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Herpetology Notes, volume 10: 209-217 (2017) (published online on 25 April 2017)

A new colour morph of bibroni (: ) and evidence for Müllerian in Tropical Indian coralsnakes

Dileep Kumar Raveendran1, V. Deepak2, Eric N. Smith3 and Utpal Smart3,*

Abstract. Meristic and molecular data provide evidence for an exceptional multi-chromatic defensive strategy in an Indian coralsnake, Calliophis bibroni from the state of Kerala. We propose a mimicry hypothesis involving a combination of an ontogenetic colour shift at maturity, from initial Müllerian mimicry with a subtropical Indian coralsnake Sinomicrurus macclellandii, to one of two very different adult dorsal colours: 1) an aposematic pattern resembling that of the sympatric tropical Indian coralsnake Calliophis castoe or 2) a cryptic dark brown colouration. To this end, we succinctly juxtapose the rich body of work on mimicry in elapids to that of their counterparts in an attempt to address the exciting yet unexplored prospect of investigating mimicry, crypsis and in Old World coralsnakes.

Key Words. Aposematism, crypsis, Indian coralsnakes, genetic distance, meristics, mimicry

Introduction Müllerian co-mimics benefit from sharing the same signal since this reduces the number of individuals that colouration provides many functions; the most have to be sacrificed per prey to educate local important among them is probably predator avoidance predators of a given aposematic colouration (Müller, and deterrence. Camouflage, or crypsis, blends 1879). Furthermore, possibly to balance prey-predator into their environment while deimatism surprises and dynamics, some mimics may replicate the colours confuses predators with the display of startling colour or patterns of dangerous models which lack bright, patterns. Taxonomically and geographical widespread, conspicuous colours and are cryptic instead (Wüster et aposematism or warning colouration is the display of al., 2004, McElroy, 2016). noxious defense by the means of cautionary colours, The majority of work on Batesian and Müllerian to deter predatory attacks. Batesian mimicry occurs mimicry complexes has been carried out on arthropods when a non-venomous or palatable species mimics such as the toxic Heliconius butterflies (e.g., Baxter et the colour and pattern of a dangerous or unpalatable al., 2008, Nadeau et al., 2014, Kronforst and Papa, 2015). species (Poulton, 1890), whereas Müllerian mimicry While it is true that mimicry is far more taxonomically is when a venomous, toxic or distasteful species, widespread in invertebrates (Pough, 1988), amongst mimics other organisms with similar disagreeable traits. vertebrates (alongside Poison frogs of the family Dendrobatidae [Yeager et al. 2012]) venomous have great potential to serve as model organism since many species are known to show striking crypsis, 1 Centre for Venom Informatics, University of Kerala, disruptive colouration and aposematism (Kikuchi et Thiruvananthapuram, Kerala, al., 2014). In effect, close to 35% of non-venomous 2 Centre for Ecological Sciences, Indian Institute of Science, species worldwide are thought to be mimics of Bengaluru, India venomous viperids (vipers and asps) and/or elapids 3 Amphibian and Diversity Research Center and Department of Biology, University of at Arlington, (cobras, kraits, coralsnakes and sea snakes) (Savage Arlington, Texas, USA and Slowinski 1992, Greene and McDiarmid, 1981). * Corresponding author e-mail: [email protected] Information on mimetic resemblances in vipers thus 210 Dileep Kumar Raveendran et al. far has lacked experimental rigor, most likely because (Reinhardt, 1844) until they attain an SVL of >400 mm. vipers lack distinctive patterns that would enable a As adults they transition to one of two phenotypes 1) quantitative approach (Pough 1988). Amongst elapids, a dorsal colouration that is a cryptic glossy bistre with the New World coralsnakes ( and ) bands of the same colour running down the flanks (plain have been amongst the earliest subjects for inquiries morph), or 2) a dorsal colouration of an iridescent bole on mimicry given their beautiful aposematic colours, brown without any dark bands on the flanks, resembling usually consisting of red, yellow and black bands (e.g. the venomous Castoe’s coralsnake C. castoe (Smith, Cope, 1860). Ever since highlighted by Wallace (1867) Ogale, Deepak