THE ANOLES (REPTILIA: SQUAMATA: DACTYLOIDAE: ANOLIS: NOROPS) OF HONDURAS. SYSTEMATICS, DISTRIBUTION, AND CONSERVATION JAMESR.MCCRANIE1ANDGUNTHERKO¨HLER2 CONTENTS Abstract 1 recentphylogeneticanalysessuggestingeightgeneraof ---------------------------------------------------------------------------- Introduction 2 anolesarerecognizable,twoofwhichoccurinHonduras -------------------------------------------------------------------- MaterialsandMethods 3 (Anolis and Norops). Each species account contains a ----------------------------------------------- A Brief History of the Study of the Anole synonymy, statements on its geographic distribution, LizardsinHonduras 8 description,diagnosis/similarspeciesstatements,alistof ----------------------------------------------- SystematicSpeciesAccounts 11 illustrations,aremarkssection,naturalhistorycomments, ------------------------------------ FamilyDactyloidae 11 anetymologysection,andalistofspecimensexamined; ------------------------------------------------- GenusAnolis 11 forafewspecies,alistofotherrecordsisincluded.The ------------------------------------------------------------- GenusNorops 17 synonymyforeachspeciesincludestheoriginaldescrip- ----------------------------------------------------------- Key to the Honduran Species of the Genera tionofthespecieswiththeoriginalproposedcombina- AnolisandNorops 221 tion, the type specimen(s), and direct quotes of the -------------------------------------------------- Clave para las Especies Honduren˜as de los species’ type locality. Each synonymy also includes the Ge´nerosAnolisyNorops 235 first use of the currently used scientific name and all -------------------------------------- Discussion of Honduran Anole Species Rela- known references to Honduran specimens and/or tionships 255 localities. Distribution mapswith the known Honduran --------------------------------------------------------------------- DistributionofAnolesinHonduras 257 localitiesplottedareincludedforeachspecies(withone ---------------------- DistributionwithinDepartments 257 exception), andother records are includedon a fewof ---------------------- DistributionwithinForestFormationsand thosemaps.Colorphotographsofanadultofeachspecies byElevation 259 are included as well as color photographs of the male --------------------------------------------------------- DistributionwithinPhysiographicRegions 262 dewlapofallspecies.Followingthespeciesaccountsare -------- DistributionwithinEcophysiographicAreas 262 twodichotomousidentificationkeys(EnglishandSpanish ------- BroadPatternsofGeographicDistribution 263 versions) to help the reader identify any specimen in ------- HistoricalUnitsofHonduranAnoleGenera hand. Following the identification keys is a section on andSpecies 264 speciesgrouprelationshipsofeachspeciesknownfrom ----------------------------------------------------------- HondurasasaDistributionalEndpoint 269 the country. A distribution section that contains the ---------- ConservationStatusofAnolesofHonduras 270 distribution of each anole species in Honduras by ------- VulnerabilityGauges 270 department, forest formation, elevation, physiographic --------------------------------------------- IUCNRedListCategories 272 region, and ecophysiographic area; broad patterns of ---------------------------------- AnoleSpeciesOccurringinProtectedAreas 272 geographicdistribution;historicalunits;andadiscussion ------- Acknowledgments 278 of Honduras as a distributional endpoint. A section on -------------------------------------------------------- LiteratureCited 279 conservation discusses vulnerability gauges, IUCN Red ----------------------------------------------------------- List categories, and a section detailing each species’ occurrenceinHonduranprotectedareas. ABSTRACT. In this review, we provide thorough descriptions of the 39 named species of the family Keywords: Anoles,Dactyloidae,Anolis,Norops,Hon- Dactyloidae (commonly referred to as anoles) we duras,Taxonomy,Systematics,Distribution,Conservation recognize as occurring in Honduras. We follow two Some years ago—I am unable to date the 110770SW164thStreet,Miami,Florida33157-2933. eventmore accurately—wordseems tohave Authorforcorrespondence([email protected]). been passed among the herpetological 2Forschungsinstitut und Naturmuseum Sencken- fraternity that I knew something about berg, Sektion Herpetologie, Senckenberganlage 25, D-60325FrankfurtamMain,Germany. anoles. How the rumor was started, I am Bull. Mus. Comp. Zool., SPS(1): 1–292, February, 