Descriptions of two nomen nudum species of Anolis lizard from northwestern South America

2009·7.3 MB·English

by  Steven Poe

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Nl ms e Tilinn ©i Comparative /^oology
US ISSN 0006-9698
Cambridge, Mass. June 2009 Number 516
1
DESCRIPTIONS OF TWO NOMEN NUDUM SPECIES OF ANOLIS LIZARD FROM
NORTHWESTERN SOUTH AMERICA
Steven Poe,1 Julian Velasco,2 Kenneth Miyata,3-4 and Ernest E. Williams3'4
Abstract. We describe two northern South American Pacific lowland species ofAnolis that historically have
beenassociatedwithnamesthatarecurrentlynominanuda. AnolislyranewspeciesissimilartoAnolisvittigerusbut
differsprimarilyinpossessingsmallerdorsalscalesontheheadandbodyandincolorpattern.Anolisanchicayaenew
speciesissimilartoAnolisperaccaeandAnolisfasciatusbutdiffersprimarilyinsizeandcolorpattern. Wescoredthe
new species for skeletal and external morphological phylogenetic characters and performed a parsimony analysis
with251 additionalAnolisspeciesusingmolecularandmorphologicaldata.AnolisanchicayaeissistertoA.peraccae
inabasalgroupofSouthAmerican"alpha"Anolis("dactyloids,""latifrons"group).Anolislyraisplacedwithother
lemurinus-groupAnoliswithin the largeclade ofmainland "beta" Anolis {"Norops").
Key words: Anolisanchicayae; Anolislyra; Colombia; Ecuador
Resumen. Describimos dos especies de Anolis de zonas bajas del Pacifico de Colombia y Ecuador que
tradicionalmente han estado asociadas con nombres en nomen nundum. Anolis lyra sp. nov. es parecido a Anolis
vittigerus pero se diferencia principalmente en que presenta las escamas del dorso de la cabeza y el cuerpo mas
pequenas, y en el patron de coloracion. Anolisanchicayae sp. nov. es parecido a Anolisperaccaey Anolisfasciatus
pero se diferenciaprincipalmenteen el tamanoyel patrondecoloracion. Secodificaroncaracteresfilogeneticos de
morfologiaexternayosteologiayserealizounanalisisdeparsimoniacon251 especiesadicionalesdeAnolisusando
unacombinationdedatosmorfologicosymoleculares.AnolisanchicayaeestaxonhermanodeA.peraccaedentrode
ungrupobasaldeespeciessuramericanasdeAnolis"alfa."Anolislyraestarelacionadoconotrasespeciesdelgrupo
lemurinus dentro de unclado mas grandecompuesto porespeciescontinentales de Anolis "beta."
1Department of Biology and Museum of Southwestern Biology, University ofNew Mexico, Albuquerque, New
Mexico 87131, U.S.A.; e-mail: [email protected]
2Grupo de Ecologia Animal, Departamento de Biologia, Universidad del Valle, A. A. 25360, Cali, Colombia
3Museum ofComparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A.
4Williams and Miyata are deceased and are included as coauthors for the followingcontributions. Poe was able to
diagnose each species and identify MCZ specimens on the basis ofWilliams' unpublished notes, which clearly were
influencedbyMiyata'sworkonthesecondspeciesdescribedherein. Additionally,WilliamsisresponsibleforFigure6.
Subsequent to submission ofthismanuscript, theeditorprovidedPoewith achapter ofMiyata'sPh.D. thesis, never
published,describingthesecondofthetwospeciesdescribedherein.Poeincorporatedinformationfromthismanuscript
intothediagnosisofthisspecies.
®The President and Fellows ofHarvard College 2009.
