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Widespread Boa Corallus Enydris Inferred from Mitochondrial DNA Sequences ROBERT W

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Widespread Boa Corallus Enydris Inferred from Mitochondrial DNA Sequences ROBERT W MOLECULAR PHYLOCENETICS AND EVOLUTION Vol. 4. No.1. March. pp. 88-92. 1995 Origin of West Indian Populations of the Geographically Widespread Boa Corallus enydris Inferred from Mitochondrial DNA Sequences ROBERT w. HENDERSON AND S. BLAIR HEDGES* Section of Vertebrate Zoology, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 53233-7478; and • Department of Biology, 208 Mueller Lab, Penn State Unil/ersity. University Park, Pennsylvania 76802 Received June 30. 1994; revised Sept~'mber 30. 1994 recognized subspecies (Fig. 1): C. e. enydris which oc­ Corallus enydris (Serpentes: Boidae: Boinae) is an ar­ curs throughout most of the Amazonian lowlands, the boreal snake with an extremely wide mainland distri­ Guianas, and the Atlantic coastal forest of Brazil and bution from southern Costa Rica to southeastern Brazil C. e. cooki which occurs in southern Central America, arid is one of two boine species that has invaded the northern Colombia, northern Venezuela (including Lesser Antilles (Grenada Bank and St. Vincent). Mito­ Isla Margarita), Trinidad and Tobago, and the Lesser chondrial DNA sequences of samples from seven geo­ Antilles as far north as St. Vincent. C. enydris is more graphically disparate localities provided evidence of variable in color and pattern than any other boine phylogenetic relationships. The monophyly of C. en­ (e.g., Figs. 1-7 in Henderson, 1993). In this paper we ydris is corroborated and a major dichotomy between present data regarding phylogenetic relationships in northern samples (Panama and Trinidad) and southern C. enydris based on mitochondrial DNA sequences of samples (Guyana, Peru, southeastern Brazil) was found sample~ from seven geographically disparate localities and corresponds to the two currently recognized sub­ and evidence for the origin of the West Indian popula­ species. Unexpectedly, the two samples from the West Indies (southern Lesser Antilles) cluster with the south­ tions. ern rather than the geographically closer northern samples (e.g., Trinidad). The results imply a fairly re­ MATERIALS AND METHODS cent Guianan-Amazonian origin of West Indian popu­ lations. tl 1986 Academic Pre... lac. DNA was extracted from small amounts «50 mg) of liver or red blood cells from the following specimens of C. enydris: MPM (Milwaukee Public Museum) INTRODUCTION 26290, Panama, Canal Zone; MPM 23596, St. Vincent, St. Patrick, 2.0 miles ENE of Layou; MPM 25445, Gre­ The neotropical mainland boine snake (Serpentes: nada, St. Andrew, Pearls; MPM 23595, Trinidad, Hol­ Boidae: Boinae) fauna, currently comprised of 10 spe­ lis Reservoir; LSUMZ (Louisiana State University Mu­ cies, includes five species with widely sympatric ranges seum of Zoology) 43139, Guyana, Jonestown; KU in Amazonia and the Guianas: Boa constrictor, Cor­ <University of Kansas) 204895, Peru, Cuzco, Cuzco all us caninus, Corollus enydris, Epicrotes cenchria, Amazonica; and IB (lnstituto Butantan) 55042, Brazil, and Eunectes murinus. The ranges of B. constrictor, C. Siio Paulo, 19uape (Fig. 1). Considering the broad enydris, and E. cenchria are even broader, extending range of Corollus enydris, we acknowledge the rela­ from Mexico and/or southern Central America to tively small number of samples in our analysis. Those southeastern Brazil and/or northern Argentina; only we have, however, are geographically strategic (Fig. B. constrictor and C. enydris have invaded the West 1), being near (1) the northern edge of the mainland Indies. Only C. caninus is regarded as monotypic, range (Panama), (2) the southern end of the mainland whereas B. constrictor and E. cenchria have 9 or 10 range (Atlantic coastal forest in southeastern Brazil), currently recognized subspecies. They are poorly de­ and (3) the western edge in the Upper Amazon of Peru fined geographically, taxonomically, and systemati­ and from (4) two West Indian islands, including one cally, although recent advances have been made (e.g., from the northernmost edge of the range (St. Vincent) Tolson, 1987; Kluge, 1991) or are in progress (R.W.H., and (5) Trinidad. The Trinidad sample (C. e. cooki) is in preparation). critical because of its geographic proximity to the C. enydris, an arboreal species ranging from south­ Lesser Antilles and because of its morphological simi­ ern Costa Rica to southern Brazil, has two currently larity to Venezuelan populations occurring between 88 1055-7903/95 $6.00 Copyright It 1995 by Academic Press. Inc. All rights of reproduction in any fonn reserved. ORIGIN OF ANTILLEAN POPULATIONS OF THE BOA Corallus enydris 89 o 300 600 900 KM 1'1 1,1 I I I I FIG. 1. Geographic distribution of the boid snake Corallus enydris, showing