Bothriechis guifarroi 2013

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A relict lineage andnew species of green palm-pitviper(Squamata, Viperidae, Bothriechis)... 77

A rc nag and nw spcs of grn pam-pvpr(Squamaa, Vprda,Bothriechis) from h Chors

Hghands of Msoamrca

Josiah H. ownsend1,2,, Melissa Medina-Flores1,3,, Larry David Wilson2,,Robert C. Jadin4,|, James D. Austin5,

1 Department of Biology, Indiana University of Pennsylvania, Indiana, Pennsylvania 157051081, USA

2 Centro Zamorano de Biodiversidad, Escuela Agrcola Panamericana Zamorano, Departamento de FranciscoMorazn, Honduras3 Escuela de Biologa, Universidad Nacional Autnoma de Honduras, Tegucigalpa, Fran-

cisco Morazn, Honduras4 Department of Ecology and Evolutionary Biology, University of Colorado Boulder,

Boulder, Colorado 80309, USA; and Amphibian and Reptile Diversity Research Center, University of Texas at

Arlington, Arlington, Texas 76019, USA 5 Department of Wildlife Ecology and Conservation, University of

Florida, Gainesville, Florida 32611, USA

urn:lsid:zoobank.org:author:16C11549-EB20-4273-B684-E143D6A9788D

urn:lsid:zoobank.org:author:F27FCAF3-1447-42B3-A0B2-47E4C967D611

urn:lsid:zoobank.org:author:98AB7342-BF24-4220-B367-29A5CCF9175F

| urn:lsid:zoobank.org:author:39AE5D95-4DAC-429E-A7FC-526105D18F88 urn:lsid:zoobank.org:author:1DE20BB5-0658-4949-8B07-6D25B2AB9499

Corresponding author:Josiah H. Townsend([email protected])

Academic editor:J. Penner | Received 7 February 2013 | Accepted 23 April 2013 | Published 13 May 2013

urn:lsid:zoobank.org:pub:0F1B0D4E-3705-452B-AFBB-337C8156B8BB

Caon: ownsend JH, Medina-Flores M, Wilson LD, Jadin RC, Austin JD (2013) A relict lineage and new species

o green palm-pitviper (Squamata, Viperidae, Bothriechis) rom the Chorts Highlands o Mesoamerica. ZooKeys 298:

77106. doi: 10.3897/zookeys.298.4834

Absrac

A new species o palm-pitviper o the genus Bothriechis is described rom Reugio de Vida Silvestreexguat in northern Honduras. Te new species diers rom congeners by having 19 dorsal scale rows atmidbody, a bright green dorsal coloration in adults, the prelacunal scale used to the second supralabial,and in representing a northern lineage that is sister to B. lateralis, which is distributed in Costa Rica andwestern Panama and is isolated rom the new taxon by the Nicaraguan Depression. Tis represents the

15th endemic species occurring in Reugio de Vida Silvestre exguat, one o the richest herpetoaunal

ZooKeys 298: 77105 (2013)

doi: 10.3897/zookeys.298.4834

www.zookeys.org

CopyrightJosiah H. Townsend et al.This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0

(CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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Josiah H. Townsend et al. / ZooKeys 298: 77106 (2013)78

sites in Honduras, itsel being the country with the highest degree o herpetoaunal endemism in CentralAmerica. We name this new species in honor o a Honduran conservationist slain in ghting against illegallogging, highlighting the sacrices o rural activists in battling these issues and the critical importance oconservation in these areas.

Rsumn

Una nueva especie de tamags verde del gnero Bothriechis se describe del Reugio de Vida Silvestreexguat en el norte de Honduras. La nueva especie diere de sus congneres por tener las de 19 escamasdorsales en la mitad del cuerpo, una brillante coloracin verde en la porcin dorsal del cuerpo en adultos,la escama prelacunal usionada con la segunda supralabial, y en representacin del linaje del norte que esclado hermano de B. lateralis, la cual se distribuye en Costa Rica y el Occidente de Panam y est aisladadel nuevo taxn por la Depresin de Nicaragua. Representa la decimoquinta especie endmica encontradaen el Reugio de Vida Silvestre exguat, uno de los lugares ms ricos de herpetoauna en Honduras, elpas con el ms alto grado de endemismo de la herpetoauna en Centroamrica. Nombramos esta nueva

especie en honor a un conservacionista Hondureo asesinado en contienda contra la tala ilegal, desta-cando los sacricios de los activistas rurales luchando rente a estos problemas y la importancia crtica dela conservacin en estas reas.

Kywords

Bothriechisguifarroisp. n., Bothriechislateralis, Bothriechismarchi, Central America, conservation, crypticspecies, endemic, Honduras, Pico Bonito National Park, exguat Wildlie Reuge

Palabras clavs

Bothriechisguifarroi

sp. n.,Bothriechis

lateralis

,Bothriechis

marchi

, Centroamrica, conservacin, ende-mismo, especies crpticas, Honduras, Parque Nacional Pico Bonito, Reugio de Vida Silvestre exguat

inroducon

In the past decade, a steady stream o taxonomic discoveries have come out o theChorts Highlands o Mesoamerica, a biogeographic region ound to the south andeast o the tectonic boundary between the Chorts and Mayan Blocks and north o the

Nicaraguan Depression (ownsend 2011, ownsend et al. 2011). Fity new species oamphibians and reptiles have been described rom the regions montane orests since2000 (Cadle 2012, Cadle and Savage 2012, McCranie and Hedges 2012, Rovito et al.2012), with literally dozens more awaiting description (ownsend 2011).

Our knowledge o the taxonomic diversity o Mesoamerican pitvipers has alsogreatly increased since the turn o the century (e.g. Campbell and Flores-Villela 2008,Jadin et al. 2011). Tree species o endemic pitvipers have been described rom theChorts Highlands since 2000: Atropoides indomitusSmith & Ferrari-Castro 2008,Bothriechis thalassinus Campbell & Smith 2000, and Cerrophidion wilsoni Jadin,

ownsend, Castoe & Campbell 2012. wo o these three taxa, B. thalassinusand C.wilsoni, had previously been concealed within more widespread taxa only to be re-vealed by more ocused sampling and phylogenetic analyses.


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A relict lineage and new species of green palm-pitviper(Squamata, Viperidae, Bothriechis)... 79

Te green palm-pitvipers (genus Bothriechis) o Mesoamerica have long been asource o taxonomic uncertainty and conusion (Campbell and Lamar 2004). Ambi-guities among type specimens and localities, the imprecise provenance o many avail-able specimens, disjunct distributions limited to ragmented highland orests, and

misleading external morphology have all contributed to a lack o taxonomic resolutionamong populations currently assigned to two species rom the Chorts Highlands; Bo-thriechis marchi(Barbour and Loveridge 1929) and B. thalassinus. As currently under-stood, these two taxa inhabit a number o disjunct localities in the Chorts Highlands(Campbell and Lamar 2004, McCranie 2011a). Available molecular data or these twotaxa are limited to a single sample assigned to each nominal orm, which indicate thetwo are sister species nested within a Nuclear Central American clade (also includingB. aurifer, B. bicolor,and B. rowleyi) that is, in turn, sister to the highland Bothriechis(i.e. B. lateralisand B. nigroviridis) ound in lower Central America (aggart et al.2001, Castoe and Parkinson 2006, Castoe et al. 2009).