2015 1 2 Bulletin of the Museum of Comparative Zoology, Special Publications Series, No. 1 unable to say, as at that time I had no specimens. Some anole species are difficult enemies who would have stooped so low in to distinguish from similar species on the retaliation for some imaginary ill-treatment basis of preserved specimens alone (i.e., at my hands. Notwithstanding, the rumor Noropslimifrons from N. zeus, N. rodrigue- grew rapidly, with the result that since that zii from N. yoroensis). McCranie has had timeagreatmajorityof theanolescollected extensive fieldwork in Honduras for about in northern Central America and in much 38yearsandduringthattimehascollectedand of Mexico have passed through my hands. photographed all anole species recognized in From these I have learned something of this work. Ko¨hler also has undertaken field- variation in the genus, but the more work from Mexico to Panama over the last specimens I have examined the more 18yearsinanefforttostudyanolesystematics. convinced I have become that the rumor The fieldwork studies by us are used as a as to my knowledge is without substantial background to identify the anole species and foundation. I take this opportunity, there- their distributions within Honduras. Finally, fore, to present the few data that I have tissues of 13 species collected by the first collected over the years on Guatemalan author in Honduras were used in the recent anoles, which should definitely silence it phylogenetic study of Nicholson et al. (2012), (Laurence Cooper Stuart, 1955: 1). and tissues of 20 other species collected in The trouble with these anoles … is that no HondurasbyMcCranieawaitstudy. one sticks to the job long enough. If you Anoles have a wide geographical distri- will make a real review of the Guatemalan bution, occurring in the southeastern Unit- ones, I have the feeling that the rest of the ed States, throughout the Bahamas, the Central American ones will fall into line Greater and Lesser Antilles, and from without so much effort (Karl Patterson northern Mexico into South America as far Schmidt in Stuart, 1955: 1). as southeastern Peru, south-central Para- guay, and southeastern Brazil (and many islands close to the mainland). Anoles also INTRODUCTION have a wide elevational distribution occur- Identifying anoles, especially preserved ring from sea level to about 3,750 m. It is specimens, can be very challenging, as so common to find several species occurring well expressed by Stuart’s quote. Few syntopically. In those cases, ecological herpetologists up to Stuart’s time were segregation between species is usually willing to take the time to tackle systematic evident. Niche partitioning in anoles has studies of anoles (as expressed in Schmidt’s been documented with respect to perch quote) occurring in their geographical area height, type of vegetation, body size, and of expertise. Thus, our knowledge of the timing of foraging activities, particularly in systematics of anoles on the mainland has the West Indies (reviewed in Losos, 2009). lagged far behind those of other speciose On the other hand, niche partitioning of lizardgroups(i.e.,Sceloporus).Additionally, anolespeciesoccurringonthemainlandhas fieldwork and examining living anole spec- been relatively poorly studied. imens is essential to gaining good knowl- Despite their wide occurrence, and the edge of this lizard group. Recording color wishful thinking of Stuart and Schmidt as notes in life and photographing living quoted above, the systematics of anole specimens (especially that of the male species remains poorly known from much dewlap) to go along with color notes is of their mainland range, including much of extremely valuable to improving one’s Central America. In recent years, Savage knowledge of some of the most important (2002) studied the anole species occurring characters of any given species. One cannot in Costa Rica and Ko¨hler et al. (2005b) overstress the importance of fieldwork and studied those occurring in El Salvador. thatofrecordingexactlocalitydataforanole Additionally, Ko¨hler and Acevedo (2004) THE ANOLESOF HONDURAS N McCranie and Ko¨hler 3 studied those species occurring at low and in England. We have also examined select moderate elevations on the Pacific versant specimens in several other museums world- of Guatemala. wide. Much variation exists in the number Meyer and Wilson (1973) provided a ofspecimensavailableforeachspecies,with checklist of the lizard species known to numbers varying from only two for Norops