Celestus macrotus Thomas & Hedges, 1989

Schools, Molly & Hedges, Blair, 2024, A new forest lizard fauna from Caribbean islands (Squamata, Diploglossidae, Celestinae), Zootaxa 5554 (1), pp. 1-306 : 78-83

Celestus macrotus Thomas & Hedges, 1989

Laselle-Baoruco Forest Lizard

(Fig. 26–27)

Celestus macrotus Thomas & Hedges, 1989:886 . Holotype: USNM 286917 collected by S. Blair Hedges and Richard Thomas from ca. 15 km W of Gros Cheval by logging roads, on the northeastern slope of Morne La Selle , Département de l’Ouest,

Haiti, on 18 November 1984 (18.3509, -71.9020; 2020 m). Celestus macrotus — Schwartz & Henderson, 1991:375. Celestus macrotus — Powell et al., 1999:105. Celestus macrotus — Schools & Hedges, 2021:220. Celestus macrolepis — Schools et al. 2022. Celestus macrotus — Landestoy et al., 2022:204.

Material examined (n=7). DOMINICAN REPUBLIC. Pedernales. MALT 00796–99 , 7 km NNE of Los Arroyos, helipad at Loma del Toro . HAITI. Ouest. ANSP 38506 , S. Blair Hedges, southeast of Pic La Selle , 20 November 2011 ; USNM 286917 , S. Blair Hedges and Richard Thomas, ca. 15 km W of Gros Cheval by logging roads, northeastern slope of Morne La Selle in the Massif de la Selle, 18 November 1984 . Sud-Est. ANSP 38505 , S. Blair Hedges, Tiffany Cloud, Miguel Landestoy, and Marcos Rodriguez, Southeast of Pic La Selle , 20 November 2011 .

Diagnosis. Celestus macrotus has (1) a dorsal pattern of chevrons/bands, (2) head markings absent/present, (3) markings in the longitudinal paramedian area present, (4) dots arranged in bars in the lateral band present, (5) a maximum SVL of 60.0– 86.1 mm, (6) ventral scale rows, 87–93, (7) midbody scale rows, 41–45, (8) total lamellae on one hand, 39–40, (9) total strigae on ten scales, 64–115, (10) relative length of all digits on one hindlimb, 30.2–31.2 %, (11) relative distance between the angled subocular and mouth, 0.640 –0.983 %, (12) relative eye length, 3.79–5.17 %, (13) relative forelimb length, 22.4–25.0 %, (14) relative ear width, 1.75–2.08 %, (15) relative rostral height, 1.61–1.95 %, (16) relative head length, 18.2–20.5 %, (17) relative mental width, 1.77 %, (18) relative postmental width, 3.00 %, (19) relative cloacal width, 7.80–9.48 %, (20) relative prefrontal width, 4.87–5.55 %, (21) relative largest supraocular width, 2.96–4.03 %, (22) relative longest finger length, 6.43–6.67 %, (23) relative distance between the ear and eye, 7.58–8.02 %, (24) relative head width, 67.6–80.6 %, (25) relative frontal width, 57.6–66.1 %, (26) relative nasal height, 1.15–1.62 %, (27) relative angled subocular height, 1.00–1.07 %, (28) relative distance between the eye and naris, 5.48–5.60 %, (29) relative canthal iii length, 1.85 %, (30) relative angled subocular width, 2.77–2.83 %, and (31) relative nasal length, 2.08–2.33 %. The species stem time is 5.58 Ma and the species crown time is 1.22 Ma (Fig. 4).

Celestus macrotus has a lower number of ventral scale rows (87–93) than most other species of the genus. This species also has a larger relative longest finger length (6.43–6.67) than most other species of the genus.

