Comptus weinlandi ( Cope 1868 )

Schools, Molly & Hedges, Blair, 2024, A new forest lizard fauna from Caribbean islands (Squamata, Diploglossidae, Celestinae), Zootaxa 5554 (1), pp. 1-306 : 126-130

Comptus weinlandi ( Cope 1868)

Hispaniolan Keeled Forest Lizard

(Fig. 45–46)

Celestus stenurus weinlandii Cope, 1868:125 . Holotype: USNM 12145 , received from T. Younglove from within 25 mi of Port-au-Prince, Haiti, between January and June 1866.

Celestus rugosus — Cope, 1879:272. Holotype: ANSP 10260 , from Puerto Plata, Dominican Republic.

Diploglossus rugosus — Boulenger, 1885:288.

Celestus rugosus — Barbour, 1930:99.

Celestus rugosus — Barbour, 1935:123.

Celestus rugosus — Barbour, 1937:138.

Diploglossus stenurus weinlandi — Schwartz, 1964:10.

Diploglossus stenurus rugosus — Schwartz, 1964:14.

Celestus stenurus rugosus — Schwartz & Henderson, 1988:100.

Celestus stenurus weinlandi — Schwartz & Henderson, 1988:101.

Celestus stenurus rugosus — Schwartz & Henderson, 1991:379.

Celestus stenurus weinlandi — Schwartz & Henderson, 1991:379.

Celestus stenurus rugosus — Hedges et al., 2019:17.

Celestus stenurus weinlandi — Hedges et al., 2019:17.

Comptus stenurus rugosus — Schools & Hedges, 2021:226.

Comptus stenurus weinlandi — Schools & Hedges, 2021:226.

Material examined (n=53). DOMINICAN REPUBLIC. Duarte. ANSP 38529–31 , Richard Thomas, Manuel Leal, and Herman Dominicia, 10.1 km NE of San Francisco de Macoris , 15 July 1993 . El Seibo. ANSP 38523 , S. Blair Hedges, Richard Thomas, and Nicholas Plummer, 4.2 km N, 8.4 km W (airline), Nisibon, Playa Cucharera (= beach W Punta Limon), 14 July 1991 ; ANSP 38528 , Richard Thomas, 5 km airline W Sabana de Nisibon , 6 August 2001 . Hato Mayor. ANSP 38513–4 , Richard Thomas, 5.6 km airline W El Valle, 2 August 2001 ; ANSP 38515–22 , Andres Perez, Dan Rabosky, and Richard Thomas, Loma del Fresca, 5.7 km airline SW Sabana de la Mar , 3 August 2001 ; ANSP 38524–7 , Andres Perez, Dan Rabosky, and Richard Thomas, 7.8 km S of Sabana de la Mar , 2 August 2001 ; SBH 267001 , Andres Perez, Dan Rabosky, and Richard Thomas, Loma del Fresca, 5.7 km airline SW Sabana de la Mar , 3 August 2001 ; SBH 267017 , SBH 267031 , Andres Perez, Dan Rabosky, and Richard Thomas, 7.8 km S Sabana de la Mar , 2 August 2001 . Independencia. ANSP 38547–8 , S. Blair Hedges, Kristin Nastase, Renee Sharp, and Patrick Parker, 5.1 km NW of La Descubierta , 30 May 1996 ; SBH 194492–3 , S. Blair Hedges, Kristin Nastase, Renee Sharp, and Patrick Parker, 5.1 km NW of La Descubierta , 30 May 1996 . Los Tabucos. ANSP 38534–7 , Richard Thomas, 8.8 km N, thence 0.5 km W Tenares , Salcedo, 21 July 1993 . Maria Trinidad Sanchez. MCZ R-126742, Alan Ross Kiester, 15 km S of Nagua , 14 September 1944 – 29 January 1971 . Puerto Plata. USNM 10260 , Puerto Plato (= San Felipe de Puerto Plata), 1878 . Salcedo. ANSP 38532–3 , Richard Thomas, 23.2 km N of thence 4.5 km W Tenares, = 0.2 km E Jaiba , 21 July 1993 ; SBH 193218 , Richard Thomas, 23.2 km N of thence 4.5 km W Tenares, = 0.2 km E Jaiba , 21 July 1993 . HAITI. Artibonite. ANSP 38542 , S. Blair Hedges, Richard Thomas, Nicholas Plummer, and Manuel Leal, 11.8 km W of Ça Soleil , 11 June 1991 ; SBH 192406 , 192408 , S. Blair Hedges, Richard Thomas, Nicholas Plummer, and Manuel Leal, 11.8 km W of Ça Soleil , 11 June 1991 . Nord’Ouest. ANSP 38541 , S. Blair Hedges and Richard Thomas, Bombardopolis , 28 April 1997 . Ouest. ANSP 38549 , S. Blair Hedges, Richard Thomas, Manuel Leal, and Nicholas Plummer, 10.1 km ENE of Petionville , 9 June 1991 ; ANSP 38551–4 , S. Blair Hedges, Richard Thomas, Nicholas Plummer, and Manuel Leal, 18.7 km E of Thomaseau , 19 May 1991 ; MCZ R-163165, E. Wade Davis, Mariani (on Port-au-Prince to Leogane road), 1–31 July ; SBH 191620 , 191649–50 , S. Blair Hedges, Richard Thomas, Manuel Leal, and Nicholas Plummer , 18.1 km E of Thomaseau, 19 May 1991 ; SBH 191622–4 , S. Blair Hedges, Richard Thomas, Nicholas Plummer, and Manuel Leal, 18.7 km E of Thomaseau , 19 May 1991 ; SBH 192424 , S. Blair Hedges Richard Thomas Manuel Leal, and Nicholas Plummer, 10.1 km ENE of Petionville , 9 June 1991 ; USNM 12145 , within 25 mi of Port-au-Prince, January–June 1866 .

FIGURE 45. (A–F) Comptus weinlandi ( USNM 12145, holotype), SVL 94.0 mm.

Diagnosis. Comptus weinlandi has (1) a dorsal pattern of irregular dots/dots in series/dots in chevrons, (2) head markings absent/present, (3) markings in the longitudinal paramedian area absent/present, (4) dots arranged in bars in the lateral band absent/present, (5) an adult SVL of 101–133 mm, (6) ventral scale rows, 81–105, (7) midbody scale rows, 37–44, (8) total lamellae on one hand, 43–55, (9) total strigae on ten scales, 167–236, (10) relative length of all digits on one hindlimb, 24.5–36.5 %, (11) relative distance between the angled subocular and mouth, 0.449 –0.901 %, (12) relative eye length, 3.33–4.43 %, (13) relative forelimb length, 20.1–27.9 %, (14) relative ear width, 0.802–2.18 %, (15) relative rostral height, 1.46–1.86 %, (16) relative head length, 15.8–18.5 %, (17) relative mental width, 1.41–1.90 %, (18) relative postmental width, 2.57–2.91 %, (19) relative cloacal width, 8.76–10.1 %, (20) relative prefrontal width, 4.29–4.88 %, (21) relative largest supraocular width, 2.31–3.49 %, (22) relative longest finger length, 5.08–6.31 %, (23) relative distance between the ear and eye, 6.03–7.95 %, (24) relative head width, 73.6–82.2 %, (25) relative frontal width, 58.8–84.6 %, (26) relative nasal height, 0.965–1.32 %, (27) relative angled subocular height, 0.568–1.23 %, (28) relative distance between the eye and naris, 4.67–5.72 %, (29) relative canthal iii length, 1.65–2.08 %, (30) relative angled subocular width, 1.64–3.36 %, and (31) relative nasal length, 1.48–1.95 %. The species stem time is 1.46 Ma and the species crown time is 0.68 Ma (Fig. 4).

We distinguish Comptus weinlandi from the other species of Comptus based on a complex of traits. From Comptus alloeides , we distinguish C. weinlandi by the total strigae on ten scales (167–236 versus 237–323). From C. arboreus sp. nov., we distinguish C. weinlandi by the relative length of digits on one hindlimb (24.5–36.5 versus 37.4–39.7) and the relative postmental width (2.57–2.91 versus 2.95–3.01). From C. badius , we distinguish C. weinlandi by the adult SVL (101–133 versus 78.2–99.1), the relative mental width (1.41–1.90 versus 1.38), the relative postmental width (2.57–2.91 versus 2.39), the relative longest finger length (5.08–6.31 versus 4.38–5.04), and the relative head width (73.6–82.2 versus 62.8–69.3). From C. maculatus , we distinguish C. weinlandi by the dorsal pattern (irregular dots/dots in series/dots in chevrons versus absent/chevrons), the adult SVL (101–133 versus 60.1–81.3), the total lamellae on one hand (43–55 versus 32–37), and the relative longest finger length (5.08–6.31 versus 4.14–5.01). From C. stenurus , we cannot distinguish C. weinlandi based on our standard suite of characters (see Remarks).

Description of holotype. USNM 12145. An adult; SVL 94.0 mm; tail nearly cylindrical, broken, 15.5 mm (16.5% SVL); axilla-to-groin distance 44.7 mm (47.6% SVL); forelimb length 22.0 mm (23.4% SVL); hindlimb length 35.3 mm (37.6% SVL); head length 16.5 mm (17.6% SVL); head width 11.3 mm (12.0% SVL); head width 68.5% head length; diameter of orbit 3.93 mm (4.18% SVL); horizontal diameter of ear opening 1.05 mm (1.12% SVL); vertical diameter of ear opening 1.44 mm (1.53% SVL); length of all toes on one foot 31.9 mm (33.9% SVL); shortest distance between angled subocular and lip 0.61 mm (0.649% SVL); shortest distance between the ocular and auricular openings 6.38 mm (6.79% SVL); longest finger length 5.75 mm (6.12% SVL); largest supraocular width 2.46 mm (2.62% SVL); cloacal width 8.66 mm (9.21% SVL); mental width 1.70 mm (1.81% SVL); postmental width 2.52 mm (2.68% SVL); prefrontal width 4.79 mm (5.10% SVL); frontal width 78.0% frontal length; nasal height 1.19 mm (1.27% SVL); angled subocular height 0.88 mm (0.936% SVL); shortest distance between the eye and naris 4.49 mm (4.78% SVL); canthal iii width 1.79 mm (1.90% SVL); angled subocular width 2.35 mm (0.250% SVL); nasal width 1.66 mm (1.77% SVL); rostral 1.86X as wide as high, visible from above, not in contact with nasals, in contact with 1 st supralabial and anterior internasal (left)/(right); anterior internasals are narrower than posterior ones; frontonasals and prefrontal fused into a single large plate with a concave posterior margin, much wider than long, bordered by posterior internasals, 1 st loreals, 1 st and 2 nd median oculars, and the frontal; frontal longer than wide; a pair of frontoparietals, separated by the posterior prolongation of the frontal and the interparietal plate; interparietal plate smaller than parietals and separating them, posteriorly touching the interoccipital, which is wider than long; parietal separated from supraoculars by 1 st temporals and frontoparietal (left)/(right); nasal single; nostril above suture between 1 st and 2 nd supralabials (left)/(right); 1 postnasal (left)/(right); 2 (left)/3 (right) loreals; 1 st loreal higher than wide (left)/small (right), in contact with postnasal, posterior internasal, prefrontal/frontonasal complex, 1 st median ocular, canthal iii, 2 nd loreal (fused with canthal iii), and 3 rd –4 th supralabials (left); 2 nd loreal shorter than 1 st, approximately as high as wide, fused with canthal iii (left)/higher than wide, in contact with posterior internasal, prefrontal/frontonasal complex, median ocular 1, canthal iii, supralabials 3–4, and the 1 st –3 rd labials (right), excluded from contact with supraocular by canthal iii (fused with loreal 2) (left); 3 rd loreal shorter than 1 st, approximately as high as wide (left); final loreal posteriorly bordering the lower preocular (left)/(right); canthal iii wider than high, fused with loreal 2 (left)/wider than high (right), contacting 1 st median ocular (fused with), anterior supraciliary, upper and lower preoculars, 1 st and 2 nd loreals, and the additional scale (left)/1 st median ocular, anterior supraciliary, upper and lower preoculars, and 1 st and 2 nd loreals (right); 10 median oculars (left)/(right), 1 st and 2 nd contacting the prefrontal (left)/(right); 1 upper preocular (left)/(right); an irregular anterior supraciliary (left)/(right); 6 lateral oculars (left)/(right); 5 temporals (left)/(right); 2 suboculars (left)/(right); posterior subocular large and elongate (left)/(right); anterior subocular small (left)/(right); 9 (left)/10 (right) supralabials, 6 to level below center of eye (left)/(right); 9 (left)/10 (right) infralabials, 6 to level below center of eye (left)/(right); mental small, followed by a single, larger post mental; 4 pairs of enlarged chin shields, followed by 1 pair of reduced chin shields; 1 st pair in contact with one another anteriorly, posteriorly separated by one scale; 2 nd pair in contact with one another anteriorly, posteriorly separated by one scale; 3 rd –5 th pairs separated by 1–4 scales; 93 transverse rows of ventral scales from mental to vent; 39 scales around midbody; 5 digits; finger lengths 3>4>5>2>1; 12 lamellae under longest finger (left)/(right); 50 total lamellae on one hand; toe lengths 4>3>5>2>1; 21 (left)/19 (right) lamellae under longest toe; strigae and median keel (more so on anterior scales, posterior have little to no keel) dorsal body and caudal scales; smooth ventral scales; 185 total strigae counted on ten scales.

Color (in alcohol): dorsal surface of head gray-brown, patternless; lateral surfaces of head grading from gray-brown to yellow-orange; dorsal surfaces of the body are gray-brown with faded brown markings in longitudinal paramedian series that continue down the body; dorsal surface of tail the same as the body; lateral areas grade from gray-brown to orange-yellow; dorsal surfaces of the limbs are gray-brown with some orange mottling; lateral and ventral areas of the limbs grade to yellow-orange, patternless; ventral surfaces of the head, body, and tail are yellow-orange, patternless.

Variation. The majority of the examined material examined have a similar pattern of large dots that are continuations of the longitudinal paramedian series. Several specimens exhibit more irregular dots in dots also arranged in chevrons. All specimens except for ANSP 38525 show marking in the longitudinal paramedian series that range from dots in series to complete longitudinal paramedian lines. Most specimens show a pattern on their head scales in the form of these scales either having darker outlines or irregular, darker dots. A minority of the specimens have a completely patternless head. All specimens that are not faded show dots arranged in bars in the lateral area except for USNM 10260, in which this trait is absent. Measurements and other morphological data for the holotype and other examined material are presented in Table 1.

Distribution. Comptus weinlandi , with the largest range in the genus Comptus , is distributed throughout most of the island and in both countries at elevations of 0–1060 m. However, it is missing from nearly all the Tiburon Peninsula, Barahona Peninsula, Central Haiti, West-Central Dominican Republic, and offshore islets (Fig. 35). It has an extent of occurrence of ~ 41,680 km 2.

Ecology and conservation. No ecological data are available for this species. We consider the conservation status of Comptus weinlandi to be Least Concern, based on IUCN Redlist criteria ( IUCN 2023). It is a common species tolerant of some habitat disturbance. However, it faces a primary threat of habitat destruction resulting from agriculture and urbanization. A secondary threat is predation from introduced mammals, including the mongoose and black rats. We need studies to determine the health and extent of remaining populations and better understand the threats to the survival of the species.

Reproduction. Ovoviviparous. Embryos observed in ANSP 38535.

Etymology. This species was named after David Friedrich Weinland for his contributions to the study of reptiles in Haiti. Weinland made a six-month visit to Jérémie, Haiti, in 1857–1858 ( Weinland 1858).

Remarks. The type locality of Comptus weinlandi was given incorrectly as Gonave island in the type description ( Cope 1868); however, this error was later noted and corrected ( Schwartz 1964). The correct type locality is from within 25 miles of Port-au-Prince, Haiti. Both C. rugosus and C. weinlandi were previously considered to be subspecies of “ Celestus ” (= Comptus ) stenurus ( Schwartz 1964) . Because of a lack of genetic and morphological differentiation, we synonymize C. stenurus rugosus under the name C. weinlandi . Schwartz (1964) also was unable to find a diagnostic character to separate the two taxa, although he found some non-diagnostic trends in pattern.

Comptus weinlandi and C. stenurus cannot be distinguished based on our suite of morphological characters; however, both species are morphologically distinct from their respective closest relatives ( C. alloeides and C. arboreus , respectively). Additionally, C. weinlandi and C. stenurus are genetically distinct (Fig. 3) and diverged 4.22 Ma (Fig. 4). Future studies should examine additional characters to morphologically distinguish C. weinlandi and C. stenurus .

Comptus weinlandi is included in our genetic dataset and has a significant support value in our ML analysis and a support value of 92% in our Bayesian analysis at the node that defines it as a species. The stem node that places C. weinlandi as the closest relative to C. alloeides has a significant support value in both our ML and Bayesian analyses. Based on our timetree (Fig. 4), C. weinlandi diverged from its closest relative 1.46 Ma, consistent with typical species of vertebrates (> 0.7 Ma; Hedges et al. 2015). We further recognize it as a distinct species because of the diagnostic trait that separates it from C. alloeides (the total strigae on ten scales). This species is also sympatric with C. alloeides at one locality on the Samana Peninsula. Comptus weinlandi was recovered as conspecific with Comptus alloeides in our ASAP analysis.

FIGURE 46. Comptus weinlandi (SBH 267796), in life. From an unnamed beach near Cueva Infierno and Cueva del Angel, Hato Mayor Province, Dominican Republic. Photo by SBH.

Genus Guarocuyus ( Landestoy et al. 2022)

Jaragua Forest Lizards

Type species. Guarocuyus jaraguanus Landestoy et al., 2022: 201 .

Diagnosis. See Guarocuyus jaraguanus Diagnosis section.

Content. One species (Table 3): Guarocuyus jaraguanus .

Distribution. See Guarocuyus jaraguanus Distribution section.