs as a variant in some derived Simose/aps, e.g. S. R77370, SVL 237 mm) to extensive (from 12th-last morrisi Homer, 1998). Hence, it is parsimoniously in­ ventral to 20th subcaudal, and extending from lowest terpreted as an apomorphy of Cacophis, convergent in laterals to the paravertebral scale rows in C. Aspidomorphus (and again in Pseudonaja and squamulosus AMS R37187, SVL 410 mm). Two indi­ Oxyuranus, which forma more distantly related clade). viduals with weak keeling (churchilli, AMS Rl 1512, Variation in frequency of contact within Cacophis is R 12480) appear to be female based on tail shape, and treated as a furtherbinary character (C7). some are unlikely to be mature based on size (churchilli A6. Te mporal scales. The practice of writers on Aus­ AMS R 11340, SVL 145 mm; squamulosus R28232, tralian elapids has generally been to recognize two SVL 187 mm). Conversely, no keeling was detected in series of temporal scales, anterior and posterior, and some likely adult males of krefftii (e.g. AMS R8 l 158, give a formula such as '2+2' or '3+4'. McDowell (1967: SVL 235 mm), but generally this seems to be a useful 500) described the widespread 2+2 condition in Aus­ indicator of sex and maturity in each species. Similar tralasian elapids, and introduced the term variability of supra-anal keels is reported within Micru­ 'temporolabial' for the lower anterior temporal "which rus by Roze ( 1996; see her Fig. 7). appears to be the homologue of the penultimate A5. Parietal and postocular scales: occurrence of supralabial of elapids with seven supralabials, but which contact. In most elapids withtwo (or sometimes three) has been squeezed out of contact with the oral border". postoculars, the uppermost contacts a temporal (or Describing the temporolabial as present or absent (e.g. sometimes labial) scale below the parietal (State 0). In Wallach, 1985) is an oversimplification, but a number some of each species of Ca cophis, the parietal contacts of distinct characters can be defined using a more de­ tailed notation. Storr ( 1968) used a three-term formula (e.g. '1+1+2'; see also Aplin & Donnellan, 1999), but without adequate explanation. The following definition gives counts consistent with Storr's on the same speci­ a mens (Scanlon, 1985), and is applicable to most Australasian elapids. Th e postsupralabial is the scale at the comer of the mouth, overlapped by the most posterior supralabial and infralabial but not counted in either series (this follows the usage of Greer & Cogger, 1985, for skinks, and is consistent with supra- and infralabial counts given by most authors). The anterior (or primary) temporal row includes the temporolabial, when distinct - regardless of whether it reaches the lip or contacts a postocular - and any other scales between supralabials and parietal which do contact the postoculars. The oblique row of scales connecting the parietal to the postsupralabial, but ex­ cluding the latter, is considered to be the last row of b c temporals (2nd, 3rd or 4th in Australian elapids), and the formula is given to a corresponding number of terms. Individual scales can be identified by row and position as 1°1, 2°1, 3 °2 etc.; thus, the temporolabial (TL) is 1 °2 in typical Australian elapids. Using this definition, all of the outgroup species (and most other Australian elapids) normally have 2+2+3 (Fig. 4d), while the condition in Cacophis spp. can be d written as 1+3 or 1+2 (Fig. 4a). McDowell (1967: 535) suggests the temporolabial is fusedto the sixth labial in Ca cophis, but this does not explain the reduction from three to two rows oftemporals. Rather, it seems simplest to assume that the single large anterior temporal of Ca cophis represents the fourscales of the ancestralpri­ mary and secondary temporal rows (as suggested by FIG. 4. Head of Cacophis churchilli (SAM R22392) in (a) Greer, 1997). If fusion of adjacent scales (failure of a lateral, (b) dorsal, and (c) ventral views, and (d) Furina ornata (AMS R I I 0357) for comparison of head scales. Scale suture to develop; Resetar & Marx, 1981) represents a bar= 5.0 mm (for a-c only). single evolutionary 'step', the inferred transformation CA COPHIS MORPHOLOGY AND PHYLOGENY 7

from four scales to one could have involved two or more interpreted as adaptations to fossoriality, related to re­ steps, but these are not counted as separate characters duction in head width and the minimization of soil here because ingroup variation is so limited. Variants in accumulation along sutures (e.g. Resetar & Marx, 1981; C. churchilli and C. krefftii do corroborate the sugges­ Savitzky, 1983). While Cacophis spp. are not strictly tion of fusion among temporals (loss of sutures rather fossorial in habit, they utilize crevices and cavities in than loss of scales). For example, AMS R75961 moist soil as refugia and in foraging nocturnally for in­ (krefftii)has '2+ 2' on each side, but on the right side of active prey, mainly skinks (Wells, 1980; Shine, l 980a; the head the anterior temporals are both elongate and Ehmann, 1992). Similar selective pressures would ap­ contact the postorbitals (interpreted as I 0 I =2°1 and ply to the primitive sea snakes, which capture gobiid TL=2°2, where '=' indicates fusion), while on the left fish within burrows on intertidal mud flats (Storr et al., there is a large L-shaped scale (representing 1986; accounts cited by Greer, 1997). TL=l 01 =2°1) with a smaller adjacent scale (2°2) widely A 7. Pale iris. Non-melanin pigmentation of the iris is separated from the postoculars. Similar conditions occur rapidly affected by preservatives or freezing, and obser­ asymmetrically in R81158, R90609, and Rl 14956 vations should be based on live specimens or clear (krefftii), and in R11512 and Rl 1362 (churchilli). Ac­ photographs ( cf. Gillam, 1979). On the other hand, cepting this evidence for fusion of the two anterior 'dark' and 'pale' eyes can usually be distinguished in temporal rows, we may write the formula for the com­ well-preserved material, so two characters are used here mon Ca cophis conditions as 1=1+3 or 1=1+2. The (see also CJ 7 below). Glyphodon and Purina have very variation in the posterior row is used below as evidence dark brown or black eyes (Hutchinson, 1990), while for relationships within the genus (C6). most other elapids, including the other outgroup taxa, Many specimens of Simose/aps and Ve rmicella spp. have the dark pigment varied by a lighter ring, spot or also have I =I +2, but the most common condition in variegations (combined as State 0). A specimen of these genera is 1+1+2 (Storr, 1968; Greer, 1997: 169; Demansiaflagellatio Wells & Wellington, 1985 (a valid Homer, 1998: Fig. 2), and when fusion occurs between species - Shea & Scanlon, unpublished data) at primary and secondary the resulting scale is either Riversleigh, north-west Queensland, had bright red eyes trapezoidal (deep anteriorly and tapering posteriorly) or iJ1 life (pers. obs.), so apparently, like Cacophis, has lit­ long and shallow, in either case quite different from that tle or no melanin in the iris. Hence Demansia is scored in Cacophis. Moreover, contrary to McDowell ( 1969a), as polymorphic, although most species - including D.

the temporolabial does occur as a separate element in simplex - have only a narrow pale ring. The almost uni­ this group (Simoselaps warro normally retains the an­ formly 'pale' iris of Cacophis (State I) appears to be cestral 2+2+3, while 2+1+2 is a common variant in the diagnostic of this genus (Hutchinson, 1990); further S. sem ifasciatus group), which has not been observed in comparison might justify defining an intermediate state Cacophis. for squamulosus, which appears to have more speckling Some specimens of Demansia spp. (like all species of or variegation than its congeners. Pseudonaja, and some Oxyuranus microlepidotus; Storr et al., 1986; pers. ohs.) have a temporal condition POSSIBLE APOMORPHIES SHARED WITH OUTGROUP TAXA even more similar to Cacophis superficially ('I +2'). A number of characters of Ca cophis spp., despite However, the complete formula in these cases is 1+2+3 being possibly or actually derived within the Australa­ or 1 +2+4, indicating retention of three distinct rows. sian radiation, also occur in both outgroup clades and This results from a single fusion between the are likely to be locally plesiomorphic (Table 2, B 1-1 O; temporolabial and 6th supralabial, as shown by the con­ see also generic diagnosis below, and Appendix 2). cave upper edge and frequent partial suture of the labial These characters will not be discussed further here. scale, and comparison with normal (or in Pseudonaja, occasional atavistic) individuals with 2+2+ 3. CHARACTERS VARIABLE WITHIN CACOPHJS Conditions precisely equivalent to those of Cacophis (I+3 or I+2, with a deep single anterior temporal) are Characters which vary among the four species of found in the primitive marine hydrophiines Ephalophis, Cacophis (referred to here by their species names alone) Parahydrophis; Hydrelaps and Disteira, and a further provide the basis of a phylogenetic analysis carried out fusion to 1+1 occurs in some Parahydrophis below. I include several autapomorphies of terminal (McDowell, 1969b, 1972, 1974; Burger & Natsuno, taxa, one of which is behavioural rather than strictly 1974; figuresin Storr et al., 1986; Cogger, 1992, 2000). morphological, in order to provide adequate diagnoses Most other sea snakes have an increased number of for species as well as higher groups. The distribution of temporals, oftenquite irregular and presumably second­ character states in outgroup genera and ingroup species arily fragmented. is given in Table 3. The state seen in Cacophis is here considered func­ Cl. Maxillary tooth number. Observed ranges of tionally analogous to those of other lineages with a tooth (alveolus) counts for the maxilla, palatine, ptery­ reduced number of temporal sutures, but to have been goid and dentary of ingroup and outgroup taxa are given derived independently from the common ancestral con­ in Table 1. The number of maxillary teeth behind the dition 2+2+ 3. Head-scale fusions in snakes have been fangs is often relatively high among Australasian and 8 J. D. SCANLON

TABLE 2. Distribution of character states (characters labelled ' A ' and 'B' in main text and Appendix 2) in outgroup genera and Cacophis. Abreviations for character types: b, binary; u, unordered; 012 (etc.), ordered multistate. Characters marked *are cladistically uninformative for analyses performed here.