From Celestus barbouri , we distinguish C. macrotus by the longitudinal paramedian lines (present versus absent), the dots arranged in bars in the lateral areas (present versus absent), the ventral scale rows (87–93 versus 118–151), the midbody scale rows (41–45 versus 47–56), the relative length of digits on one hindlimb (30.2–31.2 versus 18.2–23.5), the relative distance between angled subocular and mouth (0.640 –0.983 versus 0.437 –0.556), the relative eye length (3.79–5.17 versus 2.87–3.63), the relative forelimb length (22.4–25.0 versus 15.4–19.0), the relative head length (18.2–20.5 versus 14.6–16.6), the relative prefrontal width (4.87–5.55 versus 3.97–4.33), the relative largest supraocular width (2.96–4.03 versus 1.92–2.74), the relative longest finger length (6.43–6.67 versus 2.92–3.81), the relative distance between the ear and eye (7.58–8.02 versus 6.23–7.15), the relative nasal height (1.15–1.62 versus 0.930–1.12), the relative distance between the eye and naris (5.48–5.60 versus 4.68–4.83), the relative angled subocular width (2.77–2.83 versus 1.97–2.52), and the relative nasal width (2.08–2.33 versus 1.38– 1.65). From C. capitulatus sp. nov., we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus irregular dots/dots in chevrons), the ventral scale rows (87–93 versus 97–121), the total lamellae on one hand (39– 40 versus 25–38), the relative length of digits on one hindlimb (30.2–31.2 versus 17.6–22.3), the relative forelimb length (22.4–25.0 versus 14.3–18.1), the relative head length (18.2–20.5 versus 15.1–17.7), the relative postmental width (3.00 versus 2.62–2.97), the relative prefrontal width (4.87–5.55 versus 4.30–4.72), the relative largest supraocular width (2.96–4.03 versus 2.03–2.61), the relative longest finger length (6.43–6.67 versus 3.45–3.75), the relative frontal width (57.6–66.1 versus 78.1–81.6), the relative distance between the eye and naris (5.48–5.60 versus 4.57–5.03), the relative angled subocular width (2.77–2.83 versus 1.93–2.32), and the relative nasal width (2.08–2.33 versus 1.40–1.84). From C. crusculus , we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus absent/flecks in series/dots in chevrons), the dots arranged in bars in the lateral areas (present versus absent), the ventral scale rows (87–93 versus 98–114), the relative length of digits on one hindlimb (30.2–31.2 versus 18.7–24.7), the relative eye length (3.79–5.17 versus 2.93–3.61), the relative forelimb length (22.4–25.0 versus 12.8–20.7), the relative prefrontal width (4.87–5.55 versus 3.93–4.67), the relative largest supraocular width (2.96– 4.03 versus 1.97–2.65), the relative longest finger length (6.43–6.67 versus 2.94–4.10), the relative frontal width (57.6–66.1 versus 82.6–91.1), the relative distance between the eye and naris (5.48–5.60 versus 4.31–4.86), the relative angled subocular width (2.77–2.83 versus 2.03–2.43), and the relative nasal width (2.08–2.33 versus 1.27– 1.60). From C. duquesneyi , we distinguish C. macrotus by the longitudinal paramedian lines (present versus absent), the dots arranged in bars in the lateral areas (present versus absent), the midbody scale rows (41–45 versus 48), the total lamellae on one hand (39–40 versus 64), the relative ear width (1.75–2.08 versus 2.45), and the relative rostral height (1.61–1.95 versus 2.14). From C. hesperius sp. nov., we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus dots in chevrons), the dots arranged in bars in the lateral areas (present versus absent), the ventral scale rows (87–93 versus 111–114), the total lamellae on one hand (39–40 versus 29–34), the relative length of digits on one hindlimb (30.2–31.2 versus 21.7–26.2), the relative eye length (3.79–5.17 versus 3.61–3.74), the relative forelimb length (22.4–25.0 versus 18.6–21.3), the relative ear width (1.75–2.08 versus 1.52–1.59), the relative head length (18.2–20.5 versus 15.7–17.7), the relative largest supraocular width (2.96–4.03 versus 1.91– 2.22), the relative longest finger length (6.43–6.67 versus 3.50–4.04), the relative distance between the ear and eye (7.58–8.02 versus 6.74–7.53), the relative frontal width (57.6–66.1 versus 80.5–86.1), the relative distance between the eye and naris (5.48–5.60 versus 4.70–5.28), the relative angled subocular width (2.77–2.83 versus 2.01–2.48), and the relative nasal width (2.08–2.33 versus 1.52–1.78). From C. hewardi , we distinguish C. macrotus by the adult SVL (60.0–86.1 versus 129–171), the ventral scale rows (87–93 versus 113–137), the total lamellae on one hand (39–40 versus 50–61), the total strigae on ten scales (64–115 versus 164–315), the relative prefrontal width (4.87– 5.55 versus 4.18–4.80), the relative longest finger