and Giri, 2012). The ventral pattern as exceptional systems, numerous experimental as well retains a vivid flesh coloured red hue throughout this as qualitative studies have dealt with aposematism and transition in both cases. The complex set of chromatic mimicry in these coralsnakes (Gehlbach 1972, Smith defensive strategies described in this paper have not 1975, 1977, 1980, Greene and McDiarmid 2005, 1981, yet been reported in any other elapid, not even among Brodie 1993, Rabosky et al. 2016). Though fervently the New World pattern-diverse snakes of the contested (e.g. Gadow, 1911; Brattstrom, 1955), in Micrurus. general both empirical and experimental evidence suggests that chromatic patterns of New World Methods coralsnakes are aposematic and that they indeed serve as mimicry models for several harmless or semi-venomous Species and study area.— Calliophis is a genus of colubrid snakes; prominent examples include some Asian coralsnakes comprising ten species, distributed (Lampropeltis) and various false coralsnakes throughout India and Southeast Asia (Castoe et al., 2007; Pliocerus and ; (Dunn, 1954; Campbell Smith et al., 2008, 2012). Four out of the five species of and Lamar, 1989; Brodie and Janzen, 1995). In addition Calliophis in India have their distributions largely across to this, New World coralsnakes are also known to have the Western Ghats, a mountain range that runs almost several plausible instances of Müllerian mimicry within parallel to the western coast of the Indian peninsula themselves (Greene and McDiarmid, 1981, 2005). (Smith, 1943; Whitaker et al., 2004). One of these, Reports of mimicry in Asian elapids, on the other Bibron’s coralsnake or C. bibroni is distributed along hand, have been largely speculative. Several instances a wide altitudinal range (0–1220 m) in the evergreen include Batesian mimicry between colubrid mimics and forests in and around the Western Ghats (Deepak et coralsnake models, such as subtropical Asian coralsnakes al., 2010). Overall, there is limited information on of the genus Sinomicrurus and their putative snake the distribution, natural history and ecology of this mimics in the genus Oligodon or Kukri snakes, and long endangered species (Deepak et al., 2010; Srinivasulu, glanded tropical coralsnakes and their possible reed Srinivasulu and Molur 2014). To our knowledge the snake mimics in Calamaria (Greene and McDiarmid, first photos published of live specimens were by Sharma 2005). Additionally, the occurrence of metallic dark (1988, as C. macclellandi). Gowri Shankar and Ganesh blue or black bodies with bright red heads and tails on (2009) reported on two female individuals representing both the Malayan Blue coralsnake Calliophis bivirgata the northernmost localities known for the species and (Boie, 1827), and the sympatric Red-headed Krait presented notes on ecology and salient features of Bungarus flaviceps (Reinhardt, 1843) has been cited external morphology of the species. C. bibroni was later as an example of Müllerian mimicry in Asian elapids re-described by Deepak et al., (2010) who also provided (Slowinski, 1994; Greene and McDiarmid, 2005). records from south-west peninsular India and presented Here we report the first observational evidence of a additional information on the natural history, , complex chromatic defensive strategy in a tropical Old morphological variation and ontogenetic shift in colour World elapid, Bibron’s coralsnake Calliophis bibroni pattern. The authors brought to light the very unique (Jan, 1858), found in and along the Western Ghats of condition, in which sub-adult (<400 mm snout vent India. This species is unique in that it undergoes an length) C. bibroni tend to be aposematic and disruptive ontogenetic colour alteration that may correspond to overall but as adults these become cryptic above while a shift in Müllerian mimicry while simultaneously remaining aposematic and disruptive below. This representing a transition from aposematism to crypsis: ontogenetic shift in colour pattern had not been reported juvenile snakes with a aposematic flesh coloured in any other species of Old or New World coralsnake. dorsum and black bands possibly mimic the venomous Castoe’s coralsnake Calliophis castoe, described by MacClelland’s coralsnake Sinomicrurus macclellandii Smith et al., (2012), is found in tropical semi-evergreen A new colour morph of Calliophis bibroni 211