BREVIORA No. 516
Ernest Williams and collaborators pub- the anterior of the cloaca. Head length was
lished a series of papers from 1983 to 1992 measured from the tip of the snout to the
describing "new orproblematic" Anolis from anterior edge ofthe ear. Femoral length was
Colombia. These papers reviewed and ex- measured from the midline of the venter to
panded the Colombian anoline fauna and the knee, with the limb bent at a 90-degree
established the country as a center of angle. Head width was measured at the
diversity for the Anolis clade. The publica- broadest part of the head, between the
tion of these descriptions and other taxo- posterolateral corners of the orbits. Com-
nomic works on Colombian lizards (e.g., parisons were made with preserved material
Ayala and Castro, 1983) prompted the need of the putative closest relatives of the new
for a summary of available information. species, and with published species descrip-
Stephen Ayala (1986) obliged and produced tions, preserved material, or both of more
a list of the 205 lizard species thought to distant relatives (online Appendix 1). Scale
occur in Colombia. This list included eight terminology and characters used mainly
species ofAnolis (and species in other clades) follow standards established by Williams
listed in quotes ("") and associated with (e.g., Williams et al, 1996) for species
authors to signal that descriptions were not descriptions of anoline lizards. Skeletal
yet published but were forthcoming. Pub- description is given in terms of Poe's (1998,
lisheddescriptionsofAnolisspeciesassociated 2004) and Etheridge's (1959) characters. See
with four of these names soon followed (A. those papers for more detailed descriptions
danieli Williams 1988, A. ruizi Rueda and of skeletal conditions and alternative condi-
Williams 1986, A. lamari Williams 1992, A. tions in Anolis.
medemi Ayala and Williams 1988), but the We scored the new species for the mor-
othernames havelanguishedwithout descrip- phological phylogenetic characters of Poe
tions as nomina nuda for over 20 years. The (2004; Appendix 1) and performed a parsi-
aim of this paper is to rectify this situation mony analysis of these taxa and 251 other
with respect to two of the remaining names species of Anolis and eight outgroups with
well known to Colombian herpetologists. the use of 1,291 parsimony-informative
characters from published sources and some
MATERIALS AND METHODS ofour additional unpublished morphological
data. This dataset includes 91 morphological
We consider species to be evolutionary characters (Poe, 2004), 998 sites of mtDNA
lineages (Simpson, 1961; Wiley, 1978) and (NADH dehydrogenase subunit 2, five
operationalize this concept by identifying transfer RNAs; Nicholson et al., 2005), and
species on the basis ofconsistent differences 182 sites of nuclear DNA (internal tran-
between populations. That is, we hypothesize scribed tracer region; Nicholson, 2002).
that populations that are diagnosable by Codes for morphological data for A. anchi-
major differences in the frequencies of traits cayae and A. lyra are in online Table 1. We
are distinct evolutionary lineages or species used PAUP* version 4.0b10 (Swofford,
(see Wiens and Servedio, 2000). 2002) to perform a heuristic search with
Measurements were made with digital 100 random additions of taxa and tree-
calipers on preserved specimens and are bisection-reconnection branch swapping.
given in millimeters (mm), usually to the We also used PAUPrat (Sikes and Lewis,
nearest 0.1 mm. Snout-vent length (SVL) 2001) to perform several runs of the parsi-
was measured from the tip of the snout to mony ratchet (Nixon, 1999) to find optimal
2009 NEW SOUTH AMERICAN ANOLIS
Colombia, Valle, Rio Blanco, road between
Cali and Anchicaya (where road crosses
m
river), ca. 500 along river just above
bridge on banana plant, 12 May 1980, H.
MCZ
Carvajal, D. Harris, S. Ayala; 159767-
8, Colombia, Valle, Anchicaya Hydroelectric
Central, 3 June 1976, J. Castillo, F. Castro,
Figure 1. Anolis anchicayae new species, male. H. Chin; MCZ 159997-8, 160234-9, Colom-
Photo byJulian Velasco. bia, Valle, San Isidro, Carton, 5 and 9
MCZ
August 1977, H. Chin; 160008-12,
trees (20%-35% of characters reweighted, Colombia, Valle, Granje Bajo Central, 1976-
MCZ
150-200 replicates per run). 77, H. Chin, F. Castro; 160230,
Colombia, Valle, Bajo Calima, 14 August
DESCRIPTION OF NEW SPECIES 1977, H. Chin; MCZ 160231-3, Colombia,
Choco, Bahia Solano, near Jayita, 14-16
Anolis anchicayae, new species MCZ
October 1977, F. Castro; 160633,
Figures 1-5
Colombia, Valle, Anchicaya, 7 December
Holotype. MCZ 160234, adult male, Co- 1978, H. Chin, F. Castro; ICN 6115,
lombia, Valle, San Isidro, 5 August 1977, Colombia, Choco, Bahia Solano, near
Helen Chin. Jayita, J. Renjifo.