occur in Honduras that included 15 species carpenteri to 981 specimens for N. tropido- of anoles and 646 total anole specimens notus.Abbreviationsformuseumcollections listed in their locality records. The 40 years follow those of Leviton et al. (1985) and sincetheMeyerandWilson(1973)checklist Leviton and Gibbs (1988). Terminology have seen much additional fieldwork in for hemipenial morphology follows that Honduras, including numerous cloud forest of Myers et al. (1993) and Savage (1997). andmid-elevationbroadleafforestlocalities. Characters used in the species descriptions Meyer and Wilson almost entirely limited generallyfollowthoseusedbySmith(1967), their fieldwork to low elevation localities Williams et al. (1995), and especially the withaccessbyroads.Subsequentfieldwork, recent detailed methods of Ko¨hler (2014). especially that in isolated cloud forest Snout-ventlength(inmm)wasmeasuredusing localities,hasdiscoveredseveralnewspecies. calipers for most species, except for larger Additional fieldwork at low- and moderate- species in which a ruler was used. All other elevation localities has documented several measurementsweremadeusingcalipersand range extensions of species into Honduras were rounded to the nearest 0.1 mm. Head from nearby countries. More recently, de- length was measured from the tip of the tailed studies of several wide-ranging ‘‘spe- snout to the anterior margin of the ear cies’’ have demonstrated those species to opening. Tail height and width were mea- represent complexes composed of several suredatthepointreachedbytheheelofthe cryptic taxa (i.e., Ko¨hler, 2010; Ko¨hler and extended hind leg. Dorsal andventral scales Vesely, 2010), a few of which occur in werecountedalongthemidlineofthebody. Honduras. This report presents a systematic Methods for counting male dewlap scales reviewofthe39namedanolespeciesknown follow Fitch and Hillis (1984). Subdigital to occur in Honduras, includingthe Islasde lamellae were counted on phalanges ii–iv laBahı´a,IslasdelCisne,andthesmallislands of the fourth toe of each hind limb. in the Golfo de Fonseca. Abbreviations used in the species descrip- tions are: HL (head length); SHL (shank MATERIALS AND METHODS [tibia]length);SVL(snout-ventlength);TAL For this study we examined 5,150 spec- (tail length); TH (tail height); TW (tail imens (plus 34 skeletons, four cleared and width), and TOL (total length). The capital- stained [C&S] specimens, and four eggs) ized colors and color codes (the latter in of Honduran anoles, which represent the parentheses) are those of Smithe (1975– vast majority of those in museums world- 1981), except in a few recent instances in wide. A list of the specimens examined is which Ko¨hler (2012) was used. The latter is provided in the respective species accounts. noted when used. The numbers of specimens given for each Species synonymies contain the original species by Meyer and Wilson (1973) are description of each species, including the included in brackets in our Specimens museum number(s) for the type speci- Examined section following the number men(s), the type locality as originally stated, we examined for this revision. We have the first use of the name combination tried to examine all known Honduran recognizedherein,andallknownreferences specimens housed in U.S. museums, and to Honduran specimens or localities. The those of the Senckenberg Museum in majorityofHonduranlocalitieslistedherein Germany and the Natural History Museum are included in the Gazetteer in McCranie 4 Bulletin of the Museum of Comparative Zoology, Special Publications Series, No. 1 (2011), and the newer ones will be pub- localities and type specimens, pagination, lished in McCranie (in preparation). and titles of the literature. The dichotomous keys are meant to be This publication includes results of field- significantly detailed to allow the user to work and examination of the known litera- identify any anole in hand to species. ture through 31 June 2013. However, in some cases it might be necessary to consult the photographs, diag- Justification for the Use of the Generic nosis/similar species sections, and/or the Name Norops descriptions of the species in question. Modern studies of the systematics of the Figures are included with the keys to help anole lizards had its beginnings with Ethe- the user. ridge (1959), who placed all anole species General geographical statements of the known to him into two groups based on overall known distributions of each genus condition of the caudal vertebrae. The