Celestus macrolepis Gray, 1845:118

Schools, Molly & Hedges, Blair, 2024, A new forest lizard fauna from Caribbean islands (Squamata, Diploglossidae, Celestinae), Zootaxa 5554 (1), pp. 1-306 : 73-78

Celestus macrolepis Gray, 1845:118 . Holotype: BMNH 1946.8 .3.82 (locality unknown).

Celestus occiduus — Boulenger, 1885:290 (part).

Celestus occiduus — Schwartz & Henderson, 1988:99 (part).

Celestus macrolepis — Schools & Hedges, 2021:220.

Celestus macrolepis — Landestoy et al., 2022:204.

Material examined (n=2). JAMAICA. Localities unknown. BMNH 1946.8.3.82; BMNH 1961.1851.

Diagnosis. Celestus macrolepis has (1) a dorsal pattern bicolored (dark anteriorly, pale posteriorly), (2) head markings absent, (3) markings in the longitudinal paramedian area absent, (4) dots arranged in bars in the lateral band absent, (5) an adult SVL of 254–316 mm, (6) ventral scale rows, 112–116, (7) midbody scale rows, 46–48, (8) total lamellae on one hand, 52–54, (9) total strigae on ten scales, 398, (10) relative length of all digits on one hindlimb, 27.5–28.0 %, (11) relative distance between the angled subocular and mouth, 1.39–1.66 %, (12) relative eye length, 3.63–3.70 %, (13) relative forelimb length, 26.1–26.7 %, (14) relative ear width, 0.760–1.43 %, (15) relative rostral height, 1.53–1.75 %, (16) relative head length, 19.2–22.9 %, (17) relative mental width, 1.87 %, (18) relative postmental width, 3.81 %, (19) relative cloacal width, 11.2 %, (20) relative prefrontal width, 3.94 %, (21) relative largest supraocular width, 2.64–3.01 %, (22) relative longest finger length, 5.47–5.51 %, (23) relative distance between the ear and eye, 8.02–10.9 %, (24) relative head width, 80.5 %, (25) relative frontal width, 78.4 %, (26) relative nasal height, 1.18 %, (27) relative angled subocular height, 1.17 %, (28) relative distance between the eye and naris, 6.02 %, (29) relative canthal iii length, 1.99 %, (30) relative angled subocular width, 2.57 %, and (31) relative nasal length, 1.75 %. The species stem time is 4.64 Ma and the species crown time is unavailable (Fig. 4).

Celestus macrolepis differs from all other species of the genus in having a bicolored dorsal pattern (dark anteriorly, pale posteriorly). This species also has a smaller relative prefrontal width (3.94) than most other species of the genus. Celestus macrolepis has a larger SVL (254–316), total strigae on ten scales count (398), relative distance between the angled subocular and mouth (1.39–1.66), relative forelimb length (26.1–26.7), relative postmental width (3.81), relative cloacal width (11.2), and relative distance between the eye and naris (6.02) than most other species of the genus.

