Celestus capitulatus, Schools & Hedges, 2024

Schools, Molly & Hedges, Blair, 2024, A new forest lizard fauna from Caribbean islands (Squamata, Diploglossidae, Celestinae), Zootaxa 5554 (1), pp. 1-306 : 45-49

Celestus capitulatus sp. nov.

Southwestern Jamaican Forest Lizard

(Fig. 15)

Celestus crusculus crusculus — Grant, 1940a:102.

Celestus crusculus — Schwartz & Henderson, 1991:369 (part).

Celestus crusculus crusculus — Hedges et al., 2019:17 (part).

Celestus crusculus — Schools & Hedges, 2021:220 (part).

Celestus crusculus — Landestoy et al., 2022:204 (part).

Holotype. USNM 328168 , an adult male from Knoxwood, Saint Elizabeth Parish, Jamaica, collected by Carla Ann Hass, S. Blair Hedges, Minocal Stephenson, and Walton Stephenson on 2 October 1984 (18.00363, -77.74489; 8 m).

Paratypes (n=35). JAMAICA. Hanover. BMNH1970.1713 , Garth Underwood, Booby Cay ,1970 . Manchester. BMNH 1938.4 .13.8, Chapman Grant, Alligator Pond , southern coast, 1938 ; BMNH 1970.1721 – 1724 (one of several untagged specimens in the same jar), Garth Underwood, Alligator Pond , 1970 . Saint Elizabeth. USNM 328169 , Carla Ann Hass, S. Blair Hedges, Minocal Stephenson, and Walton Stephenson, Knoxwood , 2 October 1984 ; KU 229277 , KU 229279–82 , 3 mi SE Whitehouse , 10 July 1967 ; BMNH 1970.1719 , Garth Underwood, Black River , 1970 ; AMNH 72365 , 139138–9 , Koopman, Hecht, and Williams, 1 mi S of Black River , 13 August 1950 .

Westmoreland. BMNH 1970.1717 , Garth Underwood, Negril Hill, Holmes Bay, 1970; BMNH 1970.1718 , Garth Underwood, north of Springfield , 1970 ; BMNH 1970.1715 – 1716 , Garth Underwood, Negril , 1970 ; KU 229232–3 , 2 mi SW Old Hope , 1 July 1967 ; KU 229244 , 0.3 mi NW Whitehouse ; KU 229250–1 , 0.5 mi SE Whitehouse , 14 July 1967 ; KU 229252–8 , 229260 , 0.7 mi NW Bluefields , 17 July 1967 ; KU 229273–4 , 0.1 mi N Beeston Spring , 18 July 1967 ; KU 229275–6 , 3 mi N Kilmarnoch , 18 July 1967 ; USNM 328157 , S. Blair Hedges and Carla Ann Hass, 4.5 km W of Old Hope (at Little Bay), 29 May 1988 .

Diagnosis. Celestus capitulatus sp. nov. has (1) a dorsal pattern of irregular dots/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 62.1–81.8 mm, (6) ventral scale rows, 97–121, (7) midbody scale rows, 37–47, (8) total lamellae on one hand, 25–38, (9) total strigae on ten scales, 105–192, (10) relative length of all digits on one hindlimb, 17.6–22.3 %, (11) relative distance between the angled subocular and mouth, 0.525–1.17 %, (12) relative eye length, 2.75–3.80 %, (13) relative forelimb length, 14.3–18.1 %, (14) relative ear width, 0.671–2.04 %, (15) relative rostral height, 1.51–2.03 %, (16) relative head length, 15.1–17.7 %, (17) relative mental width, 1.28–1.84 %, (18) relative postmental width, 2.62–2.97 %, (19) relative cloacal width, 7.84–8.67 %, (20) relative prefrontal width, 4.30–4.72 %, (21) relative largest supraocular width, 2.03–2.61 %, (22) relative longest finger length, 3.45–3.75 %, (23) relative distance between the ear and eye, 6.45–7.84 %, (24) relative head width, 71.6–78.6 %, (25) relative frontal width, 78.1–81.6 %, (26) relative nasal height, 0.953–1.42 %, (27) relative angled subocular height, 0.586–1.01 %, (28) relative distance between the eye and naris, 4.57–5.03 %, (29) relative canthal iii length, 1.61–1.70 %, (30) relative angled subocular width, 1.93–2.32 %, and (31) relative nasal length, 1.40–1.84 %. The species stem time is 4.85 Ma and the species crown time is 1.27 Ma (Fig. 4).