beta and species are included. The elevational group‘‘ischaracterizedbythepresenceofa statements of Stuart (1963) are slightly pair of long, bifurcate, forward-directed modified and used in those statements (low elevations 5 sea level to 600 m; moderate processes’’ on the autotomic caudal verte- elevations5601–1,500m;andintermediate brae (Etheridge, 1959: 132), with the alpha elevations 5 1,501–2,700 m). The natural section lacking those processes. The Ethe- history comments contain information on ridge study was confined to mostly skeletal what is known about Honduran species in characteristics; thus, Etheridge (1959) did the field, and for the nonendemic species, not study scale characteristics. generalhabitatinformationfromoutsidethe Williams (1976a) studiedtheWestIndian country is also included based on the anoles and placed those lizards into three literature. Notes from the literature are also genera (Chamaeleolis Cocteau in Sagra, included on diet and reproduction when 1838; Chamaelinorops Schmidt, 1919; and available.Forestformationsusedareslightly Anolis Daudin, 1802). Anolis was further modified from those used by Holdridge divided into two groups, the alpha and beta (1967). All information in the Natural sections, following the work of Etheridge History Comments is based on Honduran (1959). Williams (1976a) also relied almost specimens unlessotherwisenoted. completely on skeletal features; no mention Distribution maps are included for each was made of scale data. Williams (1976b) species. Closedsymbols on all maps(except summarized his concepts of the species thatofNoropsnelsoni)representspecimens groups of South American anoles, again examined, whereas an open symbol repre- relying heavily on the alpha and beta caudal sents a reliable record listed in the other vertebraedivisionofEtheridge(1959).Scale records section. A single symbol may data were again almost completely ignored represent more than one nearby locality. because, in his opinion, ‘‘There is unfortu- Shadingtoillustratethegeneraldistribution nately no external characters by which is not used on those maps because of the thetwosections[alpha-beta]canbeseparat- complex topography of Honduras. ed’’ (Williams, 1976b: 263). Williams et al. Colorphotographsinlifeareincludedfor (1995) developed a protocol they titled ‘‘A all recognized species. Dewlap photographs computer approach to the comparison and are also included for males of all species. identification of species in different taxo- The specimen photographed in each case is nomicgroups.’’Williamsetal.(1995)diduse noted in the figure legend, as is the person some scale data, but as the title suggests, who took the photograph. those data were not thought to be informa- All literature cited in this work was tive above the species level. As a result, few examined to verify the correct spelling of scientists currently working on upper level scientific names, the correct citation of type anolesystematicsusescaledataintheirwork. THE ANOLESOF HONDURAS N McCranie and Ko¨hler 5 Poe (2004) used a few of the Williams et al. ally, both of us have submitted manuscripts (1995) scale characters in his phylogenetic using Norops, one of the genera recognized analysisoftheanoles,butnonethatinvolved by Guyer and Savage for most of the beta countingof more than ca. 10 scales. anolesofEtheridge(1959),butthenamewas Guyer and Savage (1987) attempted a changed to Anolis by the editors of those cladistic analysis (again, external morpholo- scientific journals. gy was almost completelyignored)ofAnolis Other authors (see Glor et al., 2001; (sensu lato) and recognized five clades as Nicholson, 2002; Poe, 2004; Nicholson et genera (Anolis; Ctenonotus Fitzinger, 1843; al., 2005, 2012; Alfo¨ldi et al., 2011) have Dactyloa Wagler, 1830; Norops Wagler, performed phylogenetic analyses that dem- 1830; and Semiurus Fitzinger, 1843 [Savage onstrated Norops, the beta anoles of Ethe- and Guyer, 1991 noted that Semiurus is a ridge(1959),toformaclade.However,some junior synonym of Xiphosurus Fitzinger, of these workers question the monophyly of 1826]). With that proposal, those authors at leastsome of the other proposedgenera. suggested that eight anole genera be recog- Recently, Nicholson et al. (2012) pro- nized (those just listed plus Chamaeleolis; duced a new phylogenetic analysis of the Chamaelinorops; and Phenacosaurus Bar- anoles, again almost completely ignoring bour, 1920; but see below). However, the scale data. Those authors also recovered Guyer and Savage