Character A 1 2 3 4 5 6 7 B 1* 2* 3* 4* 5* 6* 7* 8* 9* JO type b 012 012 b b b b b b b b b b b b 0123 012

Aspidomorphus 0 01 0 0 0 1 1 01 01 3 2 Demansia 01 01 0 0 0 01 0 01 0 01 01 01 3 01 Glyphodon 0 I 0 0 0 0 0 I 01 12 Furino 0 0 01 0 0 0 0 0 01 01 01 23 I Ca cophis 2 2 I I 3 2

TABLE 3. Distribution of character states (characters labelled 'C' in main text and Appendix 2) in out group genera and Cacophis spp. Abreviations for character types: b, binary; u, unordered; 012 (etc.), ordered multi state. Characters marked * are cladistically uninformative for analyses performed here. Character (C) I 2 3 4 5 6 7 8 9* 10 I I 12 13 14 15* I 6* I 7 18 19* type b b b b 012 b b b 012 0123 0123 b b b b b 012 b b

Aspidomorphus 0 0 0 0 0 0 I 0 I 123 23 0 0 0 01 01 0 0 ? Demansia 0 0 0 0 0 0 0 01 01 013 0 0 01 01 01 0 0 0 Glyphodon 0 0 0 01 2 0 0 0 01 01 0 0 0 0 01 0 ? Furino 01 01 I 2 0 0 12 012 0 0 0 0 0 0 0 C. churchilli 0 0 I 0 2 3 0 I 0 0 2 0 C. harriettae 0 0 0 0 0 I 2 0 0 C. krefftii I I I I 0 I 0 I 2 3 2 I 1 0 0 0 I 0 C. squamulosus 0 0 0 0 0 0 0 0 0 0 0 0 0 marine elapids (Hydrophiinae), whereas no African, C3. Pte1ygoid tooth number. Two states can be rec­ Asian or American elapids are reported to have more ognized on the basis of non-overlapping ranges: 19-24 than four (Bogert, 1943 ) . Some counts in the literature (State 0) and 11-18 (State I). The outgroup species all are probably umeliable, and dentigerous elements other have I 9 or more except for F. diadema and F. ba rnardi, than the maxilla have been mostly neglected by previous which overlap both ranges, so State 0 is considered workers, but tooth counts on all bones of many elapids plesiomorphic. have recently been tabulated by Greer (1997). C4. Dentary tooth number. Again, two non-overlap­ Nine skulls of squamulosus examined show a range ping states can be defined for Ca cophis: more than 20 of 6-8 maxillary alveoli behind the fangs (Worrell, 1963 (State 0) and fewer than 20 (State I). Most of the reported 7-10); harriettae usually has 5 (seven of eight outgroup species exhibit State 0, but Glyphodon tristis specimens examined), but only 3 in one specimen; and Furino spp. fall mainly in the range of State I, so churchilli has 6 in both skulls examined; and krefftii3- polarity is equivocal. 5 in three skulls (but Worrell, 1963; McDowell, 1967 C5. Na sal and preocular: frequency of contact. and Greer, 1997, all report 2 in this species, so the range These scales are either in contact or narrowly separated can be given as 2-5). Based on these figures, two non­ (by contact between the prefrontal and a supralabial) in overlapping ranges can be recognized: 6-8 (State 0) and Ca cophis, but variable within each species so that states 2-5 (State 1 ) . State 0 is regarded as plesiomorphic, being can be defined based on frequencies. State 0 (usually similar to counts in Glyphodon, overlapping with contacting) occurs in kreffiii ( 15 bilateral and I unilat­ Demansia and Furin o, but lower than any in eral of 16 specimens; on the remaining side, nasal and Aspidomorphus. The higher counts in Demansia and preocular separated by a distinct 'loreal ') and Aspidomorphus are a likely synapomorpy of these gen­ squamulosus (I1 and 3 of 14; again one with a loreal on era, while reduction has probably occurred one side), and can be identified with the state in independently in Furino, as in several other Australian Aspidomorphus and Demansia (separation rare or ab­ lineages. sent). State 1 (usually narrowly separated) characterizes C2. Palatine tooth number. Despite overlapping churchilli (0 bilateral contact, I unilateral of I 5) and ranges, C. squamulosus clearly tends to have a higher harriettae ( 1 bilateral of I 5; indeterminate unilaterally number of palatine teeth than the other species, and thus in another where the preocular and prefrontalare fused). two states are recognized: usually more than I I (State In G/yphodon and Furino the scales are widely sepa­ 0), usually I I or fewer (State 1 ). The mostly high counts rated (State 2). While State 0 is the only ingroup state in Aspidomorphus, Demansia and Glyphodon imply shared with outgroup taxa, polarity cannot be inferred if that State 0 is plesiomorphic, while Furina tends to ha­ the character is interpreted as an ordered morphocline ver lower tooth numbers (as on the maxilla). (0-1-2). CA COPHIS MORPHOLOGY AND PHYLOGENY 9

C6. Posterior temporal scales. As noted above (char­ phic forthis character. C. krefftiiis one of the smallest of acter A6), variation is observed in the number of scales elapids (the largest specimen examined, AMS Rl3000, in the last row of temporals. Three posterior temporals is bas SVL 345 mm, tail 39 mm), and all outgroup species the usual condition in all outgroups and most other except Furina diadema have a greater maximum SVL, elapids (State O); in a sample of C. squamulosus exam­ so that it also represents an apomorphic extreme of the ined, three posterior temporals occur in 13 of 14 genus (State 2, SVL <35 cm). As each of the apomor­ specimens, so this species is assigned the primitive state. phic states occurs in a single species, this character The frequentoccurrence of only two posterior temporals contributes no cladistic information within Ca cophis. is recognized as an apomorphy (State I) shared by the However, this 'uninformativeness' depends on the par­ remaining species: churchilli has two in 12 of 15 (and ticular outgroup arrangement adopted here, and could unilaterally in another), harriettae in 14 of 15, and possibly change if Cacophis were later determined to krefftii in every one of 16 specimens. The relative sizes have a different pattern of relationships with other taxa. of the scales indicate that it is the upper two of the three CJO. Ventral scale number. Ventral and subcaudal scales which fuse (3°1=3°2). ranges of Cacophis species and outgroups are shown in C7. Parietal and postocular scales: fi·equency of Table I; detailed frequencydistributions would be pref­ contact. Contact of parietal and lower postocular scales erable (cf. Wiens, 1998) but are not currently available is a shared derived condition of all Cacophis species for most species. All outgroup genera, and nearly all (see A5), but two distinct levels of frequency areappar­ outgroup species, have ranges overlapping from 170 to ent in samples examined for this trait. Relatively low 175 (all lower in Aspidomo1phus schlege/ii and frequency is coded as plesiomorphic (State 0), present in Demansia simplex, all higher in some other Demansia harriettae (contact bilateral in I, and unilateral in 5 of species). Three species of Cacophis also overlap in this 15 specimens) and krefftii (I and 3 of 16). High fre­ 'core' range, so the exception (krefftii) is regarded as an quency (State I) characterizes churchilli ( 1 1 and 3 of apomorphic extreme. The high ventral counts charac­ 15) and squamulosus ( 12 and I of 14). terizing some harriettae may also be apomorphic, as CB. Division of nasal scale: fr equency. The nasal they are outside the ranges of Glyphodon and scale may be either single (pierced by the nostril) or di­ Aspidomorphus. In order to utilize the maximum possi­ vided (separated into anterior and posterior scales by ble cladistic information from the data on ranges, each grooves or sutures above and below the nostril). Com­ ingroup species is assigned a distinct state, and the four plete separation of the nasal by the nostril extending its states are assumed to form a morpbocline in the same full depth, from supralabial to intemasal, occurs in some order as the maximum and minimum observed ventral other elapids (including some outgroup species), but is counts (State 0, harriettae; 1, squamulosus; 2, not observed in Ca cophis. Division of the nasal is churchilli; 3, krefftii). Outgroup species are assigned intraspecifically variable within Ca cophis, so two states the same state(s) as that of the ingroup species with are recognized on the basis of frequency of division: which it most strongly overlaps; polarity remains inde­ high (State 0) in churchilli (bilateral in 10 and unilateral terminate. in 3of15 specimens) and squamulosus (13 and 0 of 14); CJ /. Subcaudal scale number. As long-bodied and low (State I) in harriettae (2 and 0 of 15) and snakes may have short tails (and vice versa), the ventral krefftii (0 and 0 of 16). Because of the pattern of varia­ and subcaudal scale counts are considered independent tion in the outgroups, polarity can not be assigned to this characters. This character can be defined in the same character. way as the previous one - the four states ordered as the C9. Body size (m aximum snout-vent length [S VL}) . maxima for the ingroup species (State 0, squamulosus; The species of Cacophis vary considerably in size (Ta­ I, harriettae; 2, krefftii;3, churchilli) since three of the ble 1; means might be preferable as the basis for this minima are equal. Among the outgroups, counts below character, but good samples are not available for all spe­ 35 occur only in Aspidomorphus; State 0 is parsimoni­ cies). A linear size increment close to the cube root of ously considered plesiomorphic for Cacophis. two (I.26 approx.) has been reported for sympatric spe­ CJ 2. Ventral melanin pattern. This character con­ cies-pairs in numerous animal lineages by Hutchinson cerns only the distribution of dark brown or black, ( 1959) and others (see Sweet, 1980). Inthe three south­ alcohol-insoluble pigment on the ventral surface; varia­ ern (sympatric) species of Cacophis, maximum SVL tion in dorsal colour is more continuous, and differs by ratios greater than 1.4 (implying ratios of 2.75 attributable to the combination of melanin and caroten­ or more in mass), so that three distinct character states oid patterns with schemochromes or structural colours can be recognized. Most churchilli are relatively small, (the latter responsible for whites as well as the bluish but the largest examined (QM J67837; SVL 538 mm, colour common in krejftii,and as a component of greens tail 58 mm) is slightly longer than the maximum record­ in some outgroup species; cf. Fox, 1953; Bechtel, 1978). ed for harriettae (Shine, I 980a), so these two species The outgroups vary considerably in ventral colour; are assigned the same intermediate state (1). Among the Demansia spp. range from dark grey to immaculate outgroups only Glyphodon tristis and some species of white or yellowish, often with a median dark line or Demansia reach greater lengths than squamulosus, so zone. The venter is usually white in Furina and large size (State 0, SVL > 70 cm) is probably apomor- Glyphodon; 'smoky' grey, peppered more or less 10 J. D. SCANLON