length (6.43–6.67 versus 5.03–5.66), the relative angled subocular width (2.77–2.83 versus 1.63–2.23), and the relative nasal width (2.08–2.33 versus 1.56–1.88). From C. jamesbondi sp. nov., we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus absent/irregular dots/dots in chevrons), the total lamellae on one hand (39–40 versus 30–36), the relative length of digits on one hindlimb (30.2–31.2 versus 19.8–26.3), the relative forelimb length (22.4–25.0 versus 14.4–19.9), the relative postmental width (3.00 versus 2.61–2.92), the relative largest supraocular width (2.96–4.03 versus 2.16–2.79), the relative longest finger length (6.43–6.67 versus 3.66–4.33), the relative frontal width (57.6–66.1 versus 70.5–77.6), the relative angled subocular width (2.77–2.83 versus 2.09–2.76), and the relative nasal width (2.08–2.33 versus 1.42– 1.75). From C. macrolepis , we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus bicolored), the longitudinal paramedian lines (present versus absent), the dots arranged in bars in the lateral areas (present versus absent), the adult SVL (60.0–86.1 versus 254–316), the ventral scale rows (87–93 versus 112–116), the midbody scale rows (41–45 versus 46–48), the total lamellae on one hand (39–40 versus 52–54), the total strigae on ten scales (64–115 versus 398), the relative length of digits on one hindlimb (30.2–31.2 versus 27.5–28.0), the relative distance between angled subocular and mouth (0.640 –0.983 versus 1.39–1.66), the relative eye length (3.79–5.17 versus 3.63–3.70), the relative forelimb length (22.4–25.0 versus 26.1–26.7), the relative ear width (1.75–2.08 versus 0.760–1.43), and the relative longest finger length (6.43–6.67 versus 5.47–5.51). From C. microblepharis , we distinguish C. macrotus by the longitudinal paramedian lines (present versus absent), the dots arranged in bars in the lateral areas (present versus absent), the adult SVL (60.0–86.1 versus 96.4), the ventral scale rows (87–93 versus 109), the total lamellae on one hand (39–40 versus 30), the relative length of digits on one hindlimb (30.2–31.2 versus 16.6), the relative eye length (3.79–5.17 versus 1.83), the relative forelimb length (22.4–25.0 versus 14.2), the relative ear width (1.75–2.08 versus 0.446), the relative longest finger length (6.43–6.67 versus 3.11), the relative nasal height (1.15–1.62 versus 0.726), and the relative nasal width (2.08–2.33 versus 1.11). From C. molesworthi , we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus dots in chevrons), the ventral scale rows (87–93 versus 102–125), the total strigae on ten scales (64–115 versus 138–159), the relative length of digits on one hindlimb (30.2–31.2 versus 22.4–29.4), the relative eye length (3.79–5.17 versus 3.28–3.70), the relative ear width (1.75–2.08 versus 1.37–1.50), the relative largest supraocular width (2.96–4.03 versus 1.69–2.80), the relative longest finger length (6.43–6.67 versus 4.28–5.19), the relative frontal width (57.6–66.1 versus 75.9–95.5), the relative angled subocular height (1.00–1.07 versus 1.11), the relative angled subocular width (2.77–2.83 versus 2.09–2.48), and the relative nasal width (2.08–2.33 versus 1.55–1.72). From C. occiduus , we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus absent), the longitudinal paramedian lines (present versus absent), the dots arranged in bars in the lateral areas (present versus absent), the adult SVL (60.0–86.1 versus 269–367), the ventral scale rows (87–93 versus 109–134), the midbody scale rows (41–45 versus 46–56), the total lamellae on one hand (39–40 versus 50–66), the total strigae on ten scales (64–115 versus 374), the relative length of digits on one hindlimb (30.2–31.2 versus 24.4–29.7), the relative distance between angled subocular and mouth (0.640 –0.983 versus 1.26–1.27), the relative eye length (3.79–5.17 versus 2.87–3.33), the relative ear width (1.75–2.08 versus 0.948–1.39), the relative longest finger length (6.43–6.67 versus 4.77–5.46), and the relative distance between the ear and eye (7.58–8.02 versus 8.98–10.9). From C. oligolepis sp. nov., we distinguish C. macrotus by the dorsal pattern (chevrons/bands versus dots in chevrons), the dots arranged in bars in the lateral areas (present versus absent), the ventral scale rows (87–93 versus 98), the midbody scale rows (41–45 versus 35), and the total lamellae on one hand (39–40 versus 30). From C. striatus , we distinguish C. macrotus by the dots arranged in bars in the lateral areas (present versus absent), the adult SVL (60.0–86.1 versus 145), the ventral scale rows (87–93 versus 101–109), the total lamellae on one hand (39–40 versus 59–66), and the total strigae on ten scales (64–115 versus 279).