Bothriechis marchisensu lato is known rom localities in the Cordillera Nombrede Dios, Cordillera de Merendn, and Sierra de Sulaco o Honduras and adjacentareas o Guatemala, with B. thalassinusbeing ound in the Cordillera de Merendno Guatemala and Honduras, Cerro Santa Brbara and nearby highland orests inHonduras, Cerro del Mono in eastern Guatemala, and the highlands around the ElSalvador-Guatemala-Honduras border area (Campbell and Lamar 2004; McCranie2011a). Both o these taxa include a number o allopatric highland populations thathave not been assessed using phylogenetic methods. O interest here are popula-tions rom the Cordillera Nombre de Dios, ound within and around Reugio deVida Silvestre exguat and Parque Nacional Pico Bonito. Tese two cloud orestreserves are each recognized or their diverse endemic herpetoauna (McCranie andCastaeda 2005; ownsend et al. 2012) and are taxonomically and biogeographi-cally distinctive rom that o the northern Cordillera de Merendn, which includesthe vicinity o the type locality o B. marchi in the Sierra de Omoa (ownsendand Wilson 2008).

wo expeditions in 2010 provided the rst herpetoaunal inventory o the ex-

tensively orested windward portions o Reugio de Vida Silvestre exguat, one othe most endemism-rich highland orests in Mesoamerica (ownsend et al. 2012).During two visits to the windward side o Reugio de Vida Silvestre exguat inJune and July 2010, we collected a series o arboreal pitvipers representative othose assigned to B. marchi. Phylogenetic analyses revealed that the populationrom Reugio de Vida Silvestre exguat is not conspecic with the nominal taxonB. marchi, nor are they part o the Nuclear Central American clade containingB.marchiand B. thalassinus. Remarkably, this population is shown to represent a rel-ict northern lineage that is most closely related to B. lateralisrom Costa Rica and

western Panama. We herein describe the Reugio de Vida Silvestre exguat popu-lation oBothriechisas a new taxon, and discuss the implications or systematics,biogeography, and conservation.


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Materials and methods

Field-based sampling

Te type series was collected during sampling in the vicinity o La Liberacin (15.53N,87.29W; camp established at 1,030 m elevation) during 1021 June (11 participants;1,320 person-hours sampling) and 26 July2 August 2010 (13 participants; 880 personhours). issue samples were preserved in SED buer (20% DMSO, 0.25 M EDA,pH 7.5, NaCl saturated; Seutin et al. 1991, Williams 2007) and whole specimens in10% ormalin and later transerred to 70% ethanol. Specimens were deposited in theCarnegie Museum o Natural History (CM), Museum o Vertebrate Zoology, Univer-sity o Caliornia Berkeley (MVZ), National Museum o Natural History, SmithsonianInstitution (USNM), and Amphibian and Reptile Diversity Research Center, Univer-sity o exas at Arlington (UA).

DNA extraction, amplifcation, and sequencing

Genomic DNA was isolated rom muscle tissue taken rom eleven specimens oBoth-riechisusing a Qiagen DNeasy extraction kit and protocol. Four mitochondrial generagments (NADH dehydrogenase subunit 4 (ND4), cytochrome b (cyt b), 12S rRNA,and 16S rRNA) were independently PCR-amplied as described in multiple studies(Knight and Mindell 1993; Arvalo et al. 1994; Parkinsonet al. 1997; Parkinson etal. 2002) using Promega Goaq Green master mix, the primer pairs ND4 + LEU,Gludg + AtrCB3, L1091 + 12E, and 16SF + 16SR, and annealing temperatures 48C,48C, 50C, and 45C, respectively. Sequencing was perormed in both orward andreverse directions using the PCR primers on a Beckman Coulter automated capillarysequencer, and sequence chromatographs were edited using Sequencher 4.2. Sequencesor each gene were aligned separately, rst automatically using the program MUSCLE(Edgar 2004), and then manually rechecked using Se-Al v2.0a11. Gaps in alignments

were treated as missing data. No internal stop codons were ound in the two protein-coding gene ragments. Novel sequences rom this study were deposited in GenBank(KC847255289).

Previously published sequences o Bothriechiswere downloaded rom GenBankand combined with new sequence data generated in this study (able 1). Representa-tives o two Mesoamerican genera rom the diverse sister clade to Bothriechis(whichcontainsAtropoides[Mesoamerica], Bothriopsis[South America], Bothrocophias[SouthAmerica], Bothrops[Mexico to South America], Cerrophidion (Mesoamerica], and Por-thidium [Mexico to South America]) were selected or use as outgroups to root our Bo-

thriechisphylogeny (Castoe et al. 2005, 2009; Daza et al. 2010):Atropoides mexicanusand Cerrophidion wilsoni, both o which are sympatric with Bothriechisin Reugio deVida Silvestre exguat (ownsend et al. 2012).


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tabl 1. axa, vouchers, locality data, and GenBank accession numbers or sequences used in this study.Novel sequences rom this study are indicated in boldace; country codes used as ollows: CR = CostaRica; EC = Ecuador; G = Guatemala; HN = Honduras; MX = Mexico; NI = Nicaragua.

Taxon Locality Voucher

GenBank Accession Numbers

ND4 cyt b 12S 16S

Atropoides mexicanusHN: Atlntida:

exguatUSNM 578906 KC847289 KC847271 KC847268 KC847255

Bothriechis aurifer G: UA R-35031 DQ305483 DQ305466 DQ305425 DQ305448Bothriechis bicolor G: UA R-34156 DQ305484 DQ305467 DQ305426 DQ305449

Bothriechis guifarroi

HN: Atlntida:exguat

CM 156870 KC847280 KC847260

HN: Atlntida:exguat

MVZ 269305 KC847279 KC847258

HN: Atlntida:

exguat

USNM 579873 KC847286 KC847274 KC847267 KC847262

HN: Atlntida:exguat

USNM 579874 KC847288 KC847282 KC847266 KC847263

HN: Atlntida:exguat

USNM 579875 KC847287 KC847281 KC847265 KC847264

HN: Atlntida:exguat

USNM 579876 KC847276

HN: Atlntida:exguat

USNM 579877 KC847278

HN: Atlntida:exguat

USNM 579878 KC847277 KC847261

HN: Atlntida:exguat UA R-60303 KC847275 KC847259

Bothriechis lateralis CR: Acosta MZUCR-11155 U41873 AY223588 AF057211 AF057258

Bothriechis marchi

G: Zacapa:Cerro del

MonoUA R-52959 DQ305486 DQ305469 DQ305428 DQ305451

HN: Corts:Sierra de Omoa

MVZ 263604 KC847283

Bothriechisnigroviridis

CR: SanGerondo de

DotaMZUCR-11151 AY223635 AY223589 AF057212 AF057259

Bothriechis rowleyi MX: CerroBal JAC 13295 DQ305485 DQ305468 DQ305427 DQ305450

Bothriechis schlegelii

CR: Cariblancode Sarapiqu

MZUCR-11149 AY223636 AY223590 AF057213 AF057260

EC: Pichincha FHGO Live coll AF292611 AF292573 HN: Corts:

YojoaUF 157577 KC847285 KC847272 KC847270 KC847257

NI: Jinotega:Bosawas

UF 166874 KC847284 KC847273 KC847269 KC847256

Bothriechissupercilliaris

CR: San Vito DQ305487 DQ305470 DQ305429 DQ305452

Bothriechisthalassinus

G: Zacapa UA R-52958 DQ305482 DQ305465 DQ305424 DQ305447

Cerrophidion wilsoniHN: Olancho:

BotaderosUA R-52953 JQ724172 JQ724159 JQ724146 JQ627132


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Phylogenetic analyses

Bayesian inerence (BI) and maximum likelihood (ML) were implemented to recon-struct phylogenies or the Bothriechisingroup taxa. o identiy appropriate models

o nucleotide substitution or both analyses, we used the program MrModeltest v2.2(Nylander 2004), run in PAUP* v4.0b10 (Swoord 2002). We used Akaike inor-mation criterion (AIC) to select the best-t models, as estimated by MrModeltest(able 2). Te our gene ragments were concatenated (2,263 total bp), and thiscombined dataset was partitioned by gene and codon position (or cyt-b and ND4),resulting in a total o eight partitions as was shown to be justied in analysis o asimilar dataset that included these our ragments rom these species (Castoe andParkinson 2006). Stems and loops were not partitioned separately due to a lack oinormative characters.

Phylogenetic analyses using BI were conducted with MrBayes v3.0b4 (Ronquistand Huelsenbeck 2003). wo simultaneous BI runs were conducted (with the deaultMarkov chain Monte Carlo [MCMC] settings), and run or a total o 5.0 10 6 gen-erations per run, sampling trees and parameters every 100 generations. We used PSRFvalues (output by MrBayes), together with plots o cold chain likelihood values andparameter estimates visualized in racer v1.5.4 (Rambaut and Drummond 2009), toconrm stationarity and convergence o MCMC runs. Based on this evaluation, therst 1.5 105 generations rom each run were discarded as burn-in.

Phylogenetic relationships were inerred using ML as implemented in RAxML7.2.8 (Stamatakis 2006, Stamatakis et al. 2008), using the same partitioning schemedescribed above or BI. ree support was assessed using the rapid-bootstrapping algo-rithm with 1000 non-parametric bootstraps; all ML estimates and tests were run underthe GRCA model, as models available or use in RAxML are limited to variationso the general time-reversible (GR) model o nucleotide substitution.

Morphological data collection

We examined 34 preserved specimens oBothriechisor this study (Appendix). Deni-tions o scale counts and morphological eatures ollow Campbell and Lamar (2004)and bilateral characters are reported as right/let. Unsexed juvenile specimens wereconsidered separately rom adult males and emales. For the holotype, we dissected andremoved the partially everted hemipenis at the base. We then lled the hemipenis withwarm water using a blunt-tipped syringe needle in order to attempt ull eversion. Wethen removed the water and injected hot petroleum jelly with blue wax-dye until nearmaximum expansion was achieved. Finally, we tied the hemipenes and stored them

in 70% ethanol. Tis procedure is modied rom that o Myers and Cadle (2003)and Zaher and Prudente (2003) and is urther described and illustrated in Smith andFerrari-Castro (2008) and Jadin and Smith (2010). Hemipenial terminology ollows


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Dowling and Savage (1960), Keogh (1999), and Savage (2002). Comparative morpho-logical data on related species was taken primarily rom Campbell and Lamar (2004)and Solrzano (2004). Color names and codes used in descriptions o coloration in lieare rom Khler (2012); color notes in lie were derived rom a series o photographso the holotype and paratypes.

Rsuls

Bayesian and Maximum Likelihood phylogenetic analyses produced congruent topo-logical results. Our phylogeny (Fig. 1) is generally congruent with those o Castoeet al. (2009) and Daza et al. (2010), recovering two clades o nominal Bothriechisschlegelii(one Mesoamerican, one rom Ecuador) rendered paraphyletic with respectto B. supraciliaris, and showing strong support or aB. marchiB. thalassinusclade andaB. auriferB. rowleyiclade, together orming aB. auriferB. bicolorB. marchiB.rowleyiB. thalassinusclade that geographically corresponds to Nuclear Central Amer-ica (Fig. 1). Both o our analyses recovered a weakly supported clade that includes

the Costa Rica/Panama taxaB. lateralisand B. nigroviridis, along with the Bothriechispopulation rom Reugio de Vida Silvestre exguat, Honduras. Within this primarilysouthern clade, nine samples rom Reugio de Vida Silvestre exguat show virtuallyno genetic divergence rom one another, and orm a monophyletic group with a well-supported (PP = 1.0; bs = 100) sister clade to B. lateralisrom Costa Rica and westernPanama (Fig. 1).

Based on the phylogenetic results, we examined morphological variation amongpopulations oBothriechis marchi sensu lato, which conrm the evolutionary and taxo-nomic distinctiveness o the exguat population as well as its apparent morphological

anity with a specimen rom Parque Nacional Pico Bonito, approximately 75 km tothe east o Reugio de Vida Silvestre exguat. We present the ollowing description othis relict northern lineage as a new species.

tabl 2. Results rom a priorimodel selections based onAkaikeinormation criterion (AIC) conductedin MrModeltest 2.2 (Nylander, 2004) or partitions o the dataset.

Partition Total characters Parsimony-inormative characters Best-ft model

ND4 1st pos 222 39 GR+ND4 2nd pos 222 9 GR+I

ND4 3rd pos 222 128 GR+Cyt b 1st pos 231 33 GR+Cyt b 2nd pos 231 12 HKY+I

Cyt b 3rd pos 231 134 GR+I+12S 409 60 GR+16S 495 39 GR+I


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Fgur 1. Phylogeny o palm-pitvipers (genus Bothriechis), showing strong support or a species-levelclade o the exguat population (Bothriechissp. n.) that is sister to B. lateralis. Te tree was estimatedrom a Bayesian 50% majority-rule consensus composed rom a concatenated mitochondrial dataset(ND4, cyt b, 12S, and 16S; total o 2263 bp). Numbers at nodes represent values o Bayesian posteriorprobabilities (PP, let) and Maximum Likelihood bootstraps (BS, right). Nodes supported by 95% PPand 70 BS are considered highly supported.


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Sysmacs

Bothriechis guifarroisp. n.urn:lsid:zoobank.org:act:1FC0661F-B08B-4D0D-85CC-DAEAB502D0DB

http://species-id.net/wiki/Bothriechis_guiarroiFigs 23, 56

Bothrops nigroviridis(in part): Meyer 1969: 420.Bothriechis marchi(in part): Campbell 1982: 381.Bothrops marchii(in part):Wilson and Meyer 1985: 120.