densely with melanin granules, in Aspidom01phus spp. line in the other species of Cacophis, but varies in width: and some individuals of Glyphodon spp. The dark slate­ about fourscales wide in harriettae, i.e. similar in extent grey or black venter of harriettae and churchilli is to that of squamulosus but without a dark median zone, similar to the conditions in some Demansia (vestigiata, and one or two scales wide in krefftii and church ill. Two some torquata) and Aspidomorphus muel/eri, and ap­ binary characters are used, for separation vs. contact of proached by some Glyphodon, so unifonnlydis tributed the lateral pale markings, and width of the collar. Furina dark pigment is here presumed plesiomorphic (State 0). and G/yphodon are coded as not comparable for the first Southern (NSW) C. squamulosus have irregulanimals; I have not examined living Aspidomor­ the black blotches on the ventrals are narrow and also phus, but O'Shea (1996) has photographs of two species form either a midventral line, or three distinct longitudi­ in life. In Glyphodon, Furina and krefftii (in between nal rows. I have also seen a unifonnly 'peppered' dark markings) the venter is white or very pale yellow condition (precisely as in some Aspidomorphus) in a (carotenoid very faint or absent); white or yellow also Sydney specimen of squamulosus, but this is rare. The occurs in some Demansia spp. including D. simplex. state in krefftiiis also contrasting, usually with a median Aspidomorphus lineaticollis and some other Demansia line under the tail, but more regular on the body than have pink or orange ventral colours, and in squamulosus squamu/osus; the ventrals have a yellowish base and the venter varies among individuals fromorange or pink black posterolateral comers, typically forming a double to deep red, the same colour also suffusingthe light cen­ saw-tooth pattern,but oftenjo ined as a continuous dark tres on the dorsal scales and sometimes the collar. In one border across the free edge of some or most scales (as in specimen from a variable population at Greenwich, the holotype of C. fo rdei Krefft, 1869). Despite the dif­ NSW, the venter was a very deep red posteriorly, and fe rences in detail, squamu/osus and krefftii are coded there were dark orange to red centres on all of the dorsal with the same apomorphy (State 1 ). scales of the body and tail (pers. obs.). The dark-bellied CJ 3, Cl 4. Facial pattern and collar shape. The col­ species of Ca cophis and A. muelleri, appear to Jack red our pattern on the face and nape is very similar in the pigments since none are visible on the sides or dorsum. four species of Ca cophis; the dark upper surface of the Since red is present in only one ingroup species, this head is bordered by a more or less continuous pale band character is cladistically uninformative. extending from the rostrum, through and over the eyes CJ 6. Carotenoid pigment on fa ce and collar. The fa­ and onto the nape; the pale stripe is broken up by dark cial stripe and collar are normally yellow in three of the markings at scale boundaries in the labial and temporal species (State 0), but usually white (sometimes faintly regions. Similar patterns are found in Aspidomorphus yellow) in harriettae (State 1 ); in this species the pale and Demansia (most complete in some A. lineaticollis centres of the dorsal scales are also whitish (an extreme and D. torquata ), but patterns in Glyphodon and Furina condition is shown by a specimen illustrated in Wilson are unlike these, with more discrete light and dark areas. & Knowles, 1988: pl. 723). Yellow markings on the face A dark comma-shaped or 'bridle' marking from the eye and nape are considered plesiomorphic as they are usu­ to the lip is present in all Cacophis and some ally present in nearly all outgroup species. In Aspidomorphus and Demansia (also some Pseudonaja). Glyphodon, yellow pigment is nearly or completely ab­ In squamu/osus the pale facial band is continuous sent except for the nape patch of G. tristis, while in with longitudinal stripes on the neck, somewhat ex­ Aspidomorphus lineaticol/is the face stripe is white in panded towards the midline but separated from each the specimen shown by O'Shea (1996: 149). other by dark-pigmented vertebral and paravertebral Cl 7. Iris colour. This character concerns variation scale rows (sometimes greatly elongated as shown by not due to melanin (see A7 above). Because of the den­ Gow, 1989: 95 and Greer, 1997: 178, rarely connecting sity of melanin in the irides of Furina and Glyphodon, to form a complete collar). This is quite similar to the the presence or colour of other pigments in these taxa 'upper light line' present in some Aspidomorphus has not been observed; their relatives Simose/aps populations (McDowell, 1967) which are most similar bertholdi and S. littoralis have white eyes, but may have to Ca cophis in pigmentation, and possibly also compa­ apomorphically reduced carotenoid as well as melanin. rable to the pale or reddish dorsolateral streaks in some Iris colour in life is not known for all Demansia and Demansia spp. (common in D. psammophis). The 'bro­ Aspidomorphus species, but in most of them, reddish ken' collar is thus regarded as the plesiomorphic pigments combine with the melanin to produce orange­ condition for Cacophis. One species of Demansia (D. brown eyes (e.g. A. /ineaticol/is, O'Shea, 1996; torquata) has a complete, narrow pale collar continuous Demansia spp. illustrated in Storr et al., 1986; Wilson & with a pale facial stripe; pale or dark collars in other Knowles, 1988; Gow, 1989; Ehmann, 1992; Cogger, outgroups are less similar (involving contrast between 1992, 2000). Red eyes, which also occur in krejfti, are head and dorsal ground colour, or not continuous with therefore assumed to be plesiomorphic for Ca cophis facialmarking s). The collar is complete across the mid- (State 0). The iris is predominantly yellow in CA COPHIS MORPHOLOGY AND PHYLOGENY 11

squamulosus (State 1 ), and at least partly white in dered) are parsimony-informative for more inclusive harriettae and churchilli (State 2). Because the states analyses and hence retained. Of characters C 1-19 which can be ranked in order of intensity of colour, they are vary among Cacophis spp., 14 characters are binary; provisionally treated as ordered (0-1-2). characters C9 and Cl7 are assumed to form ordered CJ 8. Pupil shape. Pupils are stronglyvertically ellip­ three-state morphoclines, and C5, Cl 0 and Cl 1 have tical in squamulosus and harriettae, as in Glyphodon, four ordered states (Table 2). However, characters C9, Furina and most Aspidomorphus (only weakly so in A. C15, C16 and Cl9 are cladistically uninformative (de­ muelleri; McDowell, 1967) (State 0), but weakly oval or rived states, or combinations of states for those coded as quite round in the other species of Cacophis, and round polymorphism, occur in single terminal taxa). When un­ in all Demansia (State 1 ). informativecharacters are excluded, the effective size of CJ9. Defensive threat display. In the threat displays the data set is therefore 16 binary, 5 three-state ordered, of all Ca cophis spp. the head is raised and angled down­ and 2 four-state ordered characters. ward (displaying the pale collar); fromthis position they There are two equally most parsimonious trees, one may strike forward and downward, but almost never ac­ with the topology (Aspidomorphus ((Demansia tually bite. In three of the species the neck is held (Glyphodon, Furina)), (C. squamulosus (harriettae straight and the jaws not or only slightly expanded dur­ (churchilli, krefftii))))), and the other diffe ring only in ing the display, while squamulosus is distinctive in two the interchange of harriettae and churchil/i. For both, ways: the neck is formed into S-shaped lateral curves tree length = 46 steps, consistency index (Cl) = 0.652, and the quadrates and rear of the mandibles are spread homoplasy index (HI) = 0.348, retention index (RI)= laterally, making the head much wider than the neck 0.686, rescaled consistency index (RC) = 0.448. How­ (e.g. Grigg, Shine & Ehmann, 1985: pl. 5; the narrow ever, these treesconflict with the outgroup assumptions dark zone interrupting the pale collar, and longitudinal (Fig. I), and because the characters analysed are chosen patternon the lateral neck scales, tend to exaggerate this for informativeness relative to Cacophis rather than the visual effect). All three Furina spp. have stiff-necked outgroups, their basal nodes are considered unreliable. displays most like the smaller Cacophis spp. (e.g. Greer, Therefore, a constraint tree was used to enforce the 1997: 161 ), and Demansia spp., although relying on (Furina, Glyphodon) and (Aspidomorphus, Demansia) speed and venom in defenceand thus apparently lacking clades. These topological constraints reduce the space a comparable 'bluff display, have a similar raised-head ofunrooted treesfor eight taxa from 10 395 to 105 dis­ 'alert' posture while foraging (e.g. Scanlon, 1998). tinct alternatives. With the constraints, the single most Glyphodon tristis has a different defensive display parsimonious tree has total length 48 steps (Fig. 5); Cl = (thrashing wildly in a horizontal coil, head- and tail-hid­ 0.625, HI = 0.375, RI = 0.647, RC = 0.404. Cladistic ing), while G. dunmalli is described as inoffensive relationships among the four ingroup species are identi­ (Wilson & Knowles, 1988; Ehmann, 1992; Greer, 1997: cal to one of those found in the unconstrained analysis, 162). I know of no published descriptions of defensive where Cacophis also emerged as monophyletic. or foraging behaviour in Aspidomorphus; A. muelleri The degree of support for each grouping was strikes (and bites) froma more-or-less upright defensive measured by the support index (Bremer, 1988), posture when prevented from escaping (S. Richards, calculated in PAUP using a command filegenerated by pers. comm. 2001), but on present evidence this cannot TreeRot (Sorenson, 1999). These commands were be identified with either of the states in Cacophis. Al­ modified to use branch-and-bound rather than a though a number of other (mostly large) Australian heuristic search algorithm. Nonparametric elapids have high lateral S-bends in the defensive dis­ bootstrapping (I 0 OOO replicates, employing branch­ play (e.g. some Ps eudonaja spp., Oxyuranus and-bound search) was also used to assess the microlepidotus and Hoplocephalus spp.), the behaviour robustness of each clade ( apa11 from the outgroup cl a des in squamulosus is considered unequivocally derived where monophyly was enforced). Support for since it is not paralleled in the outgroups; however, this monophyly of Cacophis with respect to the outgroups apomorphy is not cladistically informative. (support index 4, boostrap frequency 94%) and of the (churchilli, harriettae, krefftii) clade (5, 94%) are PHYLOGENETIC ANALYSIS strong, but that for a clade comprising churchilli and krefftii is weak (I, 58%). The alternative grouping of PARSIMONY WITH ORDERED CHARACTERS harriettae with kreffiii was found in 30% of bootstrap Because of the small number oftaxa, it is not consid­ replicates, but was less parsimonious (by one step) ered useful to construct a 'hypothetical ancestor': the under the assumed constraints on outgroup outgroup genera can be included explicitly along with relationships. ingroup species, and still allow exhaustive search of tree On the preferred phylogenetic hypothesis (Fig. 5), topologies. characters Cl, C2, C4, C5, C7, CS, Cl 1, C12, Cl 7, Characters A 1-7 and B 1-10 are invariant within Cl 8, A5, and B 10 are homoplasious (Cl = 0.5 in each Ca cophis, hence uninformative for intrageneric rela­ case except C4, C7, CS [0.33], and Cl 1 [0.60]). Of tionships. However, A 1-7 (five binary, and two these, six are convergences between ingroup and three-state ordered characters) and BIO (three-state or- outgroup taxa (not discussed furtherhere), while those 12 J. D. SCANLON