forest and tropical wet evergreen forest on the western 96-Well Thermal cycler (Applied Biosystems, coast of India. The altitudinal range for C. castoe is ). Post amplification, we resolved the PCR from near sea level (ca. 10–15 m) in Goa and Karnataka products on 1.5% agarose gel and eluted using GeneJET to about 715 m in southern Maharashtra (Smith et al., Gel extraction Kit (Thermo Scientific, Massachusetts, 2012). USA). We sequenced the amplified inner PCR products An individual, initially identified as Calliophis castoe, on an ABI3730xL DNA analyzer (Applied Biosystems, was collected by the first author, DKR, on October California, USA), using the ABI PRISM BigDye the 12th at 0840 hours from Valapattanam, Kannur Terminator Cycle sequencing Kit (Perkin-Elmer Corp., District,������������������������������������������������ � USA), following the manufacturer’s instructions. brought to the attention of the authors that, despite an We cleaned the raw sequences and assembled almost exact resemblance in colour pattern to an adult consensus sequences for the gene fragments using the C castoe, the scale counts of the head and caudal area program Sequencher 4.8 (Gene Codes, Ann Arbor, corresponded strongly to meristic patterns typical to Michigan, USA). We built the alignment of the resultant C. bibroni. We therefore used molecular data from the consensus sequences using MEGA v5.2.1 (Tamura et collected specimen and compared sequence divergence al., 2011). We edited sequences by eye for accuracy and between it and several individuals of C. bibroni, and also translated them to amino acid sequences, to verify one individual of C. castoe, in order to establish its true the absence of stop codons. We detected no internal identity. stop codons and we deposited the new sequences in GenBank (Table 1). We calculated uncorrected pairwise Molecular data.—The molecular work for this study p-distances in MEGA v5.2.1 (Tamura et al., 2011). was carried out in three different laboratories in India (viz. the Centre for Venom Informatics, Evolving Phylo- Coloration and meristic data.— We took measurements lab and the Aggarwal Lab). We isolated genomic DNA of external morphology from digital images using the from six individuals (one Calliophis castoe, four C. software ImageJ (Rasband, 2004). We took photographs bibroni and the individual in question) from liver and/ with high-resolution digital cameras (> 8 megapixels), or muscle tissue (preserved in 90% ethanol) using the placing the subject at right angles with respect to the Qiagen DNeasy Kit (Qiagen, Valencia, CA, USA). The lens of the camera. We measured snout-vent length following section describes the protocol followed by the (SVL), tail length, and total length (TL) to the nearest Centre for Venom Informatics; for protocols followed mm using ImageJ or a measuring ruler or tape. We by the Evolving Phylo-Lab and the Aggarwal lab see followed standard colubroid terminology for scales (e.g. Smith (2012) and (Dutta et al., 2004) respectively. We Smith and Campbell, 1994); for counting ventrals we amplified the mitochondrial gene Cytb via PCR using followed the method of Dowling (1951). We excluded the primers GLUDGE and ATRCB3 and according to the terminal scute (tip) from the number of subcaudals. thermocycler protocol used by Castoe et al., (2007). We We counted the numbers of dorsal scale rows at one used Dream Taq™ Green PCR Master Mix, 2X (Thermo head length behind the head, at midbody and at one head

1 Scientific, Massachusetts, USA) for all amplifications length before the vent. We determined sex by observing reactions. We performed thermal cycling on a Veriti™ presence or absence of hemipenes, through dissection 2 Table 1. List of specimens used in the molecular analysis, with collection locality, sample number, and GenBank accession number.

Table 1. List of Calliophis specimens used in the molecular analysis, with collection locality, sample number, and GenBank accession number.

Species Sample # Locality Latitude Longitude GenBank Accession # C. bibroni M816 Bethary, Waynad, Kerala 11° 41' 7.4394" N 76° 7' 55.2" E KX573691 C. bibroni M817 Palode, Trivandrum, Kerala 8° 31' 26.76" N 76° 56' 11.7594" E KX573692 C. sp. M819 Valapattanam, Kannur, Kerala 11° 54' 0" N 75° 22' 12" E KX573693 C. bibroni M731 Topslip, Coimbatore, Tamil Nadu 10° 27' 51.1194" N 76° 50' 12.4794" E KX573694 C. bibroni M733 Topslip, Coimbatore, Tamil Nadu 10° 16' 53.3994" N 76° 30' 3.9594" E KX573695 C. castoe M706 Ambe Ghat, South Goa, Goa 15° 3' 50.4" N 74° 9' 56.8074" E KX573696