efforts were met with eight clades (although not the exact same strong opposition. Williams (1989) bluntly eight clades as recovered by Guyer and criticized the databases of the Guyer and Savage, 1987, 1992) they recognized as Savage(1987)analysisandCannatellaandde genera (Anolis; Audantia Cochran, 1934; Queiroz (1989) criticized the Guyer and Chamaelinorops; Ctenonotus; Dactyloa; Savage analytical procedures plus their Deiroptyx Fitzinger, 1843; Norops; and osteological categorizations. Guyer and Sav- Xiphosurus). Nicholson et al. (2012) com- age (1992) argued point by point to the bined previously published morphological critiques by Cannatella and de Queiroz and molecular data, along with new molec- (1989) and Williams (1989), added more ular data from four anole species. Pyron et data (no scale data, however) to their al. (2013: 15) provided the next phylogenet- analysis, and generated a new phylogenetic ic analysis, which was based entirely on studythatrecovered‘‘essentiallythesameset moleculardata,withtheirresultssupporting of relationships’’ as they recovered in their ‘‘the monophyly of all the genera recog- 1987publication(Crother,1999:292).Curi- nized’’ by Nicholson et al. (2012). Losos ously, there were no published responses to (2013, 2014) noted two exceptions, one of thenewGuyerandSavage(1992)effortsand which was the result of Nicholson et al. data sets. Crother (1999: 291) pointed out (2012)havinglistedonespecies(christophei ‘‘the 1992 work by Guyer and Savage is Williams, 1960, of Haiti) in two genera in almost never cited’’ even by workers citing their tables (Chamaelinorops and Xipho- and rejecting the Guyer and Savage (1987) surus). The most recent phylogenetic anal- study. Even Poe (2013), in his rejection of ysis of the anoles is that of Gamble et al. theNicholsonetal.(2012)proposedgeneric (2014; again, without any scale data). Losos scheme, referred to the Guyer and Savage (2014) reported that three of the 216 (1987;dated1986)publicationseveraltimes, species used by Gamble et al. (2014; in but did not cite Guyer and Savage (1992). addition to the error made by Nicholson Based on these types of responses, Crother with christophei noted above) fell in differ- (1999: 291) asked if the Guyer and Savage ent clades than they did in the Nicholson ‘‘studiesandsubsequentclassification[have] et al. (2012) study. As a result, those four been given a fair scientific treatment?’’ As a species make three of the Nicholson et al. result, few workers on anoles have followed generanonmonophyletic.Thetwogeneraof the Guyer and Savage taxonomy. Addition- Nicholson et al. (2012) occurring on the 6 Bulletin of the Museum of Comparative Zoology, Special Publications Series, No. 1 Latin American mainland (Dactyloa and done.OurargumenttorecognizeNoropsas Norops; Anolis allisoni Barbour, 1928, and a valid genus is supported by all phyloge- Ctenonotus cristatellus [Dume´ril and Bi- netic analyses done to date that were based bron, 1837] have been recently introduced entirely on, or nearly completely on, molec- by man to at least one Central American ulardata.Theuniquetailstructurefoundin mainland locality each) remain monophy- Noropsalsohelpstosupportamonophyletic letic in the Pyron et al. (2013) and Gamble Norops. Those two independent types of et al. (2014) studies. evidencestronglyinfluencedourdecisionto Poe(2013)respondedtotheNicholsonet elevate Norops to a full genus despite the al. (2012) study and concluded that ‘‘most’’ concerns expressed by Poe (2013) and of their proposed genera are not monophy- others. letic. Thesubsequent studies ofPyron etal. OneofourconcernswiththeNicholsonet (2013) and Gamble et al. (2014; also see al.(2012),andallotherphylogeneticanalyses Losos, 2013, 2014) have shown that ‘‘most’’ oftheanolesconductedtodate,isthatscale of the Nicholson et al. genera are, in fact, data,includingdifficultandtime-consuming monophyletic using only molecular data. scale counts, have been presumed to be Thus, the ‘‘most’’ statement used several uninformative above the species level and, times by Poe (2013) is not supported bythe thus, ignored. Ignoring a type of data that is literature. Poe’s (2013) concern about not presumed,butnotproven,tobeuninforma- enoughtaxaandnotenoughcharactersused tive is not a sound scientific approach. in the Nicholson et al. (2012) study is Unfortunately, only two of the eight genera certainly correct. Simply put, more species (clades)recoveredbyNicholsonetal.