Holotype. UA R-60303 (Figs 2, 3), an adult male rom La Liberacon (Fig. 4A,C),15.5302N, 87.2939W (DD), 1,015 m elevation, Reugio de Vida Silvestre exguat,Departamento de Atlntida, Honduras, collected 25 July 2010 by the eld team oE. Aguilar, A. Contreras, L. Gray, L.A. Herrera-B., M. Medina-Flores, A. Portillo, A.Stubbs, and J. H. ownsend. Original eld number JH 3243. Genbank accessionnumbers: 16S (KC847259), cyt b (KC847275).

Paratypes (8): HONDURAS:Departamento de Atlntida: Reugio de Vida Sil-vestre exguat: adult emale (USNM 579875) collected 19 June 2010, two adultemales (USNM 57987677) collected 29 July 2010, and two unsexed neonatescollected 18 June 2010 (USNM 57987374), all rom Cerro El Chino (Fig. 4B),15.5225N, 87.2802W (DD), 1,3601,450 m elevation, southeast o La Liberacin.wo males (CM 156870 and MVZ 269305) collected 28 July 2010 rom a ridge-toptrail above La Liberacin, 15.5418N, 87.2891W (DD), 1,290 m elevation. One male(USNM 579878) collected 30 July 2010 rom La Liberacin (Fig. 4A, C), 15.5302N,87.2939W (DD), 1,015 m elevation.

Reerred specimens (4). HONDURAS: Departamento de Atlntida: AMNH46949 rom ela, collected sometime beore April 1932; USNM 319942 romQuebrada de Oro, Parque Nacional Pico Bonito. Departamento de Yoro: Reugio deVida Silvestre exguat: USNM 33748889 rom 2.5 airline km north-northeast o

La Fortuna. See Remarks.Defnition.Bothriechisguifarroiis distinguished rom all nine congeners by theollowing combination o eatures: dorsal scales in 19-19-15 rows; ventrals in males162166 (163.8), in emales 158166 (164.0), 162166 (164.0) in neonates; sub-caudals in males 6068 (63.0), in emales 6063 (61.0), 6268 (65.0) in neonates;intersupraoculars (37); superciliary scales absent; prelacunal scale used to secondsupralabial on both sides; two known color patterns in juveniles, one brown (with apale paraventral stripe and a series o short darker dorsal blotches and a dark brownpostocular stripe bordered by yellow on its lower edge) and the other green (with a

series o pale blue blotches and a deep blue postocular stripe bordered by pale blue onits lower edge); dorsal coloration in adults green with pale blue trim on anterior edgeso dorsal scales, and pale blue postocular stripe with green along the keels in center ostripe; and iris pale green, pale gray, or pale tan.
http://zoobank.org/?lsid=urn:lsid:zoobank.org:act:1FC0661F-B08B-4D0D-85CC-DAEAB502D0DBhttp://species-id.net/wiki/Bothriechis_guifarroihttp://species-id.net/wiki/Bothriechis_guifarroihttp://zoobank.org/?lsid=urn:lsid:zoobank.org:act:1FC0661F-B08B-4D0D-85CC-DAEAB502D0DB


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Fgur 2. Photographs in lie o the adult male holotype oBothriechis guifarroi(UA R-60303), withlateral and dorsal views o the head. Photographs by JH.

Diagnosis.Bothriechis guifarroican be distinguished rom the other members othe genus Bothriechisas ollows (B. guifarroieatures indicated rst, those or speciescompared next): B. aurifer(distributed at moderate and intermediate elevations romextreme east-central Chiapas, Mexico, to east-central Guatemala) adult color pat-tern (green vs. black-bordered yellow blotches on green background and prominentblack postocular stripe) and juvenile color pattern (green with pale blue blotches orbrown with pale paraventral stripe and dark dorsal blotches vs. pale lime green withblack-bordered yellow blotches); B. bicolor (occurring marginally at low upward to

intermediate elevations rom southeastern Chiapas, Mexico, to south-central Guate-mala) number o dorsal scales at midbody (19 vs. 21) and condition o prelacunaland second supralabial scales (used vs. separate); B. lateralis(moderate to marginallyhigh elevations rom northwestern Costa Rica to western Panama) number o dor-


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sal scale rows at midbody (19 vs. modal number o 23), adult color pattern (green vs.green with pale paravertebral bars and paraventral stripe), and juvenile color pattern(bi-morph pattern o green with blue dorsal blotching or brown with pale paraventralstripe and short unicolor dark blotches vs. uni-morph pattern o brown ground color

with pale paraventral stripe and short bicolor dark and pale blotches); B. marchi(oundmarginally at low elevations up to intermediate elevations in northwestern Hondu-ras and adjacent Guatemala) condition o prelacunal and second supralabial scales(used vs. separate), number o subcaudals in emales (6063 vs. 4657); B. nigroviridis(moderate to intermediate elevations rom north-central Costa Rica to west-centralPanama) adult color pattern (patternless green vs. green with very heavy blackmottling), juvenile color pattern (green with pale blue blotches or brown with paleparaventral stripe and a series o short darker dorsal blotches vs. green with heavyblack mottling), iris color (pale green, pale gray, or pale tan vs. almost black), numberso ventral scales in both sexes (162166 and 158166 vs. 143158 and 134158),numbers o subcaudal in both sexes (6068 and 6063 vs. 4956 and 4458), andcondition o prelacunal and second supralabial scales (used vs. separate); B. rowleyi(moderate to intermediate elevations rom extreme southeastern Oaxaca to northwest-ern Chiapas, Mexico) condition o prelacunal and second supralabial scales (usedvs. almost always separate), iris color (pale green, pale gray, or pale tan vs. yellow), andjuvenile color pattern (green with pale blue blotches or brown with pale paraventralstripe and a series o short darker dorsal blotches vs. pale yellowish green with brown orpurple dorsal blotches); B. schlegelii(low to intermediate elevations rom northwesternChiapas, Mexico, southward through Central America and into northwestern SouthAmerica as ar as extreme western Venezuela and extreme northern Peru) lack osuperciliary scales in the ormer and their presence in the latter, number o suprala-bials (1012, usually 10 vs. 710, usually 8), number o midbody dorsal scale rows(19 vs. 2125, usually 23), and adult color pattern (green vs. extremely variable colorand pattern involving ground color o yellow, pink, brown, gray, or green and dorsalblotching o a sizable array o colors, but sometimes absent; contrasting postocularstripe absent vs. present); B. supraciliaris (moderate to intermediate elevations rom

southwestern Costa Rica to west-central Panama) lack o superciliary scales in theormer and their presence in the latter, number o midbody dorsal scale rows (19 vs.2123, usually 23), number o ventral scales in both sexes (162166 and 158166 vs.145150 and 141148), number o subcaudal scales in both sexes (6068 and 6063vs. 4854 and 4552), and adult color pattern (green vs. extremely variable colorand pattern involving ground color o shades o green, brown, or maroon and dorsalblotching o an array o colors contrasting with that o the ground color; contrastingpostocular stripe absent vs. present); B. thalassinus(moderate to intermediate eleva-tions rom extreme eastern Guatemala and extreme northwestern El Salvador to west-

ern Honduras) number o midbody dorsal scale rows (19 vs. 2123, usually 21),and condition o prelacunal and second supralabial scales (used vs. separate).Description o holotype. An adult male (Figs 2, 3) with hemipenes partially

everted, let removed;rostral broader than high (4.38 3.26 mm); 2 internasals an-


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Fgur 3. Dorsal, lateral, and ventral aspects o the head o the holotype oBothriechis guifarroi(UAR-60303). Photographs by RCJ.