1

212 Dileep Kumar Raveendran et al. of the base of the tail. The colour descriptions of the Discussion referred specimen in life are based on electronic images. The chromatic resemblance between the new colour We deposited these images at the Digital Collection of morph of Calliophis bibroni and C. castoe is remarkable the UTA Amphibian and Reptile Diversity Research and since the two species most likely have overlapping Center (UTADC-6724–45, 6831–32). ranges (Figure. 3), it is not hard to postulate that they are Müllerian co-mimics. Our proposal of sub-adult C. Results bibroni being co-mimics of Sinomicrurus macclellandii Molecular data.— The Calliophis species from is primarily based on the fact that the latter is the only Kannur, is genetically undifferentiated from the C. other venomous South Asian snake to display a red bibroni individuals from Kerala with a maximum p- dorsum with black bars (Whitaker et al., 2004; Deepak distance of 0.6% and a minimum p-distance of 0 % et al., 2010). It is to be noted, that the range of the two (Table 2). The C. castoe specimen is 22.0% different species does not overlap and they are separated by over from the Calliophis species from Kannur, which it 1000 km, with C. bibroni being restricted to the west shares near identical dorsal colouration with. coast of south India, while S. macclellandi is only found in the forests of north and northeastern India (Whitaker Meristic data and colouration.— The specimen et al., 2004). However, it has already been recognized from Kannur is an adult female with SVL -measuring that mimicry need not be restricted to sympatric species 587 mm (656 mm total length). Despite of superficial (Pfennig and Mullen, 2010; Pfennig et al., 2015); several resemblance, the individual differs from Calliophis New World coralsnake mimics have ranges that exceed castoe (characters in parenthesis) in the following that of their models (Endler, 1981). characteristics: no preocular (one preocular); one The two assumptions then, implicit to our reasoning postocular (two postoculars); ventrals - 224 (240–254); are: 1) banded patterns (as seen in Sinomicrurus subcaudals - 39 (45-53); single anal plate (divided anal macclellandi) function as aposematic signals (Brodie, plate) (Figure. 1). All of the meristic data mentioned fall 1993; Mochida et al., 2015) ; 2) avifauna are significant close to or within the morphological variation reported predators of Old World coralsnakes, as is the case of in C. bibroni (Deepak et al., 2010). their New World counterparts (Pough, 1988). If these The Kannur specimen has no temporal extensions of assumptions hold true then it entirely plausible that the head cap (temporal extension of head cap present); a wide-ranging or migratory avian predator would nuchal/parietal band yellow (orange); midvertebral generalize its avoidance of an unprofitable prey with sepia stripe occupying vertebral and paravertebral rows a characteristic bright banded pattern (Greene and (no stripe); ventral surface of tail as well as posterior McDiarmid, 1981). Indeed, several wide-ranging Indian body black/sepia, midventral colour orange (orange raptors are known to be forest dwellers, specializing on with no black/sepia markings); single anal plate almost snakes and other : the Crested Serpent Eagle completely sepia/black, except for orange posterior Spilornis cheela (Latham, 1790) and the Changeable border (single anal plate, white with only slightly orange Hawk Eagle Spizaetus cirrhatus (Gmelin, 1788) both lateral edges) (Figure.2 C-F). reside in tropical evergreen forests across India, and are known to actively prey on snakes; other widespread

Table3 2.Table Uncorrected 2. Uncorrected pairwise pairwise genetic distancesgenetic distances (p-distances) (p-distances) of Cytb sequencesof Cytb sequences from the fromspecimens the specimens used in this used study. in this study.

M816 C. bibroni M817 C. bibroni M819 C. sp. M731 C. bibroni M733 C. bibroni Wynad, Kerala Trivandrum, Kerala Kannur, Kerala Tamil Nadu Tamil Nadu M816 C. bibroni Wynad, Kerala

M817 C. bibroni Trivandrum, Kerala 0.006

M819 C. sp. Kannur, Kerala 0.000 0.006

M731 C. bibroni Tamil Nadu 0.002 0.008 0.002

M733 C. bibroni Tamil Nadu 0.002 0.008 0.002 0.000

M706 C. castoe Goa 0.219 0.222 0.219 0.221 0.221 4

2

A new colour morph of Calliophis bibroni 213

Figure 1. Color patterns in Bibron’s coralsnake, Calliophis bibroni, and comparison to suspected Müllerian co-mimics. Dorsum (A) and venter (B) of adult male C. bibroni, adult with cryptic dorsal colour and aposematic venter. Dorsum (C) and tail ventral surface of adult C. bibroni (D) resembling C. castoe (F). Dorsum of juvenile C. bibroni (E), resembling S. macclellandi (G). Individuals of C. bibroni presented in A-B, C-D, and E correspond to M816, M819 and M817, respectively. Photos A–E by Dileep Kumar, F by Hemant Ogale, and G by Ishan Agarwal. 214 Dileep Kumar Raveendran et al.