steps, Cl = 0.711, HI = 0.290, R1 = 0.732, RC = 0.520. -2,. Aspidomorphus As before, this treeconflicts with the outgroup assump­ tions (Fig. 1 ). Demansia When the unordered data are reanalysed with the two outgroups constrained to be monophyletic, there is a sin­ 2, . Glyphodon gle most parsimonious tree withtotal length 41 steps; Cl = 0.659, HI = 0.342, RI = 0.659, RC = 0.434. This tree Furina differs from that found in the 'ordered' analysis (Fig. 5) in the interchange of churchilli and harriettae, i.e. the - sister group of kreffiii is found to be harriettae. In a C squamulosus bootstrap analysis under the same constraints, 5, 4 monophyly of Cacophis is found in 98% of replicates, - 94 C harriettae and a clade comprising churchilli, harriettae and kreffiii 4, 5 in 90% (cf. values in Fig. 5). The sister taxon to kreffiii 94 C churchilli is foundto be harriettae in 52% and churchilli in 22.4% 0, 1 ofbootstrap replicates. 58 - C krefftii REVISED DIAGNOSES FIG. Cladogram showing most parsimonious hypothesis S. The previous diagnosis of Cacophis is that of Cogger of phylogenetic relationships among Cacophis spp., with multistate characters treated as ordered where applicable (2000; little modified from Cogger, l 975; see also (analyses described in text). Numbers above branches show Hutchinson, 1990; Greer, 1997). The revised diagnoses Bremer support for unconstrained analysis and when below list autapomorphies of Cacophis, included monophyly of both outgroups is enforced; numbers below clades, and species, and to facilitate comparison with branches bootstrap frequency (percent) in constrained other taxa, I also list many plesiomorphic conditions for analysis. Bootstrap values not applicable for the two outgroup clades (i.e. fixed at I 00% by constraint tree). the genus, including both widespread characters and those shared with only a few outgroup taxa. Character shown in bold involve convergence or reversal within states discussed in the text are identifiedby their labels Ca cophis. Characters Cl and C4 (tooth numbers on the (Al (O) etc.). Some of the characters listed but not men­ maxilla and dentary) appear as putative synapomorphies tioned elsewhere in the text, including features of the of harriettae and kreffiii, but they could easily have un­ skull, dentition and vertebrae, will be discussed else­ dergone convergence or reversal together due to where (in prep.). Diagnoses of clades and species within common genetic basis (pleiotropy) or selective factors Ca cophis, based on the most parsimonious cladogram (i.e. they are probably not independent, cf. Lee, 1998). discovered in this work, list apomorphies according to The frequency-based head-scale characters C5, C7 and the delayed transformation ( deltran) optimization as­ C8 conflict with each other as well as the optimum to­ sumption; those invariant under acctran and deltran pology, implying their independence but also the )ability assumptions (unambiguous synapomorphies) are of such variables, consistent with 'neutral drift' or fluc­ marked with an asterisk. tuating selection on the equilibrium frequenciesin each species. Cl2 (contrasting 'barred' ventral pattern),link­ CACOPHJS GUNTHER, 1863 ing squamulosus and kreffiii, is likely to be convergent Autapomorphies. Loss of paired dorsal foramina of as the patterns in these species differ in detail and may parietal bone (Al(l)*; one member of pair may occur); thus be considered to fail the similarity test of homol­ palatine choanal process relatively tall, i.e. higher than ogy. long (A2(2)*); ectopterygoid does not extend forward

ALL CHARACTERS UNORDERED past pterygoid-palatine joint (A3(2)*); supra-anal keels frequently present (A4(1)*; often extensive in adult The definitions of multi-state characters used above males, occasionally developed in females and juve­ rely on the ordering implicit in topological and numeri­ niles); parietal scale may contact lower postocular cal relationships, i.e. on abstract properties of number (A5(1); see also C7); temporal formula reduced from rather than independent assumptionsabout evolutionary 2+2+3 to 1+3 by fusion of two anterior rows (1=1+3; processes. It is nevertheless possible to analyse the A6(1)*, see also C6); iris pale (A7(1)*, melanin pig­ 'same' matrixwhile disregarding this trivially available mentation reduced to faint speckling; Cl 7(1 ), information about order, but character Cl0 (based en­ carotenoid yellow rather than reddish). tirely on rank order of meristic values) then becomes Features shared with outgroups. Small ( C9(0/l ); less parsimony-uninformative, in addition to those excluded than 75 cm snout-vent, I m total length), terrestrial, above. The shortest tree overall then has the topology oviparous hydrophiine elapid snakes, dark brown or (Aspidomorphus ((Demansia ( Glyphodon, Furina)), greyish above with pale centres on many of the dorsal (squamulosus (churchilli (harriettae, kreffiii))))), one of scales tending to form a longitudinal pattern on the the two found in the previous analysis; tree length = 38 flanks and neck; no trace of transverse bands on the CA COPHIS MORPHOLOGY AND PHYLOGENY 13