Description of holotype. USNM 286917. An adult; SVL 60.0 mm; tail nearly cylindrical, broken, 8.05 mm (13.4% SVL); axilla-to-groin distance 32.0 mm (53.3% SVL); forelimb length 13.4 mm (22.3% SVL); hindlimb length 16.9 mm (28.2% SVL); head length 12.3 mm (20.5% SVL); head width 8.31 mm (13.9% SVL); head width 67.6% head length; diameter of orbit 3.10 mm (5.17% SVL); horizontal diameter of ear opening 1.25 mm (2.08% SVL); vertical diameter of ear opening 1.28 mm (2.13% SVL); length of all toes on one foot 18.1 mm (30.2% SVL); shortest distance between angled subocular and lip 0.59 mm (0.983% SVL); shortest distance between the ocular and auricular openings 4.94 mm (8.23% SVL); longest finger length 4.00 mm (6.67% SVL); largest supraocular width 2.42 mm (4.03% SVL); cloacal width 4.68 mm (7.80% SVL); prefrontal width 3.33 mm (5.55% SVL); frontal width 57.6% frontal length; nasal height 0.97 mm (1.62% SVL); angled subocular height 0.60 mm (1.00% SVL); shortest distance between the eye and naris 3.36 mm (5.60% SVL); canthal iii width 1.11 mm (1.85% SVL); angled subocular width 1.66 mm (2.77% SVL); nasal width 1.25 mm (2.08% SVL); rostral 1.76X 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 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 much wider than long; parietal separated from supraoculars by 1 st and 2 nd temporals and frontoparietal (left)/1 st temporals and frontoparietal (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, 1 st median ocular, canthal iii, 2 nd loreal, and 3 rd and 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, lower preocular, and 1 st and 2 nd loreals (left)/(right); 9 (left)/10 (right) median oculars, 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); 3 (left)/2 (right) suboculars; posterior subocular large and elongate (left)/(right); anterior subocular small (left)/(right); 10 supralabials (left)/(right), 6 to level below center of eye (left)/(right); 10 infralabials (left)/(right), 6 (left)/6–7(right) to level below center of eye; mental small, followed by a single, larger postmental; 4 pairs of enlarged chin shields; 1 st pair in contact with one another anteriorly, posteriorly separated by one scale; 2 nd –4 th pairs separated by 1–3 scales; 89 transverse rows of dorsal scales from interoccipital to base of tail; 93 transverse rows of ventral scales from mental to vent; 41 scales around midbody; 5 digits; finger lengths 3>4>2>5>1; 10 lamellae under longest finger (left)/(right); 39 total lamellae on one hand; toe lengths 4>3>5>2>1; 17 (left)/15 (right) lamellae under longest toe; striate, angled to give impression of a faint median keel dorsal body and caudal scales; smooth ventral scales; 64 total strigae counted on ten scales.