the locations of samples used in this study. The more northern, darker stippling indicates the range of C. e. cooki, and the more southern, lighter stippling indicates the range of C. e. enydris. The numbered localities indicate sites represented in our sample: (1) Panama, (2) St. Vincent, (31 Grenada, (4) Trinidad, (5) Guyana, 161 southern Peru, and (7) southeastern Brazil. See Materials and Methods for more precise locality data. the Caribbean coast and the Rio Orinoco (Henderson Phylogenetic analysis (distance and parsimony) was and Boos, 1993). Despite long distances between Guia­ performed with MEGA (Kumar et aZ., 1993). The nan-Amazonian sample localities on the South Ameri­ Jukes-Cantor (1969) distance was used with the can mainland (Guyana-Brazil, ca. 3500 km; Guyana­ neighbor-joining method (Saitou and Nei, 1987), and Peru, ca. 2700 km) and the potential for wide rivers statistical confidence of the nodes on the trees was in­ (e.g., Rio Amazonas) to act as barriers to dispersal, it is ferred by a t test for branch-length significance from likely that the Guianan-Amazonian range (including zero, expressed as the complement of the probability, Atlantic coastal forest in Brasil) of C. enydris is contig­ or confidence probability (CP; Rzhetsky and Nei, 1992; uous (Fig. 1; Henderson, 1993). An individual of C. Kumar et aZ., 1993), and by the bootstrap method caninus (Audubon Zoo 5902, locality unknown) was in­ (Felsenstein, 1985) using 2000 replications (Hedges, cluded for comparison, and the boid Epicrates striatus 1992). Statistical significance was assessed at the 95% (S.B.H. Laboratory No. 103120, Dominican Republic, level. Maximum parsimony analysis was done with the Samana, 7 km S. of Las Galeras) was included for the modified branch and bound algorithm in MEGA. Sites purpose of rooting the tree. containing ambiguities were not included in the analy­ Methods of DNA extraction, amplification, and di­ ses. deoxy sequencing are described elsewhere (Hedges et al., 1991; Hedges and Bezy, 1993). Two oligonucleotide RESULTS primers were used to amplify and sequence both com­ plementary strands ofa 307-bp region of the mitochon­ A total of 271 aligned sites (nine taxa) could be drial cytochrome b gene (Hedges et aZ., 1992). In some scored from autoradiograms and were used in the phy­ cases, two other primers designed to amplify the same logenetic analyses (Fig. 2). Of these, 74 were variable region (Kocher et al., 1989) also were used. Aerosol­ and 28 were parsimony sites (those sites informative resistant tips were used in preparation of reagents in under the conditions of parsimony). Pairwise corrected order to reduce the probability of contamination. There distances among the specimens of C. enydris were low were no insertions or deletions and therefore align­ «0.08), supporting the use of a Jukes-Cantor distance ment was straightforward. rather than a more complicated correction (Nei, 1991). 90 HENDERSON AND HEDGES 46 1 E.strfetua C TTC GGA TCC ATA CTA CTT GCT TGe TTA TCT CTA CAA CTA CTT ACA 2 cenf,.. T •• r ••••• T •• C A.C •• T .C. G.C ••• G•••• A ••• 3 p.".. •• G ••• ••• ..C A.C •••• C. G.C T.G ••• G•••• G ••• 4 Trinided •• G •••••• • •• A.e •••• C. G.e T•• ... G•••• A ••• 5 Guyene .. A .. •• C A.C •••• CG G.C ••• G•••• G ••• 6 Bruil ..A .. •• C A.C •••• CG G.C G•••• G ••• 7 Peru •• A ••• •• C A.C •••• CG G.C G.. ..G ••• 8 Grenldli .. A ... •• C A.C •••• CG G.C G.G •• G ... 9 St.Vlncent •• A .,. •• C A.C •••• CG G.C G.... G ••• 91 1 E.strfetua GGA TTC TTC TTA GeT GTA CAC TAC ACA GCA MT ATT MC CTA GCA 2 cenl,.. •• C ••••• T C•••• C ••••• T ••• ..C •• C G•• T•• 3 Penema •• C .. r C.. ..C .. C G.. T.... G 4 Trinided •• C C.. ..C .. C ... T.... G 5 Guyena •• C '" C.. ..C .. C G.. T.. 6 Irull .. C C.. ..C .. C G.. r .. 7 Peru •• C C.. ..C .. C G.. T.. 8 Grenada .. C C.. ..C .. C G.. r .. 9 St.Vlncent .. C C.. ..C •• C G.. T .. 136 1 E.strletus TTC TCA TCT ATC ATC CAC ATT ACC CGA GAT GTC CCA TAT GGA TGA 2 cenlnus .. C G.. G.... T .T. ..C .. C 3P_ G.T G.. .TA .. C .. C 4 Trinidad ... G.T G.. .TA ..... C .. C ... 5 Guy.". ... G.T G.. .TA ..... C .. C .. 6 Brezil ... G.T G.. .TA ..... C .. C .. 7 Peru G.T G.. .TA .....C .. C 8 Grenade G.T G.. .TA .....C ..C 9 St.Vincent '" G.T G.. .TA ..... C .. C '" 181 1 E.str!etus ATA ATA CAA MC CTT CAC GeT ArT GGG GCC TCA GTA rTA TTT ATT 2 caninus •• A •• C •• A ••• A•••• C •• C •• C 3 PIINIIIIIl .. T .. C .. C .. A ... A.... C .. C .. C 4 Trinidad .. T .. C .. C .. G ... A.... C .. C .. C 5 Guy.". T.. ..T •• C •• T •• C •• A •• A ••• A•••• C •• C •• C 6 Brazil •• T •• C •• T •• C •• A ••• A•••• C •• C •• C 7 Peru .. T .. A .. T .. C .. A ... A.... C .. C .. C a Grenade •• T •• C •• T •• C •• A ••• A•••• C •• C •• C 9 St.Vlncent •• T •• C •• T •• C •• A ••• A•••• C •• C •• C 226 1 E.strietus TGT ATT TAT ATC CAT ATC GCA CGA GGC TTG TAT TAT GGG TCG TAC 2 caninus .
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