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teriorly; 2/2 canthals; 4 posterior intercanthals; supraoculars slightly more than twotimes as long as broad; 5 intersupraoculars; many scales on head o large size, includ-ing large, fat rontal and parietal scales; interrictals 25; single loreal, longer than high,bounded by upper two preoculars, canthal above, prelacunal and preoveals below,

and nasal; preoveals 3/3, suboveals 1/1; prelacunal and second supralabial used; pre-oculars 3/3, upper largest, middle large and in contact with supralacunal; suboculars2/2; postoculars 2/2; supralabials 10/10; mental broader than long (4.39 x 3.29 mm);inralabials 11/11; chin shields contacting rst our pairs o inralabials; gulars betweenchin shields and rst preventral 6/4; dorsal scale rows 19-19-15; preventrals 2; ventrals161; cloacal scute undivided; 65 undivided subcaudals; tail spine short and blunt.

Measurement o holotype.otal length 734 mm; tail length 136 mm, compris-ing 18.5% o total length; head 29.8 mm rom ront ace o rostral to posterior end omandible; head 19 mm at broadest point; neck 7 mm directly behind jaws.

Hemipenis description o holotype. Te partially everted let hemipenis o theholotype is described. Hemipenis at least 20 mm in total length and 13 mm in maxi-mum width at level o crotch; on sulcate side base with several rows o small spines(< 0.5 mm) ollowed by rows o larger spines and hooks extending or 5 mm, largestprotruding ca. 3.5 mm; asulcate side with minutely spined base up to 7 mm beorelevel o bilobation; numerous small mesial spines (< 0.5 mm) arranged in rows presentor 4 mm to the calyces, with peripheral section o each lobe containing nine spinesand hooks ( 2 mm), ve o which border lower rim o calyces; calyces ollow spinesand hooks distally; calyces scalloped, at least 10 rows at least 7 mm to apex o hemi-penis on asulcate side; sulcus spermaticus deep and biurcating ca. 4 mm beore siteo bilobation and extending upwards through spines and calyces likely to tip o eachlobe; sulcus spermaticus bordered by two, occasionally three, columns o minute scalesto the beginning o calyces, which orm the border likely to the apex o the lobes.Although the majority o the hemipenial characters o this specimen are reported, thelack o a ully everted hemipenis leaves inormation on the total length and the natureo the calyces incomplete.

Coloration o holotype in lie. Middorsal scales o the holotype Yellowish Spec-

trum Green (Color 128), ading to Light Grass Green (Color 109) laterally and be-coming Chartreuse (Color 89) ventrolaterally, with Medium Greenish Yellow (Color88) ventral scales; dorsal body scales edged anteriorly in Light Caribbean Blue (Color163), with up to approximately one-ourth o the anterior end o some scales edged inblue; skin concealed between dorsal scales Spectrum Violet (Color 186); postoccipitalstripe Light Caribbean Blue (Color 163), with keel and adjacent portion o three scalesthat lie within the postoccipital stripe Light Emerald Green (Color 142); terminal por-tion o tail Plumbeous (Color 295); iris Pale Bluish Gray (Color 287) with ne blackreticulations most heavily concentrated around the pupil.

Color pattern o holotype in preservative. Scales on dorsal suraces o the headand body blue-green, becoming more green laterally and yellow-green to yellow ven-trally. ail is mostly green with some grayish blue-green at the dorsal base. Pupil iscloudy and pale, surrounded by lime green iris heavily speckled with black.


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Fgur 4. Habitat in the vicinity o the type locality o Bothriechis guifarroi, La Liberacin, Reugiode Vida Silvestre exguat, Honduras; A riparian vegetation along the Ro Jilamito, 1,015 m elevationB small seepage pond near the top o Cerro El Chino, 1,380 m elevation C premontane rainorest withthe clearing aroundLa Liberacin (1,030 m elevation) visible in the oreground. Photographs by JH.

Variation in paratypes.We discuss scutellational variation in the three adult maleparatypes rst, the three adult emales next, and nally the two unsexed neonates. Scutel-lation varies as ollows (range ollowed by mean): ventrals (162166 [164.3], 158166[161.7], 162 and 166); subcaudals (6068 [64.0], 6063 [61.0], 62 and 68); ventrals +


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subcaudals (222233 [228.3], 221226 [222.7], 228 and 230); cloacal scute entire in allspecimens; dorsal scale row ormula 19-19-15, with the reduction to 15 rows occurringat ventrals 114162; supralabials (1011 [10.2], 1011 [10.2], 1010 and 1211);inralabials (1012 [11.0], 1013 [11.7], 1111 and 1211); preoculars 22 in all speci-

mens, except 3-3 in CM 156870; postoculars 22 in all specimens, except 32 in MVZ269305 and 44 in CM 156870; suboculars 23 [2.5], 23 [2.7], 22 and 34; relativetail length (0.1840.223 [0.199], 0. 1670.182 [0.176], 0.179 and 0.194).

wo juvenile color patterns are present in this species (Fig. 5), one we reer to asa green phase, the other as a brown phase. Both juvenile phases have distinctivelycolored tail-tips, presumably used in caudal luring, and well-dierentiated postocularstripes. Te green phase (USNM 579874) has a Chartreuse (Color 89) dorsal groundcolor with Light urquoise Green (Color 146) edging on the dorsal scales as well as ona series o irregular middorsal blotches, a Pale Green (Color 99) venter, and a Char-treuse (Color 89) head with a Jet Black (Color 300) postocular stripe, bordered aboveand below by Light urquoise Green (Color 146); tip o tail Cobalt Blue (Color 180);the iris is Pale Neutral Gray (Color 296) with ne darker reticulations. Te brownphase (USNM 579873) has a Robin Ruous (Color 29) ground color middorsally andanteriorly, becoming Salmon Color (Color 58) laterally and posteriorly, with a serieso irregular Ferrunginous (Color 35) middorsal blotches, a Pale Bu (Color 1) ventralsurace o head and venter becoming gradually darker (Pale Pinkish Bu [Color 3])posteriorly, a Dark Salmon Color (Color 59) head with a Chestnut (Color 30) postoc-ular stripe, bordered above and below by Light Bu (Color 2); tip o tail Sepia (Color286); a Pale Bu (Color 1) paraventral stripe is present on the lower hal o the rstdorsal scale row and the lateral edge o the ventrals; dark speckling along lateral edge othe ventrals; the iris is Chamois (Color 84) with ne darker reticulations.