body or tail. A yellowish band, including dark spots and vex laterally; two anterior canaliculate fangs with ridged variegations, across the snout (including most or all of or striated surfaces, followed after a diastema by 6-8 the intemasal scales), over and through the eyes and small solid or grooved teeth (Cl (O)) extending onto a temporal region, and expanding towards the midline at rod-like posterior process defined by concavities later­ the rear of the head, beginning one to several scale-rows ally and medially. Palatine with 11-17 teeth (C2(0)), behind the parietals (C13(0}, not forming a continuous extends approximately as far anteriorly as maxilla; pos­ transverse band; C14(0), extending four or more scale terior end with lateral and medial processes clasping rows back on the neck); dark variegations absent or faint anterior end of pterygoid; without lateral (maxillary) in the nuchal portion. Pale facial band broken by a dis­ process or sphenopalatine foramen, but with distinct tinct dark 'bridle' marking joining the eye to the lip. dorsomedial 'choanal' process on the posterior part of Eyes equal or smaller in diameter than their distance the shaft (A2( 1 /2)). Ectopterygoid not extending from the lip, pupil vertically elliptical (Cl 8(0)). Venter anteriorly beyond pterygoid-palatine joint (A3( 1/2)); with diffuse dark melanin pigment (C12(0)); yellowish lateral edges of ectopterygoid parallel anteriorly, an­ carotenoid pigment also present ventrally (Cl 5(0)). gling posteromedially at a slight knob-like prominence Snout short and rounded, no canthus rostralis; nasal usu­ level with the rear of the maxilla. Pterygoid with 19-24 ally divided (C8(0)), usually in contact with preocular teeth (C3(0)); lateral edge with an angular inflection or (C5(0)); preocular contacts second supralabial; six triangular process for ectopterygoid attachment; pos­ supralabials, third and fourth entering eye; seven teromedial edge usually convex, posterior tip blunt. infralabials. Internasal and prefrontal scales usually Dentary with 14-28 teeth (C4(0/l)), increasing steeply overlapping left over right (krefftii, Greer, 1993; re­ in size from inflected anterior tip to two subequally maining species, pers. obs.). 17 to 23 longitudinal rows large, robust teeth (B7(1 )) with anterolateral grooves; of dorsal scales at the first ventral (pers. obs.), reducing large 6th, 7th or 8th tooth usually followed by a gap (di­ to 15 on neck and rarely reducing again before vent astema; B8(l)) and shorter, more recumbent, posterior (B9(3), B 10( 112)). Anal and usually all subcaudals di­ teeth. vided (sometimes a few anterioror scattered subcaudals Zygosphenes of vertebrae in dorsal view trilobate, single). Dorsal scales matt to slightly glossy, lacking with rounded median Jobe; prezygapophyseal processes keels -(except in the cloaca) region, see A4). Skin be­ prominent, acuminate (terrns of Auffenberg, 1963) and tween dorsal scales pale (light brown or grey). Tongue angled anterolaterally; hypapophyses of posterior trunk with dark pigment only on middle portion, so base red­ vertebrae in lateral view with angle separating oblique dish and tips pale pink or white in life. Superficial fromhorizontal portions of ventral edge (B6). venom-gland constrictor muscle (m. adductor extemus Habitat is wet sclerophyll or rainforest; nocturnal, superficialis) without separate quadrate head from rear sheltering by day under rocks, Jogs, leaflitter, or in cavi­ of gland, and m. adductor externus medialis exposed ties associated with ant or termite nests; diet mainly of posterior to superficialis (i.e. 'Glyphodon type' of diurnal skinks captured at night under cover, also frogs, venom-gland musculature; McDowell, 1967). small snakes, and reptile eggs (Shine, l 980a ). In defen­ Hemipenis forked, apical lobes with terminal awns; ba­ sive threat display, the anterior part of the body is raised sal portion nude, bounded distally by a row of weakly stiffly and the head turned downward but not markedly enlarged spines (Keogh, 1999). flattened (Cl9(0)). Frontal bones together oval or diamond-shaped, pre­ frontalarticulated to anterolateral border but not usually CACOPHIS SQUAMUL OSUS (DUMERIL, BIBRON & reaching parietal or postorbital (so frontalnearly always DUMERIL, 1854) narrowly enters orbital margin) (B l(l)); interolfactory Parietal contacts lower postocular in maj ority of pillars of frontals as wide as the frontal-septomaxilla specimens (C7(1)*); snout-vent length may exceed 70 contact (B2(1 )). Postorbitals in edge-to-edge contact cm (C9(0)*); dark ventral pigment usually formingdis­ with parietal (allowing mediolateral but not anteropos­ tinct blotches or bars across base of each ventral scale, terior kinesis), not or barely reaching frontals, so and a zig-zag median line on the subcaudals (Cl 2(1 )*); parietal may enter orbital margin. Parietal relatively carotenoid pigment suffusing ventral and lateral scales long, narrow and slightly bulbous (not constricted), with reddish (pink or orange to deep red; C 15( l )*); in defen­ triangular supraorbital processes clasping frontals, dis­ sive threat display, neck held in lateral S-shaped coils, tinct and elongate but narrow postorbital processes, and and rear end ofjaws spread laterally to widen and flatten weak adductor crests either separated or just meeting the head (Cl9(l)*). posteriorly (not forming a sagittal crest except in the largest individuals). Parasphenoid in ventral view trian­ (CACOPHIS HARRJETTAE, C. CHURCHILL/, C. KREFFTJI) gular, not narrow and awl-like. Fenestra ovalis in opisthotic-exoccipital open laterally for its full width Palatine with 11 or fewer teeth (C2(1 )*); pterygoid (from border of prootic ), so shaft of stapes not enclosed with fewer than 19 teeth (C3(1)*); usually only two laterally by bony crista circumfenestralis. Maxilla ex­ scales in posterior temporal row (C6( l )*); subcaudal tends to or beyond posterior limit of orbit (B4( I)); count may be lower than 30 (Cl 1 (l )*); pale band con­ suborbital portion smoothly concave dorsally and con- tinuous across dorsal midline at back of head (Cl 3(1 )*). 14 J. D. SCANLON

CACOP HIS HARRIETTAE KREFFT, 1 869 most derived species, krefftii, is either churchilli or Maxilla with fewer than six teeth behind diastema harriettae, and less likely to be a clade comprising both. (Cl(!)); dentary with fewerthan 20 teeth (C4(1 )); nasal The alternativepref erred here is the treeobtained in the and preocular usually separated (C5(1 )); nasal usually 'ordered' analysis with outgroup constraints (Fig. 5). undivided (C8(1 )); ventral count not Jess than 170 and This hypothesis is also the only one in which the two may exceed 175 (Cl 0(0)*); collar (and longitudinal pale most recently separated species have disjunct geo­ graphic distributions, consistent with a vicariance stripes on body) usually white, not yellowish (Cl 6( 1 )*); occurs fromGosf ord, NSW, to Mackay iris mainly white (Cl 7(2)). process: krefftii (e.g. QM Jl4287) on the Queensland coast, while churchilli is found further north, from Townsville (CACOP HIS CHURCHJLLI, C. KREFFTll) (13640) to Mossman (J5193). The greater divergence of Ventral count Jess than 176 and may be Jess than 165 krefftii (autapomorphy in features such as head shape, (C 10(2)*); subcaudal count not exceeding 40 eye colour, and small body size) could be interpreted as (Cl 1(2)*); pale collar only one or two scales wide character displacement due to selection, since it is (Cl 4(1 )*); pupil round or only slightly elliptical broadly sympatric with both harriettae and (Cl8(1)*). squamu/osus, whereas the distribution of churchilli overlaps little, if at all, with either species (see distribu­ WELLS WELLINGTON, CACOPH!S CHURCHILLI & 1985 tion maps in Wilson & Knowles, 1988, and Ehmann, Nasal and preocular usually separated (C5(1)); pari­ 1992; those in Cogger, 1992, 2000 are less accurate, and etal usually contacts lower postocular (C7(1 )*, see also in Longmore, 1986 the maps forha rriettae, krefftii, and A5); subcaudal counts Jess than 40 (Cl 1 (3)*); iris partly squamu/osus include misidentified records of white (C l 7(2)). churchilli). Interpretation of the cladogram in terms of species­ CACOPH!S KREFFT!! GONTllER, 1 863 level historical biogeography (phylogeography) is Maxilla with fe wer than six teeth behind diastema complicated by the broad sympatry between species; (C 1 (I)); dentary with fewer than 20 teeth (C4(1 )); nasal vicariance alone is not a sufficient explanation for their scale undivided (C8(1 )); snout-vent length less than 35 present distribution. However, even non-vicariance hy­ cm (C9(2)*); ventral count does not exceed 160 potheses are testable in terms of congruence with (Cl0(3)); melanin pigment on ventral scales concen­ phylogenetic and distributional patterns in other line­ trated at posterolateral corners forming double ages, and in this case we are favoured by the strong saw-tooth pattern, or also extending medially as a con­ habitat-fidelity of Cacophis spp. We may assume sev­ tinuous dark border on each ventral (leaving base of eral cycles of interruption and reconnection of the each scale white or pale yellow), and. usually forming a eastern wet forest corridor, which have been frequent median zig-zag stripe under tail (Cl2(1)*); iris red during Plio-Pleistocene times (e.g. Bowler, 1982). One (Cl 7(0)*). possibility would involve a persistent northern popula­ tion expanding southward during three successive DISCUSSION periods of forestcontinuity (cool or moist periods, per­ The evidence for a sister-group relationship between haps the three maj or glaciations), and the southern Cacophis squamulosus and the remaining members of populations then differentiating after interruption of the the genus allows us the option of resurrecting forest corridor, giving rise to the three more divergent, Petrodymon Krefft, 1866 (cf.Wallach, 1985). However, sympatric, southern species (squamulosus, harriettae this would result in a monotypic (i.e. redundant) genus and krefftii sequentially; Fig. 6). unless populations currently assigned to C. squamulosus This model is simple in several senses: it invokes prove to belong to more than one species. Variation in only passive allopatric speciation (the preferred null hy­ ventral colour patterns within this species has been pothesis in, e.g., Brooks & McLennan, 1991 ), no noted above, and northern specimens tend to be slightly extinctions, and a known process of historical environ­ larger (Shine, l 980a), but no detailed investigation of mental change in (at this scale) an essentially geographic vanat10n has been made. The one-dimensional geographic space. The spatial asym­ autapomorphic modifications of defensive display and metry in the model (only southward range expansions) ventral pigmentation in C. squamulosus could be re­ provides a uniform explanation for the observed distri­ garded as adaptive mimicry of the sympatric Pseudechis butions based on the most parsimonious cladogram. porphyriacus . The difference in relative height of the Some other Australian elapids (Hemiaspis signata, display, and flattening of the head rather than the neck, Cryptophis nigrescens, Hop/ocephalus bitorquatus and do not contradictthis hypothesis, since C. squamulosus Trop idechis carinatus) have distributions comparable to thereby both displays its black-barred red belly to ad­ that of Ca cophis as a whole, interrupted by drier belts vantage, and reaches the height and head-width of a along the Queensland coast, but without evidence of 'Red-bellied Black' larger than itself. speciation. In H. signata, a northern form vagrans Within the (churchilli, harriettae, krefftii) clade, the Garman, 1901 is sometimes recognized as a subspecies characters used here indicate that the sister taxon to the or species, but no such distinction has been demon- CA COPHIS MORPHOLOGY AND PHYLOGENY 15

(Cadle & Gorman, 1981; Mao, Chen, Yin & Guo, 1983; Cacophls church/I/I Schwaner et al., 1985; Mengden, 1985). A view has de­ veloped that Demansia is only distantly related to other Australian taxa (Mengden, 1985; Shine, 1991; Greer, 1997), although this does not necessarily followfrom large genetic distances (phenetic data). Keogh et al. ( 1998) report DNA sequence evidence for a clade com­ prising Cacophis, Aspidomorphus, Demansia, Furina and Glyphodon, and some of the skeletal characters re­ ferred to above may also support such a group. These time results suggest that the earlier genetic studies were af­ fectedby accelerated genetic change in the whipsnake FIG. 6. Schematic distribution of Cacophis in space and lineage (as first suggested by Cadle & Gorman, 1981), time, according to phylogenetic hypothesis in Fig. 5 and and that other methods may have more success. speciation model proposed in text. Vertical axis represents latitude; present distribution of the genus is between 15° and Further studies will continue to improve understand­ 35°S, with C. churchilli north of 20° and other species ing of the adaptive radiation of Australasian elapids, and mainly or entirely further south. Horizontal axis represents external morphology, internal softanatomy, cranial and time, from before beginning of Pleistocene climatic axial skeletal morphology (including the fossil record), fluctuations (left)to the present day (right). Connectivity of genetic and molecular methods, and behavioural and 'overlapped' regions of branches is indicated by shading. ecological data can all contribute to this end.