FIGURE 26. (A–F) Celestus macrotus (USNM 286917, holotype), SVL 60.0 mm.

FIGURE 27. Celestus macrotus (ANSP 38506, SBH 269931), SVL 42.1 mm (juvenile), in life. From southeast of Pic La Selle, Sud-Est Department, Haiti. Photo by SBH.

Color (in alcohol): dorsal surface of head dark brown, patternless; lateral surfaces of head grading from dark brown to gray with darker brown eye masks; dorsal surfaces of the body are dark brown with darker brown longitudinal paramedian lines and irregular spotting; dorsal surface of tail the same as the body; lateral areas grade from dark brown to gray with darker spots in rows; dorsal surfaces of the limbs are dark brown with gray mottling; lateral and ventral areas of the limbs fade to gray with darker brown mottling; ventral surfaces of the head, body, and tail are gray with brown mottling over the entire surface.

Variation. The examined material resembles the pattern of the holotype with darker spots ranging from occurring irregularly down the extent of the dorsum in a heavy pattern (ANSP 38505) to being arranged in chevrons (ANSP 38506). Unlike the holotype, both ANSP 38505 and ANSP 38506 have head scale borders with darker outlines whereas ANSP 38505 also has irregular darker markings on its head. All specimens exhibit longitudinal paramedian lines. Measurements and other morphological data for the holotype and other examined material are presented in Table 1.

Distribution. Celestus macrotus is found in the Massif de la Selle of Haiti and the Sierra de Bahoruco of the Dominican Republic at elevations of 1930–2320 m (Fig. 11).

Ecology and conservation. The holotype was collected under rocks and logs in mature stands of pine forest, Pinus occidentalis , alongside Wetmorena surda ( Thomas & Hedges 1989) . The other Haitian specimens, south of Pic La Selle, were collected in the same habitat, alongside W. surda and Panolopus aporus . In both localities, Celestus macrotus was considerably rarer than other species.

The IUCN Redlist ( IUCN 2023) considers the conservation status of Celestus macrotus to be Endangered B1ab(i,iii) “due to its limited distribution (with an extent of occurrence of about 128 km 2), occurrence in a single location, and ongoing threats from agriculture expansion, wildfires due to anthropogenic causes and wood extraction.” Studies are needed to determine the health of remaining populations and threats to the survival of the species.

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

Etymology. From the Greek, macro, long (large in common usage), and otos, ear, in reference to the large external auditory meatus.

Remarks. Thomas &Hedges (1989) discussed the scales surrounding the eyes of diploglossid lizards, particularly as that applies to Celestus macrotus . However, this scale terminology was not used in later works ( Savage & Lips 1993; Savage et al. 2008). Thomas & Hedges (1989) also noted that, with preliminary protein electrophoretic data, Celestus macrotus was considerably divergent, falling outside of a cluster containing Celestus barbouri , Panolopus costatus , Celestus crusculus , Panolopus curtissi , Caribicus darlingtoni , and Comptus stenurus .

Celestus macrotus is included in our genetic dataset and has significant support in both Bayesian and ML likelihood analyses at the crown node of the species and the stem node that places it as the closest relative of a group containing all other species of Celestus . Based on our timetree (Fig. 4), C. macrotus diverged from its closest relative 5.58 Ma, consistent with typical species of vertebrates (> 0.7 Ma; Hedges et al. 2015). Celestus macrotus was recognized as a distinct species in our ASAP analysis.