Te type series oB. guifarroidemonstrates considerable variation in the condi-tion and shape o the scales on the dorsal surace o the head (Fig. 6), a characteristicoten considered diagnostic amongBothriechis(Campbell and Smith 2000, McCranie2011a). wo adult emales (USNM 579875 [Fig. 6C] and USNM 579877 [Fig. 6A]),one adult male (MVZ 269305 [Fig. 6B]), and one neonate (USNM 579873 [Fig. 6F])

all have multiple enlarged, unkeeled, plate-like scales present anterior to the posterior-most edge o the orbits; one adult emale ([Fig. 6D]), one adult male ([Fig. 6E]), andone neonate (Fig. 6G) all have smaller keeled scales present anterior to the posterior-most edge o the orbits. As a result o demonstrating essentially the ull range o dorsalhead scale conditions in the type series, we do not consider this characteristic to be ovalue in diagnosingB. guifarroirom other congeners.

Etymology. Te specic nameguifarroiis a patronym used to honor our colleagueand riend, Honduran environmental leader Mario Guiarro o Olancho. Don Marioearlessly led grassroots eorts to stop illegal logging in the indigenous awahka ter-

ritory o eastern Honduras, despite repeated assassination attempts and threats on hisown lie and those o his compatriots. Don Mario was murdered on 15 September2007, ironically Honduras Independence Day, while leading a mission to demarcatethe boundaries o the awahka-Asangni Biosphere and stave o urther illegal deor-


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Fgur 5. Paratypes oBothriechis guifarroiin lie; A USNM 579874, green-phase juvenile B USNM579873, brown-phase juvenile C close-up o head o USNM 579874 D close-up o head o USNM579873 e USNM 579875, emale paratype photographed in situ at Cerro El Chino, 1,420 m eleva-tion. Photographs by JH.

estation. On 21 July 2008, the only witness to Marios assassination, his son ShamirGuiarro Ramrez, was also murdered, along with Marios ather-in-law, Henry ArturoChacn, and mother-in-law, Nelda Ochoa, ater they were ollowed out o the city oJuticalpa by unknown assailants.

Distribution. Populations genetically conrmed to representBothriechisguifarroiare ound between 1,0151,450 m elevation in the western portion o the CordilleraNombre de Dios, Department o Atlntida, Honduras, within the boundaries o Reu-gio de Vida Silvestre exguat (Fig. 7). Tese localities lie within the Premontane Wet

Forest and peripherally in the Lower Montane Wet Forest ormations o Holdridge(1967; as applied by McCranie and Wilson 2002).Natural history. Te holotype was ound coiled at 2130h approximately 2.5 m

above the ground among old lea sheaths in the crown o a medium-sized understory


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Fgur 6. Variation in dorsal head scales among paratypes o Bothriechis guifarroi; A USNM 579877,adult emale B MVZ 269305, adult male C USNM 579876, adult emale D USNM 579875, adult emalee USNM 579878, adult male F USNM 579873, neonate G USNM 579874, neonate. Photographs by JH.

palm in gallery orest alongside the Ro Jilamito (Fig. 4A). Anurans o the generaDu-ellmanohylaand Ptychohylawere abundant in the immediate vicinity o the holotype.wo adult males (CM 156870 and MVZ 269305) were collected along a ridge on thenorth side o La Liberacon on the night o 28 July 2010. CM 156870 was active ona small tree rom 0.51.5 m above the ground; the second snake (MVZ 269305) wassitting coiled on the ground at the edge o the trail, and attempted to escape by crawl-ing across the path when we approached. In the immediate vicinity o these snakeswere numerous Craugastor rostralisactive on the ground and Bolitoglossasp. active onlow vegetation. wo neonates were collected on the same night (21002200 h) on 18

June 2010, and an adult emale (USNM 579875) was collected the next night, in anarea o eln orest at 1,380 m on the ridge called Cerro El Chino (Fig. 4B) above theremote ranch locality La Liberacin (at 1,030 m). Te brown-phase neonate (USNM579873) was ound atop a large, similarly-colored dead palm rond, while the green-phase neonate (USNM 579874) was sitting in essentially the same ambush positionas USNM 579873, but on top o a living green rond. Amphibian species collected inthe immediate vicinity oB. guifarroiinclude Bolitoglossasp., Nototriton sp., Craugas-tor rostralis, Plectrohyla chrysopleura, and Ptychohyla spinipollex. welve Bolitoglossasp.were encountered the same night as the two neonates, all while active on or around

dead and living palm ronds in the immediate vicinity o the neonates.Remarks. ownsend et al. (2012: 107) included a photograph o the holotypeoBothriechis guifarroias Bothriechis marchi. Bothriechis guifarroiis typically distin-guished rom B. marchiby having the prelacunal scale used to the second supralabial;


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however, one male paratype o B. guifarroi (CM 156870) has the right prelacunalseparated rom the second supralabial (they are used on the let side). Also, one speci-men oB. marchi(MCZ R-33335) rom the mountains west o San Pedro Sula alsohas used prelacunals and second supralabials on both sides, and another specimen(MCZ R-32030) rom La Cumbre has the let prelacunal separated rom the secondsupralabial (with them used on the right side).

We tentatively reer our additional specimens to Bothriechis guifarroi: two rom alocality on the leeward side o Reugio de Vida Silvestre exguat (USNM 33748889,rom 2.5 airline km NNE o La Fortuna, Dept. Yoro, 1,550 m elevation), one rom thecentral portion o the Cordillera Nombre de Dios (USNM 319942, rom Quebrada

Fgur 7. Geographic distribution o selected Bothriechisspecies and populations discussed in the text; locali-ties are based on data published herein and those o Campbell and Lamar (2004), McCranie (2011a), and Sav-age (2002); red diamond = type locality oB. guifarroiin Reugio de Vida Silvestre exguat; black diamond =reerred population oB. guifarroirom Parque Nacional Pico Bonito; circles = Bothriechis sp. inquirendapopu-

lations or the Sierra de Sulaco; squares = B. marchi; triangles = B. thalassinus, inverted triangles = B. lateralis.


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de Oro in Parque Nacional Pico Bonito, Dept. Atlntida, 1,090 m elevation), and onerom ela (AMNH 46949). All our specimens also have used prelacunals and secondsupralabials on both sides. USNM 319942 was collected as a juvenile and raised in cap-tivity (Wilson and McCranie 1992; McCranie 2011a), and exhibited a similar juvenile

color pattern as USNM 579873 beore undergoing an ontogenetic shit in coloration tothe bright green pattern exhibited by the type series oB. guifarroi. Campbell and Lamar(2004: plates 385386) also provided illustrations o juvenile B. marchi s.l. that demon-strated two color morphs similar to those exhibited byB. guifarroi; however these twoindividuals were captive born in the Houston Zoo and known only rom Honduras.