ACKNOWLEDGEMENTS strated. The genera Hypsilurus (Agamidae) and Coeranoscincus (Scincidae) each have two well-differ­ thank Jeanette Covacevich and Patrick Couper entiated allopatric species with similar distributions to (QM), Allen Greer and Ross Sadlier (AMS), Mark Cacophis churchi/li and kreffiii respectively; Litoria Hutchinson (SAM), and Ken Aplin and the late Glenn xanthomera and L. ch/oris (Anura, Hylidae) form a Storr (WAM) for permission to examine specimens in similar vicariant pair, while patterns which may be com­ their care; Rick Shine, Sam McDowell, Greg Mengden, parable to Ca cophis as a whole are seen in Ca lyptotis, and Scott Keogh for sharing their interest in elapid lampropholis, Sap roscincus (Scincidae) and phylogeny, and Mike Archer, graduate students at Mixophyes (Myobatrachidae), each with five or more UNSW, and Mike Lee for discussions of method. species (Cogger, 1992, 2000; Barker, Grigg & Tyler, Thanks to Jeanette Covacevich (again) for discussions 1995). Among small, terrestrial forest mammals, of species problems in Cacophis and Furina, Jo Sumner Dasyurus maculatus, Antechinusjlavipes and A. stuartii for discussions of Cacophis, Karen Black for photogra­ (Marsupialia, Dasyuridae), Rattus fu scipes and R. phy of Demansia, Steve Richards for behavioural lutreolus (Rodentia, Muridae) also have breaks between observations on Aspidomorphus, and David Vermeesch southern and north-Queensland populations (subspe­ and Scott Heaydon for access to additional specimens. cies; Strahan, 1983 ). As relevant sequence data become Allen Greer and Mike Lee provided detailed comments available, the estimation of divergence dates using mo­ on earlier drafts. This work was begun during a PhD lecular 'clocks' may indicate whether Cacophis and studentship at the University of New South Wales, and other such lineages have been affected by the same se­ completed as a postdoctoral research associate at quence of environmental changes (undergoing evolution (sequentially) Monash University, the University of as a community), or have followed independent timeta­ Queensland and the South Australian Museum, with bles. Based on differences in the DNA sequences of two support fromthe AustralianResearch Council via grants genes, Keogh et al. (1998) infer a split between to Mike Lee. Cacophis squamulosus and krefftii 'of considerable an­ REFERENCES tiquity', but do not give a quantitative age estimate, and comparable sequences have not yet been reported for Aplin, K. P. & Donnellan, S. C. (1999). An extended the other species. description of the Pilbara Death Adder, Acanthophis The monophyly of Cacophis is now well supported, wellsi Haser (Serpentes: Elapidae), with notes on the and the results of this analysis will allow it to be treated Desert Death Adder, A. pyrrhus Boulenger, and as a unit in futurework. For example, the evidence pre­ identification of a possible hybrid zone. Records of the sented here may help to determine the relationships of Western Australian Museum 19, 277-298. Demansia with other lineages, which seems to be one of Auffenberg, W. ( 1963). The fossil snakes of Florida. the centralproblems of elapid phylogeny. Morphologi­ Tu lane Studies in Zoology 10, 131-216. cal analyses have associated Demansia with Barker, J., Grigg, G. C. & Tyler, M. J. (1995). A Field Guide Aspidomorphus and members of the viviparous lineage to Australian Frogs. Sydney: Surrey Beatty & Sons. (McDowell, 1967, 1969b, 1985), or with Pseudechis Bechtel, H. B. (1978). Col or and pattern in snakes (Wallach, 1985), while most genetic studies have failed (Reptilia, Serpentes). Journal of Herpetology 12, 52 1- to show close relationships with any Australian lineages 32. 16 J. D. SCANLON

Blanchard, F. ( 1931). Secondary sex characters of certain Gillam, M. W. (1979). The genus Pseudonaja (Serpentes: snakes. Bulletin of the Antivenin Institute of America Elapidae) in the Northern Territory. Territory Parks 4, 95-104. and Wildlife Research Bulletin 1, 1-28. Bogert, C. M. (1943). Dentitional phenomena in cobras Gow, G. F. (1989). Complete Guide to Australian Snakes. and other elapids with notes on adaptive modifications Sydney: Angus & Robertson. of fangs. Bulletin of the American Museum of Natural Greer, A. E. (1993). Lineage-associated asymmetries in History 81, 285-360. scale overlap patterns in squamates. Herpetologica 49, Boulenger, G. A. (1896). Catalogue of the snakes in the 318-326. British Museum (Natural History) , Vol. Ill. London: Greer, A. E. ( 1997). Th e Biology and Evolution of Taylor & Francis Ltd. Australian Snakes. Sydney: Surrey Beatty & Sons. Bowler, J. M. ( 1982). Aridity in the late Tertiary and Greer, A. E. & Cogger, H. G. (1985). Systematics of the Quaternary of Australia. In Evolution of the Flora and reduce-limbed and limbless skinks currently assigned Fa una of Arid Australia, 35-45. Barker, W. R. & to the genus Anomalopus (Lacertilia: Scincidae). Greens lade, P. J. M. (Eds). Adelaide: Peacock Records of the Australian Museum 37, 1 1-54. Publications. Grigg, G., Shine, R. & Ehmann, H. (Eds.). (1985). Biology Bremer, K. ( 1988). The limits of amino-acid sequence data of Australasian Frogs and Reptiles. Sydney: Royal in angiosperm phylogenetic reconstruction. Evolution Zoological Society of New South Wales/Surrey Beatty 42, 795-803. & Sons. Brongersma, L. D. ( 1934). Contributions to Indo­ Gunther, A. ( 1863 ). Third account of new species of Australian herpetology. Zoologische Mededelingen , snakes in the collection of the British Museum. Annals Leyden 17, 161-251. and Magazine of Natural History (3) 12, 348-365. Brooks, D. R. & McLennan, D. A. (1991 ). Phylogeny, Gyi, K. K. (1970). A revision of colubrid snakes of the Ecology, and Behavior: A Research Program in Subfamily Homalopsinae. University of Kansas, Comparative Biology. Chicago & London: University Publications of the Museum of Natural History 20, of Chicago Press. 511-625. Burger, W. L. & Natsuno, T. (1974). A new genus for the Horner,P. ( 1998). Simoselaps morrisi sp. nov. (Elapidae), Arafura smooth seasnake and redefinitions of other a new species of snake from the Northern Territory. seasnake genera. Th e Snake 6, 61-75. The Beagle, Records of the Museums and Art Galleries Cadle, J. E. & Gorman, G. C. ( 1981). Albumin of the No rthern Te rritory 14, 63-70. immunological evidence and the relationships of sea Hoser, R. T. ( 1989). Australian Reptiles and Frogs. snakes. Journal of Herpetology 15, 329-334. Sydney: Pierson & Co. Cogger, H. G. (1975). Reptiles and Amphibians of Hutchinson, G. E. ( 1959). Homage to Santa Rosalia or why Australia. Sydney: A. H & A. W Reed. are there so many kinds of animals? American Cogger, H. G. (1992). Reptiles and Amphibians of Naturalist 93, 145-159. Australia. 5th Edition. Sydney: Reed Books. Hutchinson, M. N. (1990). The generic classification of Cogger, H. G. (2000). Reptiles and Amphibians of the Australian terrestrial elapid snakes. Memoirs of the Australia. 6th Edition. Sydney: Reed New Holland. Queensland Museum 29, 397-405. Cogger, H. G., Cameron, E. E. & Cogger, H. M. (1983). Keogh, J. S. ( 1998). Molecular phylogeny of elapid snakes Amphibia and Reptilia. Zoological Catalogue of and a consideration of their biogeographic history. Australia, Vol. I. Canberra: Australian Government Biological Journalof the Linnean Society 63, 177-203. Publishing Service. Keogh, J. S. ( 1999). Evolutionary implications of Cundall, D. ( 1995). Feeding behaviour in Cy lindrophis hemipenial morphology in the terrestrial Australian and its bearing on the evolution of alethinophidian elapid snakes. Zoological Journal of the linnean snakes. Journal of Zoology, London 237, 353-376. Society 125, 232-278. Cundall, D. & Sharda, J. D. (1995). Rhinokinetic snout of Keogh, J. S., Scott, I. & Scanlon, J. D. (2000). Molecular thamnophiine snakes. Jo urnal of Morphology 225, 31-50. phylogeny of viviparous Australian elapid snakes: Dumeril, A. M. C., Bibron, G. & Dumeril, A. H. A. Affinities of 'Echiopsis ' atriceps (Storr, 1980) and ( 1854 ). Erpetologie generale ou Histoire nature/le 'Drysdalia ' corona/a (Schlegel, I 83 7), with complete des Reptiles. Vol. 7, pt I. xvi + 780 pp. description of a new genus. Journal of Zoology, Paris: Librairie Encyclopedique de Roret. London 252, 317-326. Ehmann, H. (1992). Encyc/opedia of Australian Animals. Keogh, J. S., Shine, R. & Donnellan, S. (1998). Reptiles. Sydney: Angus & Robertson. Phylogenetic relationships of terrestrial Australo­ Fox, D. L. (1953). Animal Biochromes and Structural Papuan elapid snakes (subfamily Hydrophiinae) based Colours. Cambridge: The University Press. on cytochrome b and I 6S rRNA sequences. Molecular Garman, S. ( 1901 ). Some reptiles and batrachians from Phylogenetics and Evolution 10, 67-81. Australasia. Bulletin of the Museum of Comparative Kinghorn, J. R. ( 193 9). Two Queensland snakes. Records Zoology 39, 1-14. of the Australian Museum 20, 257-260. CA COPHIS MORPHOLOGYAND PHYLOGENY 17