Figure 2. Lateral and dorsal head views of selected coralsnakes of the Western Ghats showing convergent head colour patterns and scalation, adult Calliophis bibroni with cryptic dorsal colour (A and B), adult C. bibroni with Mullerian mimicry with C. castoe (C and D), and adult C. castoe (E and F).

raptors such as the Common Buzzard Buteo buteo Karnataka, India (http://walkthewilderness.net/brown- (Linnaeus, 1758) , the Oriental Honey Buzzard Pernis fish-owl-with-coral-snake-kill-in-br-hills/). ptilorhycus (Temminck, 1821) and the Booted Eagle Given that Calliophis bibroni are known to attain a Hieraaetus pennatus (Gmelin, 1788) also frequent total length of 880 mm while Sinomicrurus macclellandi forests and regularly feed on reptiles (Naoroji and in India barely get over 800 mm long (Whitaker et al., Schmitt 2007). DV has found a dead Calliophis bibroni 2004; Deepak et al., 2010), it is possible that the former with perforations made by birds on two occasions would benefit by changing their mimicry strategy if (Deepak, 2015). There is also a confirmed report of they are too large to effectively mimic S. macclellandi a Brown Fish Owl Bubo zeylonensis (Gmelin, 1788) as adults. Old World coralsnakes are known to flip feeding on a Striped coralsnake C. nigrescens from over and expose their bright bellies when threatened. A new colour morph of Calliophis bibroni 215

Figure 3. Map showing the localities of Calliophis castoe and C. bibroni in India, including those used in this study (modified from Deepak et al., 2010).

Retaining an aposematic venter could actually provide - Even though less drastic than that of Calliophis bibroni the snake a last resort opportunity to fend off any (Deepak et al., 2010), an ontogenetic transition from predator that was not fooled by crypsis and was naïve aposematism to crypsis has also been reported in enough to proceed attacking after having located the New World coralsnakes; for example, in , snake. Conceding that we have no experimental backing Allen’s coralsnake Micrurus alleni (Schmidt, 1936) to our claims of mimicry between the allopatric C. is known to undergo an ontogenetic shift wherein its bibroni and S. macclellandi, there is no means for us vivid pattern of red, yellow and black rings change to theoretically dismiss alternatives that could serve as into uniform black. Remarkably, the sympatric drivers of convergent evolution in colouration. Flicker- Milksnake Lampropeltis triangulum (Lacépède, fusion (Pough, 1976), background matching, or simply 1789) known to be a Batesian mimic of the M. alleni, the innate fear of aposematic phenotypes (Smith, 1975, displays a similar ontogenetic shift (Savage and Vial- 1977), are all hypothesized to drive the independent Kearney, 1974; Greene and McDiarmid, 2005). There evolution of parallel external appearances in snakes. is no comparable instance of ontogenetic transition Nevertheless, the complex combination of involving Müllerian co-mimics in New World aposematism and mimicry in Calliophis bibroni elapids. reported here is unique in many respects. Below we - Some New World snakes are also known to combine discuss what sets it apart from chromatic strategies features of two or more models, as seen with the seen in other relevant snake mimicry systems of the Variegated False coralsnake Pliocerus elapoides New World. (Cope, 1860) in southern and northern Guatemala, which combines the colour sequence 216 Dileep Kumar Raveendran et al.