(2012) and more characters need to be added to are represented in Honduras (Anolis and the databases of future phylogenetic analy- Norops),withAnolishavingonlyonespecies ses of the anoles. Despite those shortcom- inthecountry.Thus,weareunabletoapply ings, we recognize two genera (Anolis and scale morphology to those Nicholson et al. Norops) for the anoles occurring in Hon- clades in this study. Adalsteinsson et al. duras. In doing so, we realize that oppo- (2009), Hedges et al. (2009, 2014), Harvey nents of recognizing more than a single et al. (2012), and Hedges and Conn (2012) genusofanoles,withabout400species,will have recently incorporated time-consuming continue to argue that there is no compel- scalecounts,inmostcasesinassociationwith ling need to divide a monophyletic Anolis molecular data, in their generic divisions of intoeightgenera.However,wefeelthatthis several snake (Leptotyphlopidae, Typhlo- conservative approach is misguided. As far pidae,andDipsadidae)andlizard(Scincidae as is known, the anterolaterally directed and Teiidae) taxa, so why are we ignoring transverse process (see Etheridge, 1967, these types of data in anole phylogenetic fig. 3; Guyer and Savage, 1987, fig. 1) on studies? We have given above several the autotomic caudal vertebrae of Norops is examples of scale data having been largely ‘‘unique in structure among all lizards’’ ignored in upper level anole systematic (emphasis ours; Guyer and Savage, 1992: studies because we believe we have been 105). We argue that Norops is so distinct ignoringpotentiallyinformativedata.Oneof that it warrants its own genus (other similar the clades, along with Norops, that renders examples probably also exist, i.e., Chamae- the remaining ‘‘Anolis’’ paraphyletic is Dac- linorops barbouri Schmidt, 1919, for which tyloa, the only other anole clade occurring Schmidt proposed the new genus Chamae- naturally on the Latin American mainland. linorops, in part because of unique osteo- Individuals of that clade look so different logical characters in that form). If the fromtheremaininganolesthatitseemslikely consequence of our action renders Anolis thatasuiteofscaleandothermorphological nonmonophyletic, then that stresses the characters can be found to distinguish point that additional work does need to be Dactyloa from the remaining anoles, in THE ANOLESOF HONDURAS N McCranie and Ko¨hler 7 addition to the molecular studies that have Ecomorphs, Ecomodes, or Just alreadyaccomplishedthat(Castan˜edaandde Niche Partitioning Queiroz, 2011 [also see Castan˜eda and de Nichepartitioninginanoles,especiallyon Queiroz, 2013, who added osteological and Caribbean Islands, has been documented some basic scale data to their molecular with respect to perch height, vegetation analysis];Nicholsonetal.,2012;Pyronetal., types, body size, and timing of foraging 2013;Gambleetal.,2014).Weareinnoway activities. The wide variety of behavioral suggesting that everyone is compelled to traits exhibited by anole species has been follow our suggested recognition of Norops demonstrated to have a strong correlation as a genus. We are asking, however, that with morphology among West Indian spe- workersinvolvedinfuturetaxonomicstudies cies regardless of phylogenetic relation- ofanolesaddscalecountsandconfigurations ships. Thus, West Indian species having andskeletaldatasetstotheirmoleculardata the same or similar ecological roles have bases before performingadditional phyloge- been characterized as ‘‘ecomorphs’’ (re- netic studies. We are also hoping that with viewed in Losos, 2009). On the other hand, time(the‘‘revisionshock’’ofHedges,2013), Schaad and Poe (2010; also see Irschick et theanolecommunitywillacceptNoropsand al., 1997) were able to assign only 15 of the other anole clades as valid genera in the 123 mainland species they analyzed to a Linnaean taxonomic scheme. Also, any West Indian ecomorph. Nicholson et al. worker who disagrees with our suggested (2012) introduced the term ‘‘ecomode’’ to Linnaean taxonomy could apply the Phylo- replace ecomorph for the mainland mem- Codetoourconclusions(seedeQueirozand bers of the Dactyloidae; however, those Cantino, 2001, for an introduction to that authors did not adequately define their nomenclatural system). intended meaning of that term. Based After the above was finished and sent to largely on habitat statements in the litera- copyediting, Nicholson et al. (2014) pub- ture, Nicholson et al. (2012) placed a large lished a response to Poe’s (2013) critique of number of mainland species of anoles into their 2012 revision of anole classification. one of