MCZ
Paratypes. 112426-7, Colombia, Diagnosis. Anolis anchicayae is similar to
Choco, Rio San Juan, Carlo Docordo A. peraccae and A. fasciatus. These three
between Cucurrupi and Noanama, 24 De- species share keeled dorsal head scales, a
cember 1968, B. Malkin; MCZ 158385-7, double row of middorsal caudal scales,
fSsk
**.
Figure 2. Anolisanchicayae new species, MCZ 145362. Tail is missingin this specimen.
BREVIORA No. 516
Figure 3. Anolisanchicayae, MCZ 145362, dorsal head scales.
absence of dewlap in females, absence of supralabial. Anolis anchicayae is most easily
caudal transverse vertebral processes poste- distinguished from A. peraccae by larger size
riorly, and an elongate anterior nasal scale (to 54 mm SVL in A. peraccae; to 63 mm in
that reaches the rostral but does not contact A. anchicayae; Fig. 6) and color pattern {A.
the sulcus between the rostral and first anchicayae is predominantly yellow-green on
2009 NEW SOUTH AMERICAN ANOLIS
Figure 4. Anolisanchicayae, MCZ 145362, lateral head scales.
its flanks with an elongate ocellus from eye
to shoulder and a yellow-green dewlap in
males; A. peraccae is predominantly brown
on its flanks with no shoulder ocellus and a
cream-colored male dewlap). Anolis anchi-
cayae is most easily distinguished from A.
fasciatus by size (to 72 mm SVL in A.
mm
fasciatus; to 63 in A. anchicayae) and
male dewlap color (white with gray scales in
A. fasciatus, yellow-green in A. anchicayae).
Anolis anchicayae can be distinguished
from other South AmericanAnolis as follows
(groupings are informal according to Savage
and Guyer [1989] and others and used only
for organizational purposes; no relationships
are implied): from "beta" anoles (= "Nor-
ops"; with whichitis unlikely to beconfused)
by the absence of transverse processes on
posterior caudal vertebrae (present in beta
Anolis); from phenacosaur and tigrinus-
group Anolis {A. heterodermus, A. inderenae,
A. nicefori, A. bellipeniculus, A. carlostoddi,
A. euskalerriari, A. neblininus, A. orcesi, A.
Figure 5. Anolis anchicayae, MCZ 145362, ventral tetarii, A. vanzolinii, A. ruizi, A. solitarius, A.
head scales. menta, A. nasofrontal, A. pseudotigrinus, A.
BREVIORA No. 516
MALES FEMALES
SVL 42mm44 46 48 50 52 54 56 58 60 SVL 42mm 44 46 48 50 52 54 56 58
30 Anolis peraccae
N = 30 Norino; Colombia
20
10
50
Anolis per
40 N»I8 Plchincha; N=28
30
20
10
% K%4l%!*'&3kS£^•i.!':.:.:.:.**:'*':'*.* . .
SVL 42mm 44 46 48 50 52 54 56 58 60 SVL42mm 44 46 48 50 52 54 56 58
Figure 6. Distribution ofbody sizeswithinpopulations for samplesofA.peraccaeandA. anchicayae, showing
largersizeofA. anchicayae. Thespecimensused toconstruct thistableareunknown becausetheywerenot listed in
Williams' notes.