From Celestus barbouri , we distinguish C. macrolepis by the dorsal pattern (bicolored versus chevrons), the adult SVL (254–316 versus 78.4–93.6), the ventral scale rows (112–116 versus 118–151), the total lamellae on one hand (52–54 versus 36–49), the total strigae on ten scales (398 versus 105–136), the relative length of digits on one hindlimb (27.5–28.0 versus 18.2–23.5), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.437 –0.556), the relative forelimb length (26.1–26.7 versus 15.4–19.0), the relative head length (19.2–22.9 versus 14.6–16.6), the relative mental width (1.87 versus 1.51–1.85), the relative postmental width (3.81 versus 2.51–3.29), the relative cloacal width (11.2 versus 7.64–8.26), the relative prefrontal width (3.94 versus 3.97–4.33), the relative longest finger length (5.47–5.51 versus 2.92–3.81), the relative distance between the ear and eye (8.02–10.9 versus 6.23–7.15), the relative nasal height (1.18 versus 0.930–1.12), the relative angled subocular height (1.17 versus 0.553–1.16), the relative distance between the eye and naris (6.02 versus 4.68–4.83), the relative width of canthal iii (1.99 versus 1.54–1.93), the relative angled subocular width (2.57 versus 1.97–2.52), and the relative nasal width (1.75 versus 1.38–1.65). From C. capitulatus sp. nov., we distinguish C. macrolepis by the dorsal pattern (bicolored versus irregular dots/dots in chevrons), the adult SVL (254–316 versus 62.1–81.8), the total lamellae on one hand (52–54 versus 25–38), the total strigae on ten scales (398 versus 105–192), the relative length of digits on one hindlimb (27.5–28.0 versus 17.6–22.3), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.525–1.17), the relative forelimb length (26.1–26.7 versus 14.3–18.1), the relative head length (19.2–22.9 versus 15.1–17.7), the relative mental width (1.87 versus 1.28–1.84), the relative postmental width (3.81 versus 2.62–2.97), the relative cloacal width (11.2 versus 7.84–8.67), the relative prefrontal width (3.94 versus 4.30–4.72), the relative largest supraocular width (2.64–3.01 versus 2.03–2.61), the relative longest finger length (5.47–5.51 versus 3.45– 3.75), the relative distance between the ear and eye (8.02–10.9 versus 6.45–7.84), the relative head width (80.5 versus 71.6–78.6), the relative angled subocular height (1.17 versus 0.586–1.01), the relative distance between the eye and naris (6.02 versus 4.57–5.03), and the relative angled subocular width (2.57 versus 1.93–2.32). From C. crusculus , we distinguish C. macrolepis by the dorsal pattern (bicolored versus absent/flecks in series/dots in chevrons), the longitudinal paramedian lines (absent versus present), the adult SVL (254–316 versus 59.6–77.6), the midbody scale rows (46–48 versus 37–44), the total lamellae on one hand (52–54 versus 30–39), the total strigae on ten scales (398 versus 106–194), the relative length of digits on one hindlimb (27.5–28.0 versus 18.7–24.7), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.339 –0.884), the relative eye length (3.63–3.70 versus 2.93–3.61), the relative forelimb length (26.1–26.7 versus 12.8–20.7), the relative postmental width (3.81 versus 2.73–3.37), the relative cloacal width (11.2 versus 6.89–8.77), the relative longest finger length (5.47–5.51 versus 2.94–4.10), the relative frontal width (78.4 versus 82.6–91.1), the relative distance between the eye and naris (6.02 versus 4.31–4.86), the relative angled subocular width (2.57 versus 2.03–2.43), and the relative nasal width (1.75 versus 1.27–1.60). From C. duquesneyi , we distinguish C. macrolepis by the dorsal pattern (bicolored versus bands), the adult SVL (254–316 versus 62.1), the total strigae on ten scales (398 versus 130), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.644), and the relative ear width (0.760– 1.43 versus 2.45). From C. hesperius sp. nov., we distinguish C. macrolepis by the dorsal pattern (bicolored versus dots in chevrons), the adult SVL (254–316 versus 54.0–62.3), the midbody scale rows (46–48 versus 39–44), the total lamellae on one hand (52–54 versus 29–34), the total strigae on ten scales (398 versus 95–122), the relative length of digits on one hindlimb (27.5–28.0 versus 21.7–26.2), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.594 –0.648), the relative forelimb length (26.1–26.7 versus 18.6–21.3), the relative ear width (0.760–1.43 versus 1.52–1.59), the relative head length (19.2–22.9 versus 15.7–17.7), the relative largest supraocular width (2.64–3.01 versus 1.91–2.22), the relative longest finger length (5.47–5.51 versus 3.50–4.04), and the relative distance