their eight ecomodes (those authors Those authors concluded that Poe made also recognized a polymodal category for several errors and misrepresentations in his species they thought regularly occurred in critique. Nicholson et al. (2014) discussed more than one of their eight categories). point-by-point each of Poe’s critiques and Returningtotheecomorphcategoryusedso statedthattheyhadexplainedtheirpositions in their 2012 publication regarding the few successfully for West Indian anoles, Losos ‘‘unstable’’ species that Poe used to call the (2012) noted that those ecomorph assign- Nicholson et al. (2012) work flawed. Nichol- ments are objective because the existing son et al. (2014: 109) also stated that Poe quantitative data are testable statistically. (2013)‘‘missedtheopportunitytopresentan On the other hand, the Nicholson et al. alternative comprehensive taxonomy to re- (2012) ecomode assignments are subjective place the one against which he argues so decisions based on literature statements strenuously.’’ We completely agree with that varying from short, limited summaries to statementandwiththeNicholsonetal.(2014: well-studied species. Most anole species 109) statement that commentary regarding also use a variety of habitats. Thus, where published taxonomic revisions should be does one draw the line to distinguish one ‘‘constructive, objective, and scientifically ecomode from another (i.e., trunk anole accurate.’’ Nicholson et al. (2014) concluded versus trunk-ground anole)? that Poe’s (2013) report was lacking in those AlthoughweapplaudtheNicholsonetal. points.Timewilltellifthescientificcommu- (2012) efforts to classify mainland anole nity adopts or rejects the Nicholson et al. species into habitat categories, we think the (2012, 2014) reclassificationof theanoles. data for the vast majority of the mainland 8 Bulletin of the Museum of Comparative Zoology, Special Publications Series, No. 1 species are inadequate at present. Also, genera Polychrus and Anolis. Frost et al. based on our own fieldwork, many anole (2001) did not follow the suggested parti- species show such extensive variation in tioningofAnolisintoeightgenerabyGuyer habitat preference and ‘‘ecomode’’ variation and Savage (1987). Subsequently, Town- that it is not possible to place many species send et al. (2011) performed a phylogenetic into one of the eight ecomodes recognized analysis of iguanian lizards based on molec- byNicholsonetal.(2012).Therefore,wedo ular data and recovered a polyphyletic nottrytoforceeachanolespeciesoccurring Polychrotidae. To rectify the polyphyletic inHondurasintooneoftheNicholsonetal. Polychrotidae, Townsend et al. (2011) re- (2012) ecomode categories; rather, we just strictedthefamilyPolychrotidaetothegenus give statements about where we have Polychrus and placed Anolis (sensu lato) in observedindividualsofeachspeciesrelative the family Dactyloidae Fitzinger (1843; type to their available habitats. Statements of genus Dactyloa Wagler, 1830, family name habitatusefromtheliteraturearealsogiven originally spelled Dactyloae by Fitzinger, for most nonendemic species. 1843). Within the Dactyloidae, two of the eight genera, Anolis and Norops, occur in Environment of Honduras Honduras. The type species of the former genusisA.carolinensisVoigt,1832,aspecies McCranie (2011) recently discussed the nativetothesoutheasternUnitedStates,and general description, physiography, climate, that of the latter genus is N. auratus and forest formations of Honduras. Inter- Bonnaterre, 1789, a species that occurs in ested readers are referred to that book for South America and lower Central America. information on those subjects (also see The year 1834 saw the first descriptions distribution of anoles in Honduras herein of any anole species that occurs in Hon- for reproductions of several pertinent maps duras. Wiegmann (1834) described Norops taken from McCranie [2011]). biporcatus and N. laeviventris (both as Dactyloa), with the former having its type A BRIEF HISTORY OF THE STUDY OF locality in Guatemala and the latter in THE ANOLE LIZARDS IN HONDURAS Mexico. Fifteen of the anole species occur- Formuchofthe20thcentury,anolesand ringinHondurasweredescribedduringthe their close relatives were placed in the 1800s (Table 1), none of which have their familyIguanidaeGray(1827:56).However, type localities in Honduras. The first four Frost and Etheridge (1989) performed a anole species with type localities occurring phylogenetic analysis of iguanian lizards in Honduras were described during