AMNH 46949 was collected sometime during or beore 1932 by Douglas Marcho the Lancetilla Serpentarium, just outside o the seaside city o ela. While theela locality is considered erroneous (Wilson and McCranie 1992), it is possible thatAMNH 46949 was obtained rom somewhere in the nearby western portion o theCordillera Nombre de Dios. Given that at least some highland taxa ound at both RVSexguat and Parque Nacional Pico Bonito are endemic sister species (e.g., Oedipina

gephyraand O. petiola; McCranie and ownsend 2011), we reer USNM 319942 toB. guifarroiwith the understanding that phylogenetic evaluation o the Pico Bonitopopulation might eventually show those animals to represent a distinct taxon.

Dscusson

Conservation status o Bothriechis guifarroi. With the description o Bothriechisguifarroi, there are now at least three species o palm pitvipers endemic to the ChortsHighlands, includingB. marchiand B. thalassinus, with the potential or additionalundescribed taxonomic diversity pending phylogenetic evaluation o allopatric popula-tions in central Honduras. Based on the IUCN Red List criteria (2012), B. guifarroishould be classied as Critically Endangered (B1ab[iii]+2ab[iii]) due to its limitedknown area o occurrence and the potential or anthropogenic damage to its habitat.According to the algorithm developed by Wilson and McCranie (2004a), we calcu-

lated the Environmental Vulnerability Score (EVS) or B.guifarroias 5+8+5=18, allo-cating it to the category o a high vulnerability species. Given this conservation status,B. guifarroibecomes the 48th member o the critically endangered endemic componento the Honduran herpetoauna (Wilson et al. 2012), and the tenth snake species andthe rst viperid species so designated. Tis species also warrants immediate considera-tion or protection under CIES, given its potential or exploitation in the pet trade.

Te vicinity o the type locality oB. guifarroiis part o the relatively large and intactpremontane rainorests and cloud orests o Reugio de Vida Silvestre exguat, oneo the most important areas o herpetoaunal endemism in Mesoamerica (ownsend

et al. 2012). While deorestation in the leeward portion o Reugio de Vida Silves-tre exguat has been documented since at least the early 1990s (see summary inownsend et al. 2010), ownsend et al. (2012) reported that the windward portion othe reserve contained a large intact expanse o virtually undisturbed orest. In late 2012,


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a plot was cleared in the upper Ro Jilamito watershed to the south o La Liberacin,marking the rst time armers rom adjacent Yoro had crossed into the Ro Jilamitowatershed and illegally cleared land (L. Herrera-B., pers. comm.). Tis is an ominousdevelopment, particularly in light o the recent drastic reduction in nancial support

or conservation eorts in Reugio de Vida Silvestre exguat, which had unded thetraining and employment o a team o local park guards during 20102012.

Herpetoaunal Endemism in the Chorts Highlands. Whereas Nuclear CentralAmerica has long been accepted as a region o high biodiversity and endemicity, someobservers have urther recognized the western and eastern portions o this highlandblock as distinct biogeographic entities (Johnson 1989; Campbell 1999; ownsend2006, 2009). Eastern Nuclear Central America has been shown to have a distinctivecomponent o endemic biodiversity, particularly in amphibians and reptiles (Wilsonand Johnson 2010); however, molecular characterization o evolutionary diversica-tion patterns in this region has been limited to a ew studies o a restricted taxonomicbreadth and broader geographic ocus (e.g. Castoe et al. 2009). Tis region is geo-graphically analogous to the Chorts Block, an allochthonousgeological ormation thatcurrently orms the only modern continental portion o the Caribbean ectonic Plateand the largest terrestrial segment o the contemporary Central American land bridge(Rogers 2003; Marshall 2007). Te Chorts Block has a challengingly complex his-tory, and recently has been the subject o increased ocus, and sometimes contentiousdebate, within the geological research community (James 2007; Mann et al. 2007;Ortega-Gutirrez et al. 2007; Silva-Romo 2008; Morn-Zenteno et al. 2009).

Te majority o the geographical extent o the Chorts Highlands is ound withinthe political boundaries o Honduras, the country with the highest degree o her-petoaunal endemism o any Central American nation (Wilson and Johnson 2010).ownsend and Wilson (2010) reported 91 endemic species (47 amphibians and 44reptiles) rom Honduras. Since that work went to press, an additional ten endemicspecies have been described rom Honduras, including three new plethodontid sala-manders (N. picucha, ownsend et al. 2011; N. tomamorum, ownsend et al. 2010;Oedipina petiola, McCranie and ownsend 2011), a new black iguana (Ctenosaura

praeocularis, Hasbn and Khler 2009), a new skink (Marisora roatanae; Hedges andConn 2012), two new dwar geckos (Sphaerodactylus guanajaeand S. leonardovaldesi;McCranie and Hedges 2012), and three new colubrid snakes (Omoadiphas cannula,McCranie and Cruz-Daz 2011; Tantilla psittaca, McCranie 2011b; Tantilla olympia,ownsend et al. 2013). With these species included, the total stands at 101 species,makingB. guifarroithe 102nd described herpetoaunal species endemic to Honduras.

Bothriechismarchi and the status o populations rom Yoro. We recognizeBothriechis marchisensu stricto as occurring in localities in the Cordillera de Merendnin the Honduran departments o Corts and Santa Brbara along the border with

Guatemala, as well as or at least one isolated locality in eastern Guatemala (Fig. 7).Tis Guatemalan locality, Cerro del Mono in Departamento de Zacapa, previouslywas the source o the only sequenced sample assigned to B. marchi(UA R-52959;Castoe and Parkinson 2006). Although the population o Bothriechis rom Cerro


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del Mono does not agree morphologically with the typical orm oB. marchi(E.N.Smith, pers. comm.; see Plates 422424 in Campbell and Lamar 2004, as B. thalas-sinus), the sequence data attributed to UA R-52959 are not notably divergent roma sample o typical B. marchi(MVZ 263604) collected rom the Sierra de Omoa in

northern Honduras (Fig. 1).Te type localities oBothriechismarchiand B. thalassinusare both in the Sierra

de Caral, within approximately 20 km o one another on opposite sides o the Guate-mala/Honduras border (Campbell and Lamar, 2004) suggesting that the two speciesoccur either in parapatry or sympatry in the limited orest remaining in that mountainrange (the type locality o B. marchi is not precise; Wilson and McCranie [1992]restricted it to the orested hills above El Oro, Departamento de Santa Brbara). Indi-viduals o both taxa in the Sierra de Caral exhibit a primarily green dorsal coloration,with some scattered bluish middorsal blotches (Campbell and Smith, 2000; Campbelland Lamar, 2004: plate 425). While the presence oB. thalassinushas not been con-rmed by vouchered specimens rom the Sierra de Omoa, the proximity o the Sierrade Caral and the Sierra de Omoa in northwestern Honduras and the similarity incoloration exhibited between these nominal taxa in that vicinity suggest the possibilitythat B. thalassinusmay have gone unnoticed in the Sierra de Omoa. We have evaluatedphotographs o over a dozen individuals o green Bothriechisrom the Sierra de Omoaencountered as part o an expedition-tourism operation in that area over the past veyears, and have noted considerable variation in head scalation in the photographs.Unortunately, none o the photographed individuals were collected nor were geneticsamples taken to allow or more detailed evaluation o the Bothriechiso the Sierra deOmoa. In addition to the nominal orm oB. marchi, it is likely that more than onespecies oBothriechisoccurs in sympatry or parapatry in the Sierra de Omoa, possiblyincludingB. thalassinusand/or an unidentied sister taxon o B. guifarroi. Focusedsampling and phylogenetic analysis oBothriechisrom the Sierra de Omoa is neededto better characterize the taxonomic diversity present in that mountain range.