between the ear and eye (8.02–10.9 versus 6.74–7.53). From C. hewardi , we distinguish C. macrolepis by the dorsal pattern (bicolored versus mottled/bands), the adult SVL (254–316 versus 129–171), the total strigae on ten scales (398 versus 164–315), the relative forelimb length (26.1–26.7 versus 22.2–24.6), the relative mental width (1.87 versus 1.75–1.81), the relative postmental width (3.81 versus 2.84–3.44), the relative cloacal width (11.2 versus 8.81–9.89), the relative prefrontal width (3.94 versus 4.18–4.80), the relative head width (80.5 versus 68.4–77.1), the relative frontal width (78.4 versus 57.3–75.3), the relative nasal height (1.18 versus 1.21–1.24), the relative distance between the eye and naris (6.02 versus 5.00–5.60), and the relative angled subocular width (2.57 versus 1.63–2.23). From C. jamesbondi sp. nov., we distinguish C. macrolepis by the dorsal pattern (bicolored versus absent/irregular dots/dots in chevrons), the adult SVL (254–316 versus 54.7–72.0), the midbody scale rows (46–48 versus 35–44), the total lamellae on one hand (52–54 versus 30–36), the total strigae on ten scales (398 versus 101–173), the relative length of digits on one hindlimb (27.5–28.0 versus 19.8–26.3), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.363–1.01), the relative forelimb length (26.1– 26.7 versus 14.4–19.9), the relative postmental width (3.81 versus 2.61–2.92), the relative cloacal width (11.2 versus 6.59–9.08), the relative prefrontal width (3.94 versus 4.29–5.09), the relative longest finger length (5.47–5.51 versus 3.66–4.33), the relative distance between the ear and eye (8.02–10.9 versus 6.92–7.80), the relative frontal width (78.4 versus 70.5–77.6), and the relative distance between the eye and naris (6.02 versus 4.25–5.54). From C. macrotus , we distinguish C. macrolepis by the dorsal pattern (bicolored versus chevrons/bands), the longitudinal paramedian lines (absent versus present), the dots arranged in bars in the lateral areas (absent versus present), the adult SVL (254–316 versus 60.0–86.1), the ventral scale rows (112–116 versus 87–93), the midbody scale rows (46–48 versus 41–45), the total lamellae on one hand (52–54 versus 39–40), the total strigae on ten scales (398 versus 64–115), the relative length of digits on one hindlimb (27.5–28.0 versus 30.2–31.2), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.640 –0.983), the relative eye length (3.63–3.70 versus 3.79–5.17), the relative forelimb length (26.1–26.7 versus 22.4–25.0), the relative ear width (0.760–1.43 versus 1.75–2.08), and the relative longest finger length (5.47–5.51 versus 6.43–6.67). From C. microblepharis , we distinguish C. macrolepis by the dorsal pattern (bicolored versus chevrons), the adult SVL (254–316 versus 96.4), the total lamellae on one hand (52–54 versus 30), the total strigae on ten scales (398 versus 165), the relative length of digits on one hindlimb (27.5–28.0 versus 16.6), the relative distance between angled subocular and mouth (1.39– 1.66 versus 0.820), the relative eye length (3.63–3.70 versus 1.83), the relative forelimb length (26.1–26.7 versus 14.2), the relative ear width (0.760–1.43 versus 0.446), the relative longest finger length (5.47–5.51 versus 3.11), and the relative nasal height (1.18 versus 0.726). From C. molesworthi , we distinguish C. macrolepis by the dorsal pattern (bicolored versus dots in chevrons), the adult SVL (254–316 versus 78.1–103), the total lamellae on one hand (52–54 versus 32–44), the total strigae on ten scales (398 versus 138–159), the relative distance between angled subocular and mouth (1.39–1.66 versus 0.653 –0.845), the relative forelimb length (26.1–26.7 versus 17.5– 24.2), and the relative longest finger length (5.47–5.51 versus 4.28–5.19). From C. occiduus , we distinguish C. macrolepis by the dorsal pattern (bicolored versus absent), the relative distance between angled subocular and mouth (1.39–1.66 versus 1.26–1.27), the relative eye length (3.63–3.70 versus 2.87–3.33), the relative forelimb length (26.1–26.7 versus 23.5–23.9), and the relative longest finger length (5.47–5.51 vs 4.77–5.46). From C. oligolepis sp. nov., we distinguish C. macrolepis by the dorsal pattern (bicolored versus dots in chevrons), the head markings (absent versus present), the longitudinal paramedian lines (absent versus present), the ventral scale rows (112–116 versus 98), the midbody scale rows (46–48 versus 35), and the total lamellae on one hand (52–54 versus 30). From C. striatus , we distinguish C. macrolepis by the dorsal pattern (bicolored versus absent/chevrons), the adult SVL (254–316