From Eastern Cuba

A New Species of Anolis (Sauria: Iguanidae) from Eastern Cuba

l 2 A LBERTO R. ESTRADA AND S. BLAIR H EDGES

1 lnstituto de Investigaciones Forestalls, Apartado 5152, La Habana 5, C.P. 10500, Cuba ‘Department of Biology, 208 Mueller Laboratory, Pennsylvania State University, University Park, Pennsylvania 16802, USA

ABSTRACT. – Anolis alayoni is described from upland localities in eastern Cuba. It is a cryptic species, and the fourth member of the twig ecomorph of Cuban Anolis. Morphological data suggest that its closest relative is the partly sympatric species A. angusticeps.

The largest amniote genus, Anolis (>300 National Museum of Natural History, sp.), is represented in Cuba by more than Smithsonian Institution), and CARE (A. R. 50 species (Estrada and Garrido, 1991; Gar- Estrada Collection). rido and Hedges, 1992; Hass et al., 1993). One of the most unusual groups of this Anolis alayoni sp. nov. widespread genus on Cuba is the angusti- (Fig. 1A) ceps group, of which there are three spe- Holotype. —MNHNCU 2746, an adult cies: Anolis angusticeps, A. guazuma, and A. male from La Fangosa, Yateras, Guanta- paternus (Williams, 1976; Burnell and namo Prov., Cuba, collected by Alberto R. Hedges, 1990). All three are members of Estrada on 22 June 1990. the twig ecomorph (Williams, 1983), char- Paratypes. —(29) CARE 60800, male, same acterized by a cryptic morphology (resem- collecting data as holotype; MCZ R178768 bling a twig) and behavior. (male) and CARE 60721 (female), between Anolis angusticeps has been the focus of Cayo Fortuna and Riito, Yateras, Guanta attention of several previous studies. Har- namo Prov., 11 March 1989, collected by dy (1966) studied variation in the Cuban A. R. Estrada; MCZ R178769, CARE 60801, and Bahama Islands populations, and de- female, Arroyon, San Antonio del Sur, scribed A. angusticeps paternus from Isla de Guantanamo Prov., 22 June 1990, collected la Juventud. Schwartz and Thomas (1968) by A. R. Estrada; CARE 60872, female, Cayo examined variation in A. angusticeps in more Fortuna, Yateras, Guantanamo Prov., 16 detail, including in their analysis speci- March 1987, collected by Antonio Perez- mens from eastern Cuba. Later, Garrido Asso; CARE 60759, 60819, females, Piedra (1975) analyzed variation in western pop- la Vela, Yateras, Guantanamo Prov., 22 ulations and treated A. paternus as a distinct March 1989, collected by A. R. Estrada and species. Finally, Garrido (1983) described A. Perez-Asso; CARE 60299, female, Far- A. guazuma from Pico Turquino in Santiago allones de la Italiana, Levisa, Mayari, Hol- de Cuba province. In this paper, we de- guin Prov., 18 September 1987, collected scribe a fourth Cuban twig anole from lo- by A. R. Estrada and Orlando H. Garrido; calities in two eastern provinces of Cuba: CARE 60327 female, Cayo Guan, Mea, Hol- Holguin and Guantanamo. guin Prov., 3 October 1987, collected by A. In the account below, the following ab- R. Estrada; CARE 60797, 60799 females, Es- breviations are used: MNHNCU (Museo pana Chiquita, Sierra de Canasta, El Sal- National de Historia Natural de Cuba), vador, Guantanamo Prov., 20 June 1990, CZACC (Colecciones Zoologicas, Instituto collected by A. R. Estrada; CARE 60186, de Ecologia y Sistematica, Academia de male, Arroyo Culebra de Hacha, Mea, Hol- Ciencias de Cuba), USNM (United States guin Prov., 21 September 1986, collected 66 A. R. ESTRADA AND S. BLAIR HEDGES

FIG. 1. (A) Anolis alayoni (El Molino, Guantinamo Prov.), and (B) A. angusticeps (9 km W Trinidad, Sancti Spiritus Prov.). by A. R. Estrada; MNHNCU 31, male, 3 km Prov.. 20 September 1980. collected by Noel E La Melba, Mea, Holguin Prov., 23 Sep- Gonz~lez; CZACC 7302 (old number IZ tember 1987, collected by Carlos Pefia and 4220), male, Arroyo Blanco, Baracoa, Guan- Orlando H. Garrido; MNHNCU 2762, fe- tiinamo Prov., 27 May 1974, collected by male, Sumidero del Rio Cuzco, El Salvador, O. H. Garrido; CZAZZ 9778 (old number Guant&amo Prov., 16 June 1990, collected IZ5272), male, Sabanilla, Baracoa, Guan- by Alfonso Silva Lee; USNM 335020, El tiinamo Prov., May 1978, collected by An- Molino (ca. 7 km W Palenque), Yateras, filoquio Suirez; CZACC 7325, female, La Guantinamo Prov., 21 June 1990, collected Florida, Baracoa, Guanttinamo Prov., 14 by S. Blair Hedges; MNHNCU 51,70, males, September 1965, collected by O. H. Garri- La Poa, Sabanilla, Baracoa, Guant5namo do; CZACC 7326, 7332, female and male, Prov., between November 1986 and March Base de Monte Iberia, Baracoa, Guant6na- 1987, collected by O. H. Garrido; MNHNCU mo Prov., 9-10 March 1972, collected by 1607, male, Yumuri, Maisi, Guantzinamo Jorge de la Cruz and Luis F. de Armas; Prov., 27 July 1989, collected by Riberto CZACC 7377, male, Base de Monte Iberia, Arencibia; CZAZZ 7400, (old number IZ Baracoa, Guanttinamo Prov., 9 March 1972, 6267), male, Yumuri, Mais!, Guant&amo collected by O. H. Garrido; CZACC 7264 NEW CUBAN ANOLIS 67

(old number IZ 5941), male, Gran Tierra, less keeled, smallest anteriorly; nostril cir- Maisi, Guantanamo Prov., 10 May 1972, cular; nasal scale separated from rostral by collector unknown; CZACC 7266 (old 2-3 (modally two; two in holotype) irreg- number IZ 5929), male, Nibujon, Baracoa, ularly shaped scales; supraorbital semicir- Guantanamo Prov., Lorenzo Zayas; CZACC cles large, convex, rugose laterally, sepa- 7329-30, female and male, 9 km S from Ase- rated by 1-2 (modally one; one in holo- rrio. type) rows of scales of small size; canthal Diagnosis. —A small species (2 SVL 42.2 ridge with five scales well-defined, second mm males; 36.6 mm females) of Anolis with canthal longest, diminishing in size ante- short limbs, long snout, short semiprehen- riorly; supraorbital ridge well-defined in sile tail, and enlarged scales on dorsal sur- both sexes; frontal ridge well-defined, con- face of head. Anolis alayoni is a member of fluent anteriorly; 10-29 loreal scales (mod- the angusticeps species group of Anolis, ally 24; 22 in holotype); temporal scales which includes A. angusticeps, A. guazuma, small; interparietal about 1.4 times as long and A. paternus. as wide, separated from supraorbital sem- From A. guazuma, it can be distinguished icircles by one scale or in contact (one in by a tail longer than SVL (shorter in gua- holotype); 8-13 scales around the inter- zuma), 1-2 scales between supraorbital parietal (modally 11; 10 in holotype); ear semicircles (modally 0 in guazuma), keeled opening small, elliptical or subcircular, be- head scales (smooth in guazuma), well-de- hind mouth commisure; 5-9 (modally 7; 7 veloped yellow dewlap (small and whitish in holotype) scales between second can- in guazuma), and other scale and pattern thals; 3-5 (modally four; five in holotype) features. From A. paternus, it can be distin- scales between nasal scales; suboculars di- guished by smooth ventral scales (keeled rectly in contact with supralabials, ante- in paternus) and larger body size. riorly grading into loreals; 6-10 (modally From its closest relative, A. angusticeps, it eight; eight in holotype) supralabial scales; can be distinguished by large size of the mental large, divided, in contact with 4-6 males (X SVL = 42.2 mm in alayoni; 39.9 small, granular postmarital scales (modally mm in angusticeps) and the color of the four; five in holotype). dewlap: dark yellow in alayoni, pale orange Trunk with dorsal scales small, granular; or peach in angusticeps. Scale characters two middorsal rows larger, 25-35 scales in which can distinguish most A. alayoni from 0.5 cm on middorsal region (modally 28; most A. angusticeps are: modally 11 (8-12) 26 in holotype); ventral scales larger than scales around the interparietal in males, 10 dorsals, smooth, rounded, arranged in (9-13) in females of alayoni (modally 13 transverse rows, 18–28 scales in 0.5 cm on [11-16] in males, bimodally 13 and 14 [12- midventral region (bimodally 20 and 22; 15] in females of angusticeps); modally two 22 in holotype); dewlap large (Fig. 2), pres- [2-3] scales between naris and rostrum in ent in both sexes, but vestigial in females; alayoni (modally three [2-3] in angusticeps), scales large and arranged in rows, larger modally four (3-5) scales between nares in than throat scales; limbs short, tibial length alayoni (modally five [3–6] in angusticeps), shorter than femoral length, quotient of modally 15 (13–16) lamellae under phalan- humeral length/tibial length 0.74- 1.0 (0.88 ges II and III of fourth toe in alayoni (mod- in holotype); 13–16 (15 in holotype) la- ally 13 [12-20] in angusticeps), modally four mellae under phalanges II and III of fourth (2-6) postmarital scales in alayoni (modally toe; scales of supracarpal smooth; scales of six [4-7] in males, four [4-6] in females of hind limbs smooth, keeled on supratarsals; angusticeps), bimodally 20 and 22 (18-28) tail laterally compressed, broader at the middorsal scales in alayoni (bimodally 35 base. and 37 [26-37] in angusticeps). In combi- Coloration in life pale grayish-brown (in nation, these color, measurement and light phase), with black and brown lin- modal scale differences clearly distinguish chenate markings; 4-5 brown or black all specimens of alayoni and angusticeps. crossbands or blotches with “X” form on Description .—Head narrow and elongate; middorsal line; 5–7 more distinct brown head scales enlarged, hexagonal, more or bands on tail, alternating with gray bands; 68 A. R. ESTRADA AND S. BLAIR HEDGES

TABLE 1. Measurements of Anolis alayoni and A. angusticeps. Snout-vent length (SVL); head length (HL); head width (HW); jaw length (JL); interorbital distance (IO); naris-snout distance (NS); snout-orbit distance (SO); internarial distance (IN); interparietal scale length (IPL); interparietal scale width (IPE); ear opening vertical diameter (EVD); ear opening horizontal diameter (EHD); humeral length (HUL); tibial length (TL); femoral length (FL); tail length (TAL); axilla-groin distance (AG); mean (1); standard deviation (SD); minimum- maximum (m-M); sample size (n).

A. alayoni A. angusticeps Males Females Males Females 2 .7 z ? SD SD SD SD Character n m-M n m-M n m-M n m-M SVL 42.2 36.6 39.9 38.7 2.89 1.74 2.7 2.22 16 35.8 -46.8 13 33.3 -38.9 10 34.3 -44.8 6 36.6 -42.2 HL 12.6 10.2 11.6 10.1 0.79 0.47 0.99 0.44 16 10.9-13.8 13 8.9-10.7 10 10-13.1 6 9.5-10.8 HW 7.5 6.0 6.5 5.8 0.66 0.59 0.52 0.52 16 6.2-8.8 13 5.2-7.4 10 5.6-7.2 6 5.4-6.7 JL 10.9 8.8 9.9 8.8 0.77 0.41 0.93 0.56 16 9.2-12.2 13 7.8-9.3 10 8.2-11.5 6 8.2-9.4 IO 4.3 3.6 3.9 3.5 0.38 0.28 0.42 0.28 16 3.7-5.0 13 3.1-4.0 10 3.2-4.5 6 3.1-3.8 NS 1.3 1.1 1.2 1.1 0.10 0.09 0.19 0.18 16 1.1-1.5 13 1.0-1.3 10 1.0-1.6 6 0.7-1.2 SO 5.7 4.5 5.3 4.8 0.40 0.33 0.72 0.35 16 5-6.5 13 3.8-5.1 10 4.5-6.8 6 4.4-5.2 IN 1.6 1.3 1.4 1.5 0.16 0.10 0.17 0.17 15 1.2-1.8 13 1.2- 1.5 9 1.1-1.6 6 1.3-1.6 IPL 1.8 1.49 1.7 1.4 0.26 0.21 0.33 0.34 16 1.1-2.2 12 1.2- 1.9 10 1.4-2.0 6 1.0-1.8 IPW 1.2 1.0 1.4 1.1 0.26 0.15 0.25 0.19 16 0.9-2 12 0.9-1.4 10 1.0-1.8 6 0.8-1.3 EVD 0.8 0.6 0.6 0.5 0.1 0.1 0.09 0.08 16 0.5-1.0 12 0.4-0.9 10 0.6-0.9 6 0.4-0.6 EHD 0.7 0.6 0.5 0.5 0.1 0.1 0.1 0.1 16 0.5-1.0 12 0.4-0.9 10 0.3-0.7 6 0.4-0.7 HUL 6.6 5.7 6.1 5.4 0.5 0.3 0.7 0.2 16 5.6-7.6 13 5.1-6.2 10 4.3-7.2 5 5.1-5.6 — — NEW CUBAN ANOLIS 69

T ABLE 1. Continued.

A. alayoni A. angusticeps Males Females Males Females .-i f ? .7 SD SD SD SD Character n m-M n m-M n m-M n m-M TL 7.4 6.4 7.1 6.3 0.5 0.3 0.5 0.19 16 6.2-8.6 13 5.7-7.6 10 6.0-8.1 6 6-6.6 F1 8.9 7.5 8.3 7.5 0.6 0.6 0.6 0.3 16 7.6-9.8 13 6.4-8.7 10 6.7-8.8 6 7.2-8 TAL 46.4 44.5 45.3 54.6 5.15 5.83 5.68 4.66 6 41.9-53.3 8 36.0 -55.0 7 38.6-54.8 3 50.5 -59.7 AG 16.9 15.3 15.3 15.4 1.5 1.3 1.9 1.3 16 13.5 -19.4 13 13.5 -16.9 10 11.5-18.6 6 12.9 -16.6

a simple black or brown dot postventral to flanks above and below lateral stripe; pale ear opening; about 4 narrow dark brown yellow postmarital and throat, with 5-6 lines radiating from eye; orbital region brown bands on each side; yellow from reddish-brown; dorsal surface of head light chest to belly, with brown spots; base of brown; yellowish-beige lateral stripe from tail reddish; dewlap yellow ochre in males, ear to hind limb insertion; grayish-brown pale yellow anteriorly; scales in margin

TABLE 2. Meristic characters of A. alayoni and A. angusticeps. Scales around interparietal (SAI); scales between naris and rostral (SNR): middorsal scales in 0.5 cm (SMD); midventral scales in 0.5 cm (SMV); rows of scales between supraorbitals (RSS); scales between interparietal and supraorbital (SSI); supralabial scales (SSL); infralabial scales (SIL); loreal scales (SL); internarial scales (SIN); lamellae under II-III fourth toe phalanges (L4T); postmarital scales (SPM); scales between second canthal (SSC). Mean (z); mode (M); sample size (n).

~ M n. SAI SNR SMD SMV RSS SSI SSL SIL SL SIN L4T SPM SSC A. alayoni Males 10 2 28 22 1 8 9 22 4 15 57 11 2 28 20-22 1 1/1 8 8 24 4-5 15 47 15 14 16 16 16 16 16 16 16 16 16 16 16 Females 11 2 30 19 1 8 8 19 4 15 58 10 2 28 20 1 0/0 8 8 — 4 15 47 13 13 13 13 13 13 13 13 13 13 10 13 13 A. angusticeps Males 13 3 32 26 1 8 9 26 5 15 57 13 3—— 1 1/1 8 8 24 5 13-14 6 7 10 10 10 10 10 10 10 10 10 10 10 10 10 Females 14 3 31 21 1 9 10 20 5 15 57 13-14 3 — — 1 1/1 — 9 — 5 13 46 6 66 6 666666 6 66 70

FIG. 2. Extended dewlaps of (A) Anolis alayoni (El Molino, Guantin.nm Prov.), and (B) A. angusticeps (9 km W Trinidad, Sacti Spiritus Prov.). yellow, light brown inside. Animals with with A. angusticeps are given in Tables 1 dark phase have the same patterns, but with and 2. some differences general coloration dark Distribution (Fig. 3).—Known only from reddish-brown; dark brown orbits; dorsal upland areas of Holgtdn and Gwmt&mmo surface of head and lores dark brown; mid- provinces in eastern Cuba. Alitudinal dorsal brown stripe with a light fine line range 200-900 m. in the middle; brown belly with dark spots Etymology. –The new species is dedicat- (CARE 60800). Some animals have black or ed to our friend, and distinguished Cuban dark brown blotches on sides below the arachnologist, Giraldo Alay6n. mid-dorsal stripe and above the lateral light Natural History. –Anolis alayoni occurs in stripe (MCZ R178769, CARE 60798). various kinds of forest. In the Sierra de la Measurements of Holotype.— SVL 46.8, Canasta (Espafia Chiquita), it was observed 13.8, 7.1, tibial length 8.0, femoral length on tree trunks and twigs at two or three 9.4, and tail length 53.3. Complete mea- meters above the ground, in very dry surements for A. alayoni and comparison woods. Animals from Cayo Fortuna, Piedra NEW CUBAN ANOLIS 71

FIG. 3. Distribution of two species of Anolis in eastern Cuba. Localities for A.alayoni (circles) are (from west to east): Farallones de la Italiana (Sierra del Cristal), Sumidero del Rio Cuzco (western Meseta del Guaso), Espana Chiquita (Sierra de Canasta), Culebra de Hacha (Cuchillas de Mea), Piedra la Vela (Cuchillas de Tea), Cayo Fortuna (Cuchillas de Toa), Riito (Cuchillas de Toa), Cayo Cuan (Alturas de Baracoa), 3 km E de la Melba (Cuchillas de Toa), Arroyon (Sierra del Purial), Base de Monte Iberia (Alturas de Baracoa), Aserrio Nuevo Mundo (Alturas de Baracoa), Nibujon (Alturas de Baracoa), Nibujon and La Florida (Alturas de Baracoa), Sabanilla (Cuchillas de Baracoa), Boca del Yumuri y Gran Tierra (Meseta de Maisi). The localities of A. angusticeps (triangles) are: San Ramon, Bajada del Pesquero de la Alegria (Meseta de Cabo Cruz), Finca La Bahia, Pico Turquino, La Mula, 4 km W del Uvero, Santiago de Cuba, Playa Siboney, Playa Jaragua, Sardinero, Cuchillas de Guajimero, 9 km SE Mea, Ovando Arriba. - la Vela a Culebra de Hacha, and Cayo Guan, with a yellow dewlap. The dewlap color were collected in very dry vegetation as- of alayoni is always yellow, and females sociated with Pinus cubensis, termed “Char- have a yellow throat. rascal.” These animals were perched on Although A. alayoni has not been taken twigs and tree trunks at about the same syntopically with A. angusticeps, the distri- height. The specimens from Arroyon were butions of these two species overlap broad- collected on twigs very close to the river ly (Fig. 3). Three localities for A. angusticeps in secondary rainforest. The same situation lie within the range of A. alayoni and are was observed for specimens from La Mel- only a few kilometers from localities of A. ba, Rio Cuzco, and Farallones de la Italiana. alayoni (Fig. 3), therefore these two species Animals collected at El Molino were sleep- should be considered sympatric. ing on twigs at night several meters above Acknowledgments. —For collecting per- ground next to a small stream. mits and general support, we thank Gil- Remarks. -Ruibal (1964:488) reported a berto Silva and the staff of the Museo Na- peach dewlap color for Cuban examples of cional de Historia Natural de Cuba. Luis A. angusticeps, and in our experience this V. Moreno and Luis F. de Armas (I.E.S. is the typical color throughout the range Habana, Cuba) loaned us comparative ma- of that species. However, there is some terial from CZACC, Academia de Ciencias variation, and Schwartz and Thomas (1968: de Cuba. The curatorial staffs of the Amer- 55-56) reported a male from Pinar del Rio ican Museum of Natural History, Museum 72 A. R. ESTRADA AND S. BLAIR HEDGES of Comparative Zoology, and the National a species list. Breviora, Museum of Comparative Museum of Natural History (Smithsonian zoology 440:1-21. ——. 1983. Ecomorphs, faunas, island size, and Institution) kindly allowed the senior au- diverse end points in islands radiations of Anolis. thor to examine specimens in their collec- In R. B. Huey, E. R. Pianka, and T. W. Schoener tions. This research was supported in part (eds.), Lizard ecology, pp. 326-370, Harvard Uni- by grants from the U.S. National Science versity Press, Cambridge. Foundation (BSR 8906325 and DEB 9123556) to S.B.H. APPENDIX

LITERATURE CITED Specimens Examined Burnell, K. L., and S. B. Hedges. 1990. Relationships Anolis angusticeps (25).—CUBA: Bosque of West Indian Anolis (Sauria: Iguanidae): an ap- de La Habana, Ciudad de La Habana, CARE proach using slow-evolving protein loci. Carib. J. Sci. 26:7-30. 60814- 17; Cienaga de Zapata, Matanzas, Estrada, A. R., and O. H. Garrido. 1991. Dos nuevas CZACC 7420-21 (IZ 273-74); San Miguel especies de Anolis (Lacertilia: Iguanidae) de la re- de los Banos, Jovellanos, Matanzas, gion oriental de Cuba, Carib. J. Sci. 27:146-161. MNHNCU 1259, 1261-62; Cayo Cantiles, Garrido, O. H. 1975. Variacion de Anolis angusticeps Archipielago de los Canarreos, CZACC Hallowell (Lacertilia: Iguanidae) en el Occidente de Cuba. Poeyana 144:1-18. 7335 (IZ864), Cayo Juan Garcia, Cayeria de ——. 1983. Nueva especie de Anolis (Lacertilia: San Felipe, CZACC 7336 (IZ865), Rio Cal- Iguanidae) de la Sierra del Turquino, Cuba. Carib. abazas, Placetas, Villa Clara, MNHNCU J. Sci. 19(3-4):71-76. 3238; Bajada al Pesquero de la Alegria, Ni- ——, and S. B. Hedges. 1992. Three new grass anoles (Sauria: Iguanidae) from Cuba. Carib. J. Sci. quero, Granma, CARE 60536; La Mula, 4 28:21-29. km W de Uvero, Guama, Santiago de Cuba, Hardy, J. D. 1966. Geographic variation in the West CZACC 6120, Santiago de Cuba, CZACC Indian lizard, Anolis angusticeps, with the descrip- 7263; Playa Siboney, Santiago de Cuba, tion of a new form, Anolis angusticeps paternus, subsp. CZACC 7445; Sardinero, Santiago de Cuba, nov., from the Isle of Pines, Cuba (Reptilia: Igua- nidae). Carib. J. Sci. 6(1-2):23-31, CZACC 7428; San Ramon, Campechuela, Hass, C. A., S. B. Hedges, and L. R. Maxson. 1993, Granma, MCZ 59248-50 (skeleton); Finca Molecular insights into the relationships and bio- La Bahia, 32 km SW de Victoria de las Tu- geography of West Indian anoline lizards. Bio- nas, La Tunas, MCZ 59251-52; 9 km SE of them. Syst. Ecol. 21:97-114. Ruibal, R. 1964. An annotated checklist and key to Mea, Holguin, MCZ 59253; Rio Ovando the anoline lizards of Cuba, Bull. Mus. Comp. Zool. Arriba, Maisi, Guantanamo, MCZ 42469, 130(8):476-520. 42526; Cuchillas de Guajimero, 23.37N, Schwartz, A., and R. Thomas. 1968. A review of 75.04W, near Guayabal de Yateras, Guan- Anolis angusticeps in the West Indies. Quart, J, Flor- tanamo, MCZ 42558; Pico Turquino, Gua- ida Acad. Sci. 31(1):52-69. Williams, E. E. 1976. West Indian anoles: a taxonom- ma, Santiago de Cuba, MCZ 50154; Playa ic and evolutionary summary. 1. Introduction and Jaragua, Santiago de Cuba, AMNH 96500.