Krefft, G. (1866). On snakes observed in the 'Mengden, G. A. (1983). The taxonomy of Australian neighbourhood of Sydney. Transactions of the elapid snakes: a review. Records of the Australian Philosophical Society of NSW 1862-65, 34-60. Museum 35, 195-222. Krefft, G. (1869). Th e Snakes of Australia: an illustrated Mengden, G. A. (1985). Australian elapid phylogeny: a and descriptive catalogue of all the known sp ecies. summary of the chromosomal and electrophoretic data. Sydney: Thomas Richards, Government Printer: In Biology of Australasian Frogs and Reptiles, 185- (Facsimile edition, 1984. Brisbane: Lookout 192. Grigg, G., Shine, R. & Ehmann, H. (Eds). Publications) Sydney: Royal Zoological Society of New South Lee, M. S. Y. ( 1997). Phylogenetic relationships among Wales/Surrey Beatty & Sons. Australian elapid snakes: the soft anatomical data Mertens, R. ( 1936). Ober liuf3ere Geschlechts-Merkmale reconsidered. Herpetological Journal 7, 93-102. einiger Schlangen. Senckenbergiana 19, 169-174. Lee, M. S. Y. (1998). Convergent evolution and character O'Shea; M. (1996). A Guide to the Snakes of Pap ua New correlation in burrowing reptiles: towards a resolution Guinea. Port Moresby: Independent Publishing. of squamate phylogeny. Biological Journal of the Queensland Museum. (2000). Wildlife of Tropical North Linnean Society 65, 369-453. Queensland. Brisbane: Queensland Museum. Longmore, R. (1986). Atlas of elapid snakes of Australia. Rasmussen, A. R. (2002). Phylogenetic analysis of the Australian Flora and Fauna Series (7). Canberra: "true" aquatic elapid snakes Hydrophiinae (sensu Bureau of Flora and Fauna. Smith et al., 1977) indicates two independent Maddison, W. P., Donoghue, M. J. & Maddison, D. R. radiations into water. Steenstrupia 27, 47-63. ( 1984 ) . Outgroup analysis and parsimony. Systematic Resetar, A. R. & Marx, H. (1981). A redescription and Zoology 33, 83-103. generic reallocation of the African colubrid snake Maddison, W. P. & Maddison, D. R. (2000). MacClade Elapocalamus gracilis Boulenger with a discussion of 4.0. Sunderland, Massachusetts: Sinauer Associates. the union of the brille and postocular shield. Journal of Mao, S. H., Chen, B. Y., Yin, F. Y. & Guo, Y. W. (1983). Herpetology 15, 83-89. lmmunotaxonomic relationships of sea snakes and Roze, J. A. (1996). Coral Snakes of the Americas: Biology, terrestrial elapids. Comparative Biochemistry and Identification, and Venoms. Malabar, Florida: Krieger Physiology A 74, 869-872. Publishing Co. McDowell, S. B. (1967). Aspidomorphus, a genus of New Savitzky, A. H. (1983) . Coadapted character complexes Guinea snakes of the family Elapidae, with notes on among snakes: fossoriality, piscivory, and durophagy. related genera. Journal of Zoology, London 151, 497-543. American Zoologist 23, 397-409. McDowell, S. B. (J 969a). Toxicocalamus, a New Guinea Scanlon, J. D. (1985). Phylogeny and relationships of the genus of snakes of the family Elapidae. Journal of elapid snake genus Simoselaps Jan, 1859: the Zoology, London 159, 443-5 1 1. evolution of a group of burrowing snakes. (BSc McDowell, S.B. (J969b). Notes on the Australian sea­ Honours thesis: University of Sydney). snake Ep halophis greyii M. Smith (Serpentes: Scanlon, J. D. (1988). Phylogeny and adaptation in Si­ Elapidae: Hydrophiinae) and the origin and moselaps and related genera (Serpentes: Elapidae: classification of sea-snakes. Zoological Journal of the Hydropheinae). Australian Bicentennial Herpetologi­ Linnean Society, London 48, 333-349. cal Conference, Abstracts. (Queensland Museum: McDowell, S. B. ( 1970). On the status and relationships Brisbane). of the Solomon Island elapid snakes. Journal of Scanlon, J. D. (1995). An elapid snake (Hydropheinae) Zoology, London 161, 145 -190. from the Middle Miocene Encore Site, Riversleigh. McDowell, S. 8. ( 1972). The genera of sea-snakes of the 5th Conference on Australasian Vertebrate Evolution, Hydrophis group (Serpentes: Elapidae). Transactions Palaeontology and Systematics, Abstracts, 20. of the Zoological Society of London 32, 1 89-247. Scanlon, J. D. (1996). Studies in the palaeontology and McDowell, S. 8. (1974). Additional notes on the rare and systematics of Australian snakes. (PhD thesis, primitive sea-snake, Ep halophis greyii. Jo urnal of University of New South Wales: Sydney). Herpetology 8, 123-128. Scanlon, J. D. (1998). Prey-scaring by visual pursuit McDowell, S. B .. (1985). The terrestrial Australian elapids: predators: a new use for tail-waving in snakes. general summary. In Biology of Australasian Frogs Herpetofauna 28(2), 5-10. and Reptiles, 26 1 -264. Grigg, G., Shine, R. & Schwaner, T. D., Baverstock, P. R., Dessauer, H. C. & Ehmann, H. (Eds). Sydney: Royal Zoological Society Mengden, G. A. (1985). Immunological evidence for of New South Wales/Surrey Beatty & Sons. the phylogenetic relationships of Australian elapid McDowell, S. B. (1987). Systematics. In Snakes: snakes. In Biology of Australasian Frogs and Reptiles, Ecology and Evolutionary Biology, 1-50. Seigel, R. 177- 1 84. Grigg, G., Shine, R. & Ehmann, H. (Eds). A., Collins, J. T. C. & Novak, S. S. (Eds). New York: Sydney: Royal Zoological Society of New South MacMillan. Wales/Surrey Beatty & Sons. 18 J. D. SCANLON

Shea, G. M. & Sadlier, R. A. (1999). A catalogue of the Storr, G. M., Smith, L. A. & Johnstone, R. E. (1986). non-fossil amphibian and reptile type specimens in the Snakes of Western Australia. Perth: Western collection of the Australian Museum: types currently, Australian Museum. previously and purportedly present. Technical Reports Strahan, R. (Ed.) (1983). Th e Australian Museum of the Australian Museum (I 5 ), 1-91. Complete Book of Australian Mammals. Sydney: Shine, R. (I98 0a). Comparative ecology of three Angus & Robertson. Australian snake species of the genus Cacophis Sweet, S. S. (1980). Allometric inference in morphology. (Serpentes: Elapidae). Copeia 1980, 83 I -838. American Zoologist 20, 643-652. Shine, R. (I980b). Ecology of eastern Australian Swofford, D. L. (2002). PAUP*. Phylogenetic Analysis wh ipsnakes of the genus Demansia. Jo urnal of Using Parsimony (*and Other Methods). Version 4. Herpetology 14, 381-389. Sunderland, Massachusetts: Sinauer Associates. Shine, R. (I 98 1 ). Ecology of Australian elapid snakes of Underwood, G. (1967). A Contribution to the the genera Purina and Glyphodon. Jo urnal of Classification of Snakes. London: British Museum Herpetology J 5, 219-224. (Natural History). Shine, R. (1985). Ecological evidence on the phylogeny of Wallach, V. ( 1985). A cladistic analysis of the terrestrial Australian elapid snakes. In Biology of Australasian Australian Elapidae. In Biology of Australasian Frogs Frogs and Reptiles, 255-260. Grigg, G., Shine, R. & and Reptiles, 223-253. Grigg, G., Shine, R. & Ehmann, H. (Eds). Sydney: Royal Zoological Society Ehmann, H. (Eds). Sydney: Royal Zoological Society of New South Wales I Surrey Beatty & Sons. of New South Wales/Surrey Beatty & Sons. Shine, R. ( 1991 ). Australian Snakes: a natural history. Wells, R. (I 980). Notes on Krefft's Dwarf snake Sydney: Reed Books. ( Cacophis krefltii). Herpetofauna J J, 18- I 9. Shine, R. & Keogh, J. S. (1996). Food habits and Wells, R. W. & Wellington, C. R. ( 1985). A classification reproductive biology of the endemic Melanesian of the Amphibia and Reptilia of Australia. Australian elapids: are tropical snakes really different? Journal of Jo urnalof Herpetology (Supplement Series) J, 1-6 I. Herpetology 30, 238-247. Wiens, J. J. (1998). The accuracy of methods for coding Slowinski, J. B. & Keogh, J. S. (2000). Phylogenetic, and sampling higher-level taxa for phylogenetic relationships of elapid snakes based on Cytochrome b analysis: a simulation study. Systematic Biology 47: mtDNA sequences. Molecular Phylogenetics and 397-4 13. Evolution 15, 157- 164. Wilson, S. K. & Knowles, D. G. (1988). Australia 's Sorenson, M. D. ( 1999). Tree Rot Version 2. Distributed Reptiles. Sydney: Collins. by the author, Boston University. Worrell, E. ( 1955). A new elapine snake from Queensland. Storr, G. M. (I 968). The genus Vermicella (Serpentes: Proceedings of the Royal Zoological Society of New Elapidae) in Western Australia and Northern Territory. South Wa les 1953-54, 41-43. Jo urnal and Proceedings of the Royal Society of Worrell, E. ( 1963). Reptiles of Australia. Sydney: Angus Western Australia 50, 80-92. & Robertson. Storr, G. M. (1978). Whipsnakes (Demansia : Elapidae) Worrell, E. (1970). Reptiles of Australia. 2nd Edition. of Western Australia. Records of the Western Sydney: Angus & Robertson. Australian Museum 6, 287-301. Storr, G. M. (I98 1 ). The genus Purina (Serpentes: Elapidae) in Western Australia. Records of the Western Australian Museum. 9, I I 9-123. Storr, G. M. (1985). Phylogenetic relationships of Australian elapid snakes: external morphology with an emphasis on species in Western Australia. In Biology of Australasian Frogs and Reptiles, 221 -222. Grigg, G., Shine, R. & Ehmann, H. (Eds). Sydney: Royal Zoological Society of New South Wales/Surrey Beatty & Sons. Accepted: 22. 8. 02 19