the based on morphology and divided the early 1900s. Barbour (1914) described N. former Iguanidae into several families. The nelsoni (as Anolis) from the Islas del Cisne, phylogeny recovered by Frost and Ethe- GraciasaDios;Barbour(1928)describedA. ridge (1989) suggested that the ‘‘anoloid allisoni from Isla de Roata´n, Islas de la iguanid lizards’’ of Etheridge and Williams Bah´ıa; Dunn and Emlen (1932) described (1985: 1) formed a monophyletic lineage N. sminthus (as Anolis) from what is now that they placed in the family Polychridae partofParqueNacionalLaTigra,Francisco (5 Polychrotidae) Fitzinger (1843, type Moraza´n; and Schmidt (1936) described N. genus Polychrus Cuvier, 1816). Frost et al. loveridgei (as Anolis) from Portillo Grande, (2001) performed a new phylogenetic anal- Yoro. Three other species of anoles occur- ysisofthepolychrotidlizardsbasedonboth ringinHondurasweredescribedduringthe morphological and molecular data that period from the late 1930s to the early demonstrated the Polychrotidae recovered 1970s. These are N. wellbornae Ahl (1939; intheFrostandEtheridge(1989)studywas as Anolis ustus wellbornae, type locality in not monophyletic. Thus, Frost et al. (2001) El Salvador), N. heteropholidotus Mertens restricted the family Polychrotidae to the (1952a; as Anolis, type locality in El THE ANOLESOF HONDURAS N McCranie and Ko¨hler 9 TABLE1. ALISTINGOFTHEORIGINALDESCRIPTIONSOFTHEHONDURANSPECIESOFANOLISANDNOROPS,AUTHORS,ANDABBREVIATED TYPELOCALITIES. OriginalName Author(s) AbbreviatedTypeLocality Anolisallisoni Barbour,1928 IsladeRoata´n,Honduras Noropsamplisquamosus McCranie,Wilson,andWilliams,1992 ElCusuco,Honduras Anolisbeckeri Boulenger,1882 Yucata´n,Mexico Noropsbicaorum Ko¨hler,1996b IsladeUtila,Honduras Dactyloabiporcata Wiegmann,1834 SantaRosadePansos,Guatemala Anolis(Draconura)capito Peters,1863 ‘‘CostaRica’’ Anoliscarpenteri Echelle,Echelle,andFitch,1971 nearTurrialba,CostaRica Anoliscrassulus Cope,1864 Coban,Guatemala Anoliscupreus Hallowell,1861 ‘‘Nicaragua’’ Noropscusuco McCranie,Ko¨hler,andWilson,2000 ElCusuco,Honduras Anolisheteropholidotus Mertens,1952a SantaAna,ElSalvador Anolisjohnmeyeri WilsonandMcCranie,1982 WSWofBuenosAires,Honduras Noropskreutzi McCranie,Ko¨hler,andWilson,2000 nearLaFortuna,Honduras Dactyloalaeviventris Wiegmann,1834 ‘‘Mexico’’ Anolis(Gastrotropis)lemurinus Cope,1861 Veragua5Panama Anolis(Dracontura)capito Cope,1862 Veragua5Panama Anolisloveridgei K.Schmidt,1936 PortilloGrande,Honduras Anolismorazani J.TownsendandWilson,2009 Cataguana,Honduras Noropsmuralla Ko¨hler,McCranie,andWilson,1999 CerrodeEnmedio,Honduras Anolisnelsoni Barbour,1914 SwanIslands,Honduras Noropsocelloscapularis Ko¨hler,McCranie,andWilson,2001 nearQuebradaGrande,Honduras Anolisoxylophus Cope,1875 easternCostaRica Anolispetersii Bocourt,1873,inA.H.A.Dume´ril, VeraPaz,Guatemala Bocourt,andMocquard,1870–1909 Noropspijolense McCranie,Wilson,andWilliams,1993 PicoPijol,Honduras Noropspurpurgularis McCranie,Cruz,andHolm,1993 nearLaFortuna,Honduras Anolisquaggulus Cope,1885 SanJuanRiver,Nicaragua Noropsroatanensis Ko¨hlerandMcCranie,2001 IsladeRoata´n,Honduras Anolisrodriguezii Bocourt,1873 ‘‘PansosGuatemala’’ Noropsrubribarbaris Ko¨hler,McCranie,andWilson,1999 SanLuı´sdelosPlanes,Honduras Anolissagrei Cocteau,inA.M.C.Dume´riland ‘‘Cuba’’ Bibron,1837 Anolissminthus DunnandEmlen,1932 SanJuancitoMountains,Honduras Anolistropidonotus W.Peters,1863 Huanusco,Mexico Anolisuniformis Cope,1885 ‘‘Guatemala’’and‘‘Yucatan’’ Anolisunilobatus Ko¨hlerandVesely,2010 Awasbila,Honduras Noropsutilensis Ko¨hler,1996a IsladeUtila,Honduras Noropswampuensis McCranieandKo¨hler,2001 R´ıosAner-Wampu´,Honduras Anolisustuswellbornae Ahl,1939 ‘‘ElSalvador’’ Noropsyoroensis McCranie,Nicholson,andKo¨hler,2002 nearLaFortuna,Honduras Noropszeus Ko¨hlerandMcCranie,2001 Liberia,Honduras Salvador), and N. carpenteri Echelle, species new to science (Norops amplisqua- Echelle, and Fitch (1971; as Anolis, type mosus McCranie, Wilson, and Williams, locality in Costa Rica). 1992; N. cusuco McCranie, Ko¨hler, and During 1979, McCranie made his first of Wilson, 2000; and N. johnmeyeri Wilson what was to become numerous trips to and McCranie, 1982). Ko¨hler began field- isolated cloud forest and many broadleaf work in Honduras during 1993. The forest localities throughout Honduras. That combined fieldwork of McCranie and first cloud forest trip, to El Cusuco, Corte´s, Ko¨hler, along with examination of Hon- resulted in the discovery of three anole duran anoles in museum collections, re-