of two sympatric coralsnake species (Pough, Rica. Evolution 47: 227–235. 1988). However, this dual mimicry is displayed Brodie, E. D., Janzen, F. J. (1995). Experimental Studies of Coral simultaneously and is not sequentially associated Snake Mimicry: Generalized Avoidance of Ringed Snake Patterns by Free-Ranging Avian Predators. Functional Ecology with ontogeny as seen with C. bibroni. 9: 186–190. - While apostasy (frequency-dependent selection of Campbell, J. A., Lamar, W. W. (1989). The venomous reptiles of rare morphs[Clark, 1969]) is expected to maintain Latin America. polymorphism in Batesian mimics, Müllerian co- Castoe, T. A., Smith, E. N., Brown, R. M., Parkinson, C. L. (2007). Higher-level phylogeny of Asian and American coralsnakes, their mimics typically tend to be monomorphic (Endler and placement within the Elapidae (Squamata), and the systematic Greenwood, 1988; Joron and Mallet, 1998; Mallet and affinities of the enigmatic Asian coralsnake Hemibungarus Joron, 1999). Calliophis bibroni is unique in that it calligaster (Wiegmann, 1834). Zoological Journal of the Linnean not only displays polymorphism (i.e. the plain morph Society 151: 809–831. and the C. castoe morph) while being a Müllerian co- Deepak, V., S. Harikrishnan, Vasudevan, K., Smith. E. N. (2010). mimic, but that the two colour morphs are sympatric Redescription of Bibron’s coralsnake, Calliophis bibroni Jan (Figure. 3). And even though conspicuous exceptions 1858 with notes and new records from south of the Palghat and Shencottah gaps of the Western Ghats, India. Hamadryad 35: to this rule are frequently seen in several species of 1–10. butterflies (Mallet and Joron, 1999), polymorphism Dunn, E. R. (1954). The “Mimic” Problem in . in Müllerian mimics within New World coralsnakes Evolution 8: 97–102. is yet to be reported. Dutta, S. K., Vasudevan, K., Chaitra, M. S., Shanker, K., Aggarwal. R. K. (2004). Jurassic frogs and the evolution of amphibian In summary, the new colour morph of Calliophis endemism in the Western Ghats. Current Science 86: 211–216. bibroni described here provides a unique glimpse into Endler, J. A. (1981). An overview of the relationships between the complex eco-evolutionary interactions occurring mimicry and crypsis. Biological Journal of the Linnean Society between an enigmatic group of tropical venomous snakes 16: 25–31. and their environment, which have not been reported Endler, J. A., Greenwood. (1988). Frequency-Dependent , before in any coralsnakes. The hypotheses we discuss Crypsis and Aposematic Coloration [and Discussion]. above are testable with rigorous field experiments, Philosophical Transactions of the Royal Society of London B: involving clay or plasticine models. We hope that the Biological Sciences 319: 505–523. Gadow, H. (1911). Isotely and coral snakes. Zoologische Jahrbücher evidence provided herein will motivate ecologists für Systematik, Ökologie und Geographie der Tiere 31: 1–24. to initiate pioneering experimental studies, drawing Gehlbach, F. R. (1972). Coral snake mimicry reconsidered: the inspiration from the elegant work on Neotropical snake strategy of self-mimicry. Forma et Functio 5: 311–320. fauna, exploring mimicry, crypsis and aposematism in Gowri Shankar P., Ganesh S. R. (2009). Sighting record and range Old World coralsnakes. extension of Calliophis. Herpetological bulletin 108: 10–13. Greene, H. W., McDiarmid, R. W. (1981). Coral Snake Mimicry: Acknowledgments. We would like to thank Dr. Praveen K. Does It Occur? Science 213: 1207–1212. Karanth and his lab (Center for Ecological Sciences, Indian Greene, H. W., McDiarmid, R. W. (2005). Wallace and Savage: Institute of Science, Bengaluru) and Dr. Ramesh K Aggarwal and heroes, theories and venomous snake mimicry. In: Ecology his lab (Center for Cellular and Molecular Biology, Hyderabad) and Evolution in the Tropics: A Herpetological Perspective, p. for sequencing the Calliophis castoe and C. bibroni individuals 190-208.���������������������������������������������������� � . respectively. We would like to thank Dr. Karthikeyan Vasudevan, White, M.E., Ed., ��������������������������������������������� (Lab for the Conservation of Endangered Species, Hyderabad) for Press providing specimens of C. bibroni used in this study. We would Joron, M., Mallet, J. L. B. (1998). Diversity in mimicry: paradox or like to thank Ishan Agarwal and Hemant Ogale for allowing us paradigm? Trends in Ecology & Evolution 13: 461–466. to use the photos of Sinomicrurus macclellandi and C. castoe Kikuchi, D. W., Seymoure, B. M., Pfennig, D. W. (2014). Mimicry’s respectively. VD is grateful to Silamban and Karuppaswami for palette: widespread use of conserved pigments in the aposematic their assistance in the field. 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Accepted by Kotambylu Gururaja