APPENDIX l Glyphodon dunmalli: QM 123 178*; G. tristis : SAM R13998*, MV unreg.*; Boulenger (1896), Worrell Sources of data for ingroup and outgroup species. (1955), Shine (1981) , Scanlon (unpublished data). Skeletal material examined marked with asterisk (*); specimens examined only for external characters omit­ ted for taxa other than Cacophis spp. and Demansia APPENDIX 2 simplex (available from author on request). Useful List of characters used in the phylogenetic analysis. sources of further data and illustrations for Australian species include Cogger ( 1975, 1992, 2000), Storret al. Autapomorphies ofCacophis. See Table 2 for distri­ (I 986), Wilson & Knowles (1988), Gow ( 1989), Hoser A. bution of states in outgroup genera. (I 989), Ehmann (1992), and Greer ( 1997).

I. Paired parietal foramina: present, at least in small Aspidomorphus lineaticollis: AMS R125021 * '. specimens (O); normally absent (1). AMNH 42376*; A. muelleri: AMS R16614* R19013* 2. Palatine choanal process: absent (O); present but low' A. sch/egelii : MCZ 73 11* (AMNH and MCZ speci� i.e. 'short' (1); higher than long, i.e. 'tall' (2). mens examined by M. Lee); Brmrnersma (1934), 3. Ectopterygoid anterior extent: anterior to palatine­ McDowell (1967), Shine and Keogh (1996), O'Shea pterygoid joint (O); approximately level with (lateral ( 1 996). to) joint (1 ); entirely posterior to palatine (2). Cacophis churchilli: AMS R6489, RI0732, Rl 1340, 4. Supra-anal keels: lateral scales of cloacal region simi­ Rl 1362, RJ 1506, Rl 1512, R12480, R12482, R12914, lar in gross morphology to those of rest of body (O); R17035, R20207, R53726, R63 163, R63836, patch of keeled lateral scales present in adult males QM13640, 14295, 15 193, 15292, 15296, J5720, 15722, (I). 15723, 15724, 15725, 15954, J7339, J13674, 121 206, 5. Parietal and lower postocular: separated by contact of 124539, 153282*; SAM R22392, JS63*. upper postocular with anterior temporal (O); some­ Cacopllis harriettae: AMS R648, R5844, R6182, times in contact, separating upper postocular from R8507, R9399, Rll687, R12734, R12854, R13027, temporal (1 ) . Rl3701, R13716, Rl8277, R4 7545, R86763, R88468, 6. Temporals: 2+2+3, three distinct rows (O); 1=1+3 (or QM 14443*, 120278*, J26544*, J46288*, 147658*, I=I+ 2), single large anterior temporal incorporating 147982*, 150600*, SAM R26989*; McDowell (1967), temporolabial(I) . Shine (I980a). 7. Iris colour in preservative (melanin): entirely or Cacophis krefftii: AMS R877 (x2), Rl502, R2303, . mamly dark (O); pale, with at most dark flecks or faint R24 J 1, R4 1J8, R6545, R7492, R8026, R8768, R9 198, variegation (1). Rl0013, Rl 1812, R12356, R12738, R12919, R12995, R13000, R1300 1, R13064, RJ3662 (x2), R13743, B. Characters possibly derived within Australasian R13799, RJ3823, R13824, R13869, R13982, RJ4344, elapids but shared by Cacophis with both outgroup R14359 (x2), R14422, R14815, R15690, R17917, clades. See Table 2 for distribution of states in R18482, R47474, R58486, R75961, R74466, R77370, outgroup genera. R8 1158, R81 159, R8 1160, R8 1161, R86762, R86797, R90609, R97275, R106956, RJ 10341, Rl 14956, 1. Prefrontal and postorbital bones: widely separated R1254 10, AMS unreg.*, QM J966, J14287, J32725, and frontal broadly entering orbital margin (O); pre­ J34031, 146583*, SAM R26974*; McDowell (1967), frontal and postorbital approach or meet, effectively Shine {1980a), Wells (1980). . excludmg frontal from margin ( 1). Cacophis squamulosus: AMS R28232, R29733, 2. Interolfactory pillars offronta ls: distinctly constricted R30336, R37 187, R4 1801, R47471, R47544, R47546, (O); as wide as the septomaxillary-frontal contact R4 7779, R48108, R50220, R52964, R627 10, R64975, widely separating olfactory openings of frontal ( J ) . ' AMS unreg. (x4)*, QM147659*, 147976*, J47983*, 3. Maxilla anterior process: short and blunt (O); promi­ SAM R2263A*, 1S3*, JS14*; Shine (1980a). nent or acute in ventral view (I). Demansia psammophis: QMJ7 134*, 126907*, 4. Maxilla posterior extent, relative to postorbital in lat­ 146291 *, 147978*, AMS R- 1 3-672*, JS 44*; D. eral view: short, not beyond orbit (O); long, beyond vestigiata : AMS R- 13-667*; D . . sp. cf. olivacea : posterior margin of orbit as definedby the postorbital JS 169*; D. sp. cf. torquata : QMJ46289*, SAM (I). R20483*; D. simplex : AMS RJ3045, R13046, 5. Coronoid eminence of mandible: absent, dorsal mar­ R13702, RJ4030, R14029, R 128403, R128404, NTM gin of compound smoothly curved (O); eminence R18625*; Storr (1978), Shine (1980b), Shea and present as slight to strongconvex angulation of dorsal Scanlon (unpublished data). edge of surangular ( l ). Fu ri11 a barnardi: SAM R27022*, AMS unreg.*; F. 6. Hypapophysis shape in posterior trunk vertebrae: diadema : AMS R98 165*, SAM R6075*, R6703*, smoothly sigmoid in lateral view (O); some vertebrae 1S32*; ornata : WAM Rl 5088*; Shine ( 1981 ), Storr F. with a distinct horizontal portion defined by an (1981), Scanlon (1985, unpublished data). anteroventral angle (I). 20 J. D. SCANLON

7. Dentary teeth: uniform or with smooth gradient of 8. Nasal scale: divided in maj ority (O); single in maj ority

size (O); distinctly larger anteriorly (1) . {I). 8. Dentary tooth row: lacks a diastema (O); diastema 9. Maximum snout-vent length: greater than 70 cm (O); commonly present behind enlarged teeth (1 ). 40-65 cm {I); less than 35 cm (2). Ordered 0- 1-2. 9. Number of midbody scale rows: 19 or more (O); 17 10. Ventral scale number (range): 176-200 (0); 170-175 (l); 15-17, intraspecifically variable (2); 15 (3). Po­ (1); 165-169 (2); 161-164 (3); 140-160 (4). Ordered. larity follows Wallach (1985), but state 0 is almost 11. Subcaudal scale number: 41-50 (O); 38-40 ( 1 ); 30-37 certainly derived within Glyphodon (21 rows in G. (2); 25-29 (3). Ordered 0- 1-2-3. dunma/li). 12. Ventral melanin pigment: uniformly distributed, 10. Posterior scale-row reduction: one or more reduc­ 'peppered' or generally dark grey (O); strongly con­ tions always present (O); variable, reduction trasting pattern, usually transverse dark and light sometimes present (1); reduction rare or absent (2). bands on each ventral scale ( 1 ). 13. Upper light line or nape band: pale lines on neck lon­ C. Characters varying within Cacophis. See Table 3 for gitudinal, separated across midline (O); transverse and distribution of states in Cacophis species and connected across midline, forming a complete collar outgroup genera. {I). 14. Upper light line or nape band: occupies at least 4 1. Maxillary teeth posterior to fangs: 6-8 alveoli (O); 2-5 transverse scale rows (0), or 1-2 only (I). (I). 15. Carotenoid on body: pale yellow or absent (O); or­ 2. Palatine teeth: usually more than 11 alveoli (O); 11 or ange to red ( 1 ). fewer (I). 16. Carotenoid on face and nape: yellow (O); very pale

3. Pterygoid teeth: 19-24 alveoli (0): 11-18 (I). yellow or white (1 ) . 4. Dentary teeth: 21 or more (O); less than 20 (I). 17. Iris colour in life(non-melani n): red (O); yellow ( 1 ); 5. Nasal and preocular: usually in contact (O); usually partly or mainly white (2). Ordered 0- 1 -2. separated, contact rare ( 1 ); normally widely sepa­ 18. Pupil shape: vertically elliptical (O); weakly oval or rated, contact not observed (2). Ordered 0-1-2. round (I). 6. Posterior temporals: nearly always three in fi nal row 19. Forebody in high defensive display: held straight

(2+2+3 or 1=1+3) (O); reduced to two in most indi­ (O); lateral curves ( 1 ) . viduals (1=1+2) (I). 7. Parietal-postocular contact, frequency: minority (O); majority ( 1 ).