We distinguish Celestus capitulatus sp. nov. from congeners based on a complex of traits. From Celestus barbouri , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus chevrons). From C. crusculus , we distinguish C. capitulatus sp. nov. by the relative frontal width (78.1–81.6 versus 82.6–91.1). From C. duquesneyi , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus bands), the midbody scale rows (37–47 versus 48), the total lamellae on one hand (25–38 versus 64), the relative length of digits on one hindlimb (17.6–22.3 versus 31.4), the relative eye length (2.75–3.80 versus 4.36), the relative forelimb length (14.3–18.1 versus 24.4), the relative ear width (0.671–2.04 versus 2.45), the relative rostral height (1.51–2.03 versus 2.14), the relative head length (15.1–17.7 versus 21.6), the relative mental width (1.28–1.84 versus 2.35), the relative postmental width (2.62–2.97 versus 3.19), the relative cloacal width (7.84–8.67 versus 9.98), the relative prefrontal width (4.30–4.72 versus 5.41), the relative largest supraocular width (2.03–2.61 versus 2.66), the relative longest finger length (3.45–3.75 versus 6.52), the relative head width (71.6– 78.6 versus 64.6), the relative frontal width (78.1–81.6 versus 75.2), the relative angled subocular height (0.586– 1.01 versus 1.61), the relative distance between the eye and naris (4.57–5.03 versus 5.46), the relative angled subocular width (1.93–2.32 versus 2.90), and the relative nasal width (1.40–1.84 versus 2.01). From C. hesperius sp. nov., we distinguish C. capitulatus sp. nov. by the relative forelimb length (14.3–18.1 versus 18.6–21.3) and the relative width of canthal iii (1.61–1.70 versus 1.77–1.93). From C. hewardi , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus mottled/bands), the adult SVL (62.1–81.8 versus 129– 171), the total lamellae on one hand (25–38 versus 50–61), the relative length of digits on one hindlimb (17.6–22.3 versus 24.1–30.6), the relative forelimb length (14.3–18.1 versus 22.2–24.6), the relative cloacal width (7.84–8.67 versus 8.81–9.89), the relative longest finger length (3.45–3.75 versus 5.03–5.66), and the relative frontal width (78.1–81.6 versus 57.3–75.3). From C. jamesbondi sp. nov., we distinguish C. capitulatus sp. nov. by the relative frontal width (78.1–81.6 versus 70.5–77.6) and the relative width of canthal iii (1.61–1.70 versus 1.75–2.16). From C. macrolepis , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus bicolored), the adult SVL (62.1–81.8 versus 254–316), the total lamellae on one hand (25–38 versus 52–54), the total strigae on ten scales (105–192 versus 398), the relative length of digits on one hindlimb (17.6–22.3 versus 27.5–28.0), the relative distance between angled subocular and mouth (0.525–1.17 versus 1.39–1.66), the relative forelimb length (14.3–18.1 versus 26.1–26.7), the relative head length (15.1–17.7 versus 19.2–22.9), the relative mental width (1.28–1.84 versus 1.87), the relative postmental width (2.62–2.97 versus 3.81), the relative cloacal width (7.84–8.67 versus 11.2), the relative prefrontal width (4.30–4.72 versus 3.94), the relative largest supraocular width (2.03–2.61 versus 2.64–3.01), the relative longest finger length (3.45–3.75 versus 5.47–5.51), the relative distance between the ear and eye (6.45–7.84 versus 8.02–10.9), the relative head width (71.6–78.6 versus 80.5), the relative angled subocular height (0.586–1.01 versus 1.17), the relative distance between the eye and naris (4.57–5.03 versus 6.02), and the relative angled subocular width (1.93–2.32 versus 2.57). From C. macrotus , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus chevrons/bands), the ventral scale rows (97–121 versus 87–93), the total lamellae on one hand (25–38 versus 39–40), the relative length of digits on one hindlimb (17.6–22.3 versus 30.2–31.2), the relative forelimb length (14.3–18.1 versus 22.4–25.0), the relative head length (15.1–17.7 versus 18.2–20.5), the relative