Paraphyly in Bothriechis marchisensu lato in terms o populations rom the Cor-dillera de Merendn and the Cordillera Nombre de Dios, the latter now known to

represent B. guifarroi, calls into question the taxonomic status o populations rom iso-lated localities in the Sierra de Sulaco in Departmento de Yoro (Fig. 7). Tese popula-tions are represented in collections by one specimen rom Cerro de Pajarillos (USNM561085), three specimens rom the Montaa de Mataderos (FMNH 21777, MCZR-3878586), 14 specimens rom Portillo Grande near Montaa Macuzal (FMNH3473235, 35895, 35999601, 37217, 38542, 41621; MCZ R-3878991), and twospecimens rom Subirana Valley (FMNH 21892, MCZ R-38788). Four o thesespecimens examined by us or this paper (MCZ R-3878586, 3879091) dier romB. guifarroiin having 211915 dorsal scale rows (versus 191915) and having vary-

ing conditions o usion o the prelacunal and second supralabial (used on both sidesin MCZ R-38790, separate on both sides in MCZ R-38786, and used on one side andseparate on the other in MCZ R-38785 an R-38791). Given the considerable phyloge-netic diversication presented by analysis oB. guifarroiand B. marchi s. s., we cannot


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justiy assignment o the Yoro populations to either taxon in the absence o molecu-lar characterization o those populations. Tereore, we tentatively reer to the Yoropopulations as Bothriechis species inquirendapending collection o resh material andphylogenetic characterization. Eorts to secure this material are currently underway.

Evolutionary and biogeographic implications. Te phylogenetic position oB.guifarroihas signicant implications or our understanding o the biogeography andevolution o palm-pitvipers. Crother et al. (1992) rst presented a phylogenetic hy-pothesis or the genus Bothriechissupporting an eyelash pitviper clade (B. schlegeliiandB. supraciliaris) that is sister to a clade containing the remaining species oBothriechis,all o which have highland-associated distributions (typically


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mybp, based on the hypothesis that B. lateralisrepresented the sister lineage to theNuclear Central American clade. Given the discovery oB. guifarroi, the timing o di-vergence across the Nicaraguan Depression could be much more recent, similar to thatseen between Cerrophidion sasaiand C. wilsoni(3.066.03 mybp; Castoe et al. 2009).

Alternatively, it is possible that a Lower Central American Bothriechis(i.e., B. guifarroi,B. lateralis, B. nigroviridis) may have split rom Nuclear Central American taxa via theNicaraguan Depression earlier than suggested by Castoe et al. (2009) and Daza et al.(2010). Following additional sampling and the evaluation o populations presently re-erred to as B. sp. inquirenda, a reevaluation o Mesoamerican pitviper biogeography iswarranted, and may shed urther light on the complicated biogeographic relationshipbetween northern and southern Central America.

Acknowldgmns

Research was carried out under permits issued by ICF (Resolucin DE-MP-086-2010and Dictamen DVS-ICF-045-2010). We are indebted to the ollowing individuals andorganizations or supporting our work in Reugio de Vida Silvestre exguat: Allan J.Fuentes (PROLANSAE); Alcalde Adolo Pagoada-Saybe (Municipalidad de Arizona),Sad Lanez, Iris Acosta, Roberto Downing, Andrs Alegra (Instituto Nacional de Con-servacin y Desarrollo Forestal, reas Protegidas y Vida Silvestre [ICF]);guardabosquesErain Aguilar (San Jos de exguat), Alonso Contreras (Mezapita), and Alionso Portil-lo (Jilamito Nuevo); and Jos Dubn, majordomo at La Liberacin. We thank BenjaminK. Atkinson, Cesar A. Cerrato, Levi N. Gray, Luis A. Herrera, Paul House, Mayron Mc-Kewy Meja, Ciro Navarro-Umaa, Alexander L. Stubbs, and Hermes Vega-Rodrguezor their valued assistance and hard work in the eld during 2010. Fieldwork was sup-ported in part by a grant to Kirsten E. Nicholson (Central Michigan University; Na-tional Science Foundation DEB-0949359). For acilitating deposition o the type series,we thank Jonathan Campbell, Carl Franklin, and Eric Smith (UA), Roy McDiarmid,Steve Gotte, and James Poindexter (USNM), Carol Spencer, Jim McGuire, and ed Pa-

penuss (MVZ), and Jos Padial and Stephen Rogers (CM). We would also like to thankSean M. Rovito or sharing data and a tissue sample o aB. marchicollected rom theSierra de Omoa, and Eric N. Smith or sharing his insights about highland Bothriechisineastern Guatemala. William Brenneman (IUP) provided helpul comments on a drat othis manuscript, and Ileana Luque-Montes created the map used or Figure 7.

Rfrncs

Arvalo ES, Davis SK, Sites JW Jr. (1994) Mitochondrial DNA sequence divergence and phylo-genetic relationships among eight chromosome races o the Sceloporus grammicuscomplex(Phrynosomatidae) in central Mexico. Systematic Biology 43: 387418. doi: 10.1093/sysbio/43.3.387
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Appndx

Specimens examined; museum acronyms ollow Sabaj-Prez (2012).

Bothriechis guifarroi (12) Honduras: ALNIDA: Quebrada de Oro, USNM319942; ela, AMNH 46949; La Liberacin, CM 156870, MVZ 269305,USNM 57987378, UA R-60303. YORO: 2.5 airline km NNE o La Fortuna,USNM 33748889.

Bothriechis marchi(19) Honduras: CORS: near Corada, MCZ 28014; LaCumbre, AMNH 4695457, MCZ 3202931; Sierra de Omoa, MCZ 3333436, 3356064, USNM 83454. SANA BRBARA: Quimistn, MCZ 27260,27510; between Corada and Quimistn, MCZ 2756768, UMMZ 90677(was MCZ 27569); Santa Brbara, MCZ 28014.

Bothriechis thalassinus (3) Honduras: COPN: Quebrada Grande, KU 203094;OCOEPEQUE: 21.7 km east o Nuevo Ocotepeque, LSUMZ 23821; SANABRBARA: southeastern slope o Cerro Santa Barbara, LSUMZ 11638.

Bothriechis sp. inquirenda (4) Honduras: YORO: Montaas de Mataderos, MCZ3878586; Portillo Grande, MCZ R-3879091.

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Bothriechis guifarroi 2013