tigrinus, A. menta, A. lamari, A. vaupesianus, scales separating the supraorbital semicircles
A.jacare,A. umbrivagus, A.paravertebralis,A. (supraorbital semicircles in contact in these
williamsmitteremeierorum) by possessing species); from other punctatus- and aequa-
strongly keeled head scales superior to the torialis-group Anolis (A. chocorum, A. cali-
orbits (mostly smooth or pustulose in mae, A. antioquiae, A. fitchi, A. aequator-
phenacosaur and tigrinus-group Anolis); ialis, A. megalopithecus) in the lack of a
from latifrons- and some aequatorialis-group dewlap in females (female dewlap present in
Anolis (A. latifrons, A. frenatus, A. fraseri, these species); from A. punctatus and the
A. danieli, A. apollinaris, A. casildae, A. proboscis Anolis (A. proboscis, A. laevis, A.
squamulatus, A. princeps, A. purpurescens, phyllorhinus) by the absence of swelling on
A. agassizi, A. eulaemus, A. maculigula) by the snout of the male (swelling present in
much smaller size (> 100 mm SVL in these males of these species); from mirus-gvoup
species); from some punctatus-group Anolis Anolis {A. mirus, A. parilis) by the presence
(A. cuscoensis, A. soinii, A. huilae, A. of a broad, raised toepad (toepad narrow
transversalis, A. boettgeri, A. deltae, A. and continuous with first phalanx in mirus
dissimilis, A. sanlamartae) by displaying group); from A. chloris in body color and
2009 NEW SOUTH AMERICAN ANOLIS
pattern (solid bright green in A. Moris; especially posteriorly; interparietal length 1.3
banded pattern of browns and greens in A. (1.2-1.6); three (3-4) scales separate interpa-
anchicayae); from A. festae and A. nigroli- rietal and supraorbital semicircles; preoccip-
neatus (the name is a probable synonym of ital absent; six (5-6) supralabials to center of
A. festae; Williams, 1982) in male dewlap eye; nine (8-10) postrostrals; six (6-9) post-
pattern (dark elongate blotch in these mentals; some slightlyenlarged scales present
species; absent in A. anchicayae); from A. in supraocular disc, decreasing gradually in
caquetae in absence of contact between size, bordered medially by a partial row of
interparietal and supraorbital semicircles; small scales; mental partially divided poste-
from A. propinqus in number of subdigital riorly, extending posterolaterally beyond
lamellae (27 in A. propinqus, 16-20 in A. rostral, with posterior border in concave
anchicayae); from A. gemmosus in the arc; two (2-A) enlarged sublabials in contact
condition of the nasal scale (elongate with infralabials (or posteriorly separated by
anterior nasal contacts sulcus between small scales); dewlap reaches well posterior
rostral and first supralabial in A. gemmosus; to axillae in males, absent in females; scales
is separated from sulcus by small scale on dewlap in rows of approximately three
inferior to naris in A. anchicayae). scales with some scattered scales; no axillary
ExternalDescription ofHolotype (paratype pocket; pair ofenlarged postcloacal scales in
variation in parentheses; description is based contact (or separated by small scales; absent
MCZ
on adults in good condition: 160234, in females); nuchal and dorsal crests absent;
158386-7, 159767, 159997-8, 160009-10, dorsal scales keeled; approximately 10 (5-12)
160230, 160232-3, 160238). Snout-vent enlarged middorsal rows, six (6-8) longitu-
length 52.0 (50.0-53.6); head length 12.0 dinal rows in 5% of SVL; ventral scales in
(11.8-13.2; 0.23-0.25 SVL), width 7.1 (7.0- mostly transverse but some diagonal rows,
7.8; 0.14-0.15 SVL); ear height 1.3 (1.1-1.6; smooth, eight (7-9) scales in 5% of SVL;
0.02-0.03 SVL); femoral length 12.9 (13.6- anterior thigh scales large, keeled, overlap-
16.0; 0.25-0.30 SVL); tail length 115 (102- ping, becoming smaller and nonoverlapping
116; 1.96-2.26 SVL); fourth toe length 10.5 posteriorly; supradigitals multicarinate; toe-
(10.7-11.9; 0.21-0.22 SVL), width 1.0 (1.0- pads expanded; 19 (16-20) expanded lameal-
1.3; 0.02 SVL). lae under third and fourth phalanges of
Dorsal head scales uni- to multicarinate fourth toe (second and third phalanges of
above orbits, mostly smooth in frontal area, Williams [e.g., Williams et al, 1995]); tail
unicarinate on snout; frontal depression with a double row ofmiddorsal scales.
present; rostral slightly overlaps mental Skeletal Description (Based on Dry Skele-
MCZ
anteriorly; nine (8-12) scales across snout tons 112426-7, 159767, 160012,
between second canthals; supraorbital semi- ICN6115). Parietal roof flat, with trapezoi-
circles separated by two (1-2) scales; sub- dal crests, with a very narrow posterior
oculars incontactwith supralabials; one (1-2) border, with no casquing, lacking crenula-
elongate supraciliary scale followed by small tion on edges, not extending posteriorly over
undifferentiated scales; seven (5-7) loreal supraoccipital, with anterolateral corners
rows; elongate anterior nasal scale contacts flush with posterolateral edges of frontal;
rostral, is separated from sulcus between pineal foramen at parietal-frontal suture (n
rostral and first supralabial by one small = 4) or in parietal (n = 1); dorsal skull bones
scale inferior to naris; supraorbital semicir- smooth; postfrontal present; prefrontal sep-
cles evident but not strongly differentiated, arated from nasal by anterior extension of
BREVIORA No. 516
frontal (n = 2) or in contact with nasal (n = ually lost posteriorly (Etheridge's [1959]
2); frontal sutures anteriorly with nasals; "alpha" condition).
parallel crests on nasals absent; external Color in Life (Adaptedfrom Field Notes
nares bordered posteriorly by nasals; dorsal and Color Photos by Velasco). Males brown
aspect ofjugal terminates on lateral surface middorsally interrupted by approximately 10
of postorbital; jugal does not contact squa- dark blotches that extend laterally; flanks
mosal; posterodorsal ramus of squamosal greenish-yellow with narrow diagonal rows
shorter than posteroventral ramus, separated of partially connected small brown spots;
from parietal by supratemporal; posterior elongate brown mark extending back from
aspect ofjugal mostly straight; epipterygoid eye, bordered by black; upper head part
contacts parietal dorsally; pterygoid and yellow with brown, lateral head brown;
palatine teeth absent; lateral edge of vomer ocular border yellow; yellow with brown
is smooth, without posteriorly directed lat- blotches on lips; dewlap yellow green with
eral processes; maxilla extends posteriorly brown spots; iris blue; limbs and tail banded.
beyond ectopterygoid on ventral surface of Females with broad middorsal stripe bor-
skull (n = 3) or reaches ectopterygoid (n = dered laterally by black; flanks yellow green
2); crest between basipterygoid processes of to yellow with longitudinal very narrow dark
basisphenoid absent; lateral shelfofquadrate brown stripes similar to middorsal stripe.
absent; no black pigment on surface ofskull; Distribution. Anolis anchicayae is found
nasals do not overlap premaxilla dorsally; throughout the Pacific coastal lowlands of
posterior edge of parietal is approximately Colombia and Ecuador in primary and
even with vertical level of parietal-frontal secondary humid forest (Fig. 7). Many spec-
suture; posteriormost mandibular tooth is imens were collected by day on trunks
partially anterior (n = 3 mandibles) or between 2 and 5 m.
partially posterior (n = 7 mandibles) to Etymology. The species is named for its
anterior mylohyoid foramen; large splenial type locality, the Anchicaya River valley.
present; ventral aspect of anteromedial pro-
cess ofcoronoid extends posteriorly (n = 4) Anolis lyra, new species
or slopes anteriorly (n = 1); external opening Figures 8, 9, 10A
of surangular foramen is entirely within
surangular (n = 1) or bordered by dentary Holotype. MCZ 80955, adult female,
and surangular (n = 3); posterior suture of Ecuador, Pichincha, Finca Victoria, 37 km
dentary is pronged; anteriormost aspect of SE of Santo Domingo, 16 July 1964, F.
posterior border of dentary is anterior to (n Vuilleumier.
= 3 mandibles) or within (n = 7 mandibles) Paratypes. MCZ 77458-60, Colombia,
mandibular fossa; labial process ofcoronoid Valle, Lower Rio Calima, tributary of Rio
MCZ
present; coronoid does not extend postero- San Juan, 1960, I. Cabrera; 80954,
laterally beyond surangular foramen; jaw Ecuador, Pichincha, Finca Victoria, 37 km
sculpturing absent; angular absent; angular SE of Santo Domingo, 16 July 1964, F.
MCZ
process of articular present; teeth unicuspid Vuilleumier; 124407, Ecuador, Pi-
anteriorly, tricuspid posteriorly; nine (n = 3) chincha, Santo Domingo de los Colorados,
or 11 (n = 1) premaxillary teeth. 2 August 1968, J. Lynch; MCZ 145263,
Four postxiphisternal ribs attached dor- Ecuador, Pichincha, Hotel Tinalandia,
sally, none free (4:0 rib formula); transverse 15 km N Santo Domingo de los Colorados,
processes on anterior caudal vertebrae grad- 6 November 1964, A. Kiester, E. Williams;
2009 NEW SOUTH AMERICAN ANOLIS
200 Kilometers
Figure 7. Mapofcoastal regionsofColombiaandEcuadorshowingcollectingsitesofAnolisanchicayae(filled
triangles) and Anolis lyra (open squares) according to type series and personal observations ofauthors. 1. Nuqui,
Choco Department, Colombia; 2. Pueblo Rico, Risaralda Department, Colombia; 3. Cafio Docordo between
Cucurrupi and Noanama, San Juan River, Choco, Colombia; 4. Bajo Calima, Buenaventura, Valle del Cauca
Department, Colombia; 5. Cisneros, Buenaventura, Valle del Cauca Department, Colombia; 6. Bajo Anchicaya,
Buenaventura, Valle del Cauca Department, Colombia; 7. Cajambre river, Buenaventura, Valle del Cauca
Department, Colombia; 8. Malaga Bay, Buenaventura, Valle del Cauca Department, Colombia; 9. Tangareal,
Nariiio Department, Colombia; 10. Centro Cientifico Rio Palenque, Los Rios Province, Ecuador; 11. Finca La
Victoria, southeast from Santo Domingo de los Colorados, Pichincha Province, Ecuador; 12. Puerto Quito,
PichinchaProvice,Ecuador; 13. HotelTinalandia,approx. 15 kmNofSantoDomingodelosColorados,Pichincha
Province, Ecuador.
MCZ
145867, Ecuador, Los Rios, Centro lyra differs from A. vittigerus in possessing
Cientifico Rio Palenque, 23 June 1974, K. smaller middorsal scales (mean number of
Miyata; MCZ 146994-5, Ecuador, Pin- longitudinal scales in 5% SVL for A. lyra =
chincha, 1 km N Buena Fe, 31 July 1975, 8.5, A. vittigerus = 5.8; P = 0.006, Mann-
R. Webster; MCZ 152433-4, Ecuador, Pi- Whitney U test), more scales between supra-
chincha, Hotel Tinalandia, 15 km N Santo orbital semicircles (mean = 2.3 in A. lyra, 0.6
Domingo de los Colorados, 27 April 1977, in A. vittigerus; P = 0.001, Mann-Whitney U
MCZ
K. Miyata; 164416, 165209-10, test), and more scales between interparietal
171160, Ecuador, Pinchincha, Puerto Quito, and supraorbital semicircles (x = 4.1 in A.
April and September 1983, December 1984, lyra, 2.4 in A. vittigerus; P = 0.001, Mann-
MCZ
G. Onore; 164420-1, Ecuador, Pinch- Whitney f/test; Fig. 10). Also, A. lyra tends
incha, Santo Domingo de los Colorados, to display much weaker lateral striping (i.e.,
April 1983, G. Onore. broken lines only extending to the axillae
Diagnosis. Anolislyradiffers from all other versus broad stripes to groin in A. vittigerus)
SouthAmericanAnolisexceptA. vittigerusin and a more prominent lyre-shaped posterior
possessing a small red dewlap with a dark head marking (Fig. 8; absent to weak in A.
central spot in males and a lyre-shaped vittigerus). In western Colombia, the only
symbol in the occipital region (Fig. 8). Anolis area in which both species can be expected to
10 BREVIORA No. 516
Figure 8. Anolis lyra new species, male, from 2 km N ofBoca Ana, Valle, Colombia. Photo by Steven Poe.
figure 9. Anolis lyra new species, female, from 2 km N ofBoca Ana, Valle, Colombia. Photo by Steven Poe.