postmental width (2.62–2.97 versus 3.00), the relative prefrontal width (4.30–4.72 versus 4.87–5.55), the relative largest supraocular width (2.03–2.61 versus 2.96–4.03), the relative longest finger length (3.45–3.75 versus 6.43–6.67), the relative frontal width (78.1–81.6 versus 57.6– 66.1), the relative distance between the eye and naris (4.57–5.03 versus 5.48–5.60), the relative angled subocular width (1.93–2.32 versus 2.77–2.83), and the relative nasal width (1.40–1.84 versus 2.08–2.33). From C. microblepharis , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus chevrons), the adult SVL (62.1–81.8 versus 96.4), the relative length of digits on one hindlimb (17.6–22.3 versus 16.6), the relative eye length (2.75–3.80 versus 1.83), the relative forelimb length (14.3–18.1 versus 14.2), the relative ear width (0.671–2.04 versus 0.446), the relative head length (15.1–17.7 versus 14.7), the relative postmental width (2.62–2.97 versus 2.47), the relative longest finger length (3.45–3.75 versus 3.11), the relative nasal height (0.953–1.42 versus 0.726), the relative angled subocular width (1.93–2.32 versus 2.90), and the relative nasal width (1.40–1.84 versus 1.11). From C. molesworthi , we distinguish C. capitulatus sp. nov. by the relative length of digits on one hindlimb (17.6–22.3 versus 22.4–29.4), the relative cloacal width (7.84–8.67 versus 8.73–9.35), the relative longest finger length (3.45–3.75 versus 4.28–5.19), the relative distance between the ear and eye (6.45–7.84 versus 7.97–8.83), the relative angled subocular height (0.586–1.01 versus 1.11), the relative distance between the eye and naris (4.57–5.03 versus 5.32–5.50), and the relative width of canthal iii (1.61–1.70 versus 1.99–2.09). From C. occiduus , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus absent), the adult SVL (62.1–81.8 versus 269–367), the total lamellae on one hand (25–38 versus 50–66), the total strigae on ten scales (105–192 versus 374), the relative length of digits on one hindlimb (17.6–22.3 versus 24.4–29.7), the relative distance between angled subocular and mouth (0.525–1.17 versus 1.26–1.27), the relative forelimb length (14.3–18.1 versus 23.5–23.9), the relative head length (15.1–17.7 versus 20.4–20.6), the relative mental width (1.28–1.84 versus 1.86), the relative postmental width (2.62–2.97 versus 3.57), the relative cloacal width (7.84–8.67 versus 9.00), the relative prefrontal width (4.30–4.72 versus 4.76), the relative longest finger length (3.45–3.75 versus 4.77–5.46), the relative distance between the ear and eye (6.45–7.84 versus 8.98–10.9), the relative frontal width (78.1–81.6 versus 63.8), the relative angled subocular height (0.586–1.01 versus 1.30), the relative distance between the eye and naris (4.57–5.03 versus 6.51), and the relative angled subocular width (1.93–2.32 versus 2.52). From C. oligolepis sp. nov., we distinguish C. capitulatus sp. nov. by the midbody scale rows (37–47 versus 35). From C. striatus , we distinguish C. capitulatus sp. nov. by the dorsal pattern (irregular dots/dots in chevrons versus absent/chevrons), the adult SVL (62.1–81.8 versus 145), the total lamellae on one hand (25–38 versus 59–66), the total strigae on ten scales (105–192 versus 279), the relative length of digits on one hindlimb (17.6–22.3 versus 37.8), the relative eye length (2.75–3.80 versus 3.85), the relative forelimb length (14.3–18.1 versus 26.1), the relative head length (15.1–17.7 versus 18.9), the relative prefrontal width (4.30–4.72 versus 5.68), the relative largest supraocular width (2.03–2.61 versus 2.63), the relative longest finger length (3.45–3.75 versus 7.48), the relative distance between the ear and eye (6.45–7.84 versus 9.00), the relative head width (71.6–78.6 versus 82.1), the relative frontal width (78.1–81.6 versus 76.5), the relative angled subocular height (0.586–1.01 versus 1.12), and the relative distance between the eye and naris (4.57–5.03 versus 6.16).

Description of holotype. USNM 328168. An adult male; SVL 71.7 mm; tail nearly cylindrical, 6.97 mm (9.72% SVL); axilla-to-groin distance 40.3 mm (56.2% SVL); forelimb length 12.9 mm (18.0% SVL); hindlimb length 18.3 mm (25.5% SVL); head length 12.7 mm (17.7% SVL); head width 9.98 mm (13.9% SVL); head width 78.6% head length; diameter of orbit 2.44 mm (3.40% SVL); horizontal diameter of ear opening 1.46 mm (2.04% SVL); vertical diameter of ear opening 2.10 mm (2.93% SVL); length of all toes on one foot 15.7 mm (21.9% SVL); shortest distance between angled subocular and lip 0.42 mm (0.586% SVL); shortest distance between the ocular and auricular openings 5.62 mm (7.84% SVL); longest finger length 2.66 mm (3.71% SVL); largest supraocular width 1.87 mm (2.61% SVL); cloacal width 5.62 mm (7.84% SVL); mental width 0.92 mm (1.28% SVL); postmental width 1.88 mm (2.62% SVL); prefrontal width 3.29 mm (4.59% SVL); frontal width 78.1% frontal length; nasal height 1.02 mm (1.42% SVL); angled subocular height 0.42 mm (0.586% SVL); shortest distance between the eye and naris 3.32 mm (4.63% SVL); canthal iii width 1.22 mm (1.70% SVL); angled subocular width 1.66 mm (2.32% SVL); nasal width 1.21 mm (1.69% SVL); rostral 1.62X as wide as high, barely 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 straight posterior margin, wider than long, bordered by posterior internasals, 1 st loreals, canthal iii, 1 st median oculars, and the frontal; frontal much 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 and 2 nd temporals (missing the 1 st on the left) and frontoparietal (left)/(right); nasal single; nostril above suture between 1 st and 2 nd supralabials (left)/(right); 1 postnasal (left)/ (right); 2 loreals (left)/(right); 1 st loreal higher than wide (left)/(right), in contact with postnasal, posterior internasal, prefrontal/frontonasal complex, canthal iii, 2 nd loreal, and 3 rd –4 th supralabials (left)/(right); 2 nd loreal shorter than 1 st, approximately as high as wide (left)/(right), excluded from contact with supraocular by canthal iii (left)/(right); 2 nd loreal posteriorly bordering the lower preocular (left)/(right); canthal iii wider than high (left)/(right), contacting 1 st median ocular, anterior supraciliary, upper and lower preoculars, prefrontal/frontonasal complex, and 1 st and 2 nd loreals (left)/(right); 10 (left)/9 (right) median oculars, 1 st contacting the prefrontal (left)/(right); 1 upper preocular (left)/(right); an irregular anterior supraciliary (left)/(right); 5 (left)/6 (right) lateral oculars; 5 temporals (left)/(right); 2 suboculars (left)/(right); posterior subocular large and elongate (left)/(right); anterior subocular small (left)/(right); 9 supralabials (left)/(right), 6 to level below center of eye (left)/(right); 9 infralabials (left)/(right), 6 to level below center of eye (left)/(right); mental small, followed by a single, larger postmental; 4 pairs of enlarged chin shields; 1 st pair in contact with one another; 2 nd –4 th pairs separated by 1–3 scales; 118 transverse rows of dorsal scales from interoccipital to base of tail; 113 transverse rows of ventral scales from mental to vent; 44 scales around midbody; 5 digits; finger lengths 3>4>2>5>1; 8 lamellae under longest finger (left)/(right); 32 total lamellae on one hand; toe lengths 4>3>5>2>1; 13 lamellae under longest toe (left)/(right); striate and slightly keeled dorsal body and caudal scales; smooth ventral scales; 105 total strigae counted on ten scales.

Color (in alcohol): dorsal surface of head pale brown, patternless; lateral surfaces of head grading from pale brown to yellow-cream with darker brown eye masks and markings on the labial scales; dorsal surfaces of the body are pale brown with two darker broken longitudinal paramedian lines and dotted chevrons; dorsal surface of tail the same as the body; lateral areas grade from pale brown to cream with the chevrons from the dorsum continuing; dorsal surfaces of the limbs are medium brown, patternless; lateral and ventral areas of the limbs fade to cream, patternless; ventral surfaces of the head, body, and tail are cream with some darker markings on the chin shields.

FIGURE 15. (A–F) Celestus capitulatus sp. nov. (USNM 328168, holotype), SVL 71.7 mm.

Variation. The dorsal ground color of all examined specimens is pale to medium brown with darker brown dots that form broken chevrons that stretch across the dorsums. All have cream, virtually patternless venter. The throats of both paratypes have a darker gray tone than the rest of their cream-colored venter. Markings in the longitudinal paramedian area range from absent to mottled to bearing small lines with the holotype having the most pronounced longitudinal paramedian lines. The majority of specimens do not exhibit dots arranged in bars in the lateral bands, many of those that do show mottling as opposed to clear dots. Measurements and other morphological data for the holotype and other examined material are presented in Table 1.

Distribution. Celestus capitulatus sp. nov. is distributed on the southwestern coast of Jamaica in Westmoreland and St. Elizabeth at elevations of 0–530 m (Fig. 12). It has a range size of 1,130 km 2.

Ecology and conservation. Little is known of the ecology of this species other than the holotype and one other animal collected at the same time were both collected in rotting vegetation. A different specimen (USNM 328157) was collected at dusk in sea grape leaf litter. Given the large number of locations (Fig. 12) and occurrence in coastal dry areas as well as inland mesic areas, this species is likely common and tolerant of human disturbance.

We consider the conservation status of Celestus capitulatus sp. nov. to be Least Concern, based on IUCN Redlist criteria ( IUCN 2023). However, its relatively small range is of concern, and therefore studies are needed to determine the health and extent of the populations, and any threats to the survival of the species.

Reproduction. No data on reproduction are available for this species.

Etymology. The species name ( capitulatus ) is a masculine nominative singular adjective (Latin) meaning “small head,” in reference to the relatively small average head length of adults in this species.

Remarks. Specimens of this species were observed in the majority of the museum collections examined (AMNH, BMNH, KU, and USNM), having been collected as recently as the 1980s. Celestus capitulatus sp. nov. is included in our genetic dataset and has significant support in both Bayesian and ML likelihood analyses at the crown node. The stem node that places it outside of the group including Celestus duquesneyi , Celestus hesperius sp. nov., Celestus hewardi , Celestus jamesbondi sp. nov., Celestus macrolepis , Celestus molesworthi , Celestus occiduus , and Celestus striatus had significant support in our ML analysis and 81% in our Bayesian analyses. Based on our timetree (Fig. 4), C. capitulatus sp. nov. diverged from its closest relative 4.85 Ma, consistent with typical species of vertebrates (> 0.7 Ma; Hedges et al. 2015). Celestus capitulatus sp. nov. was recognized as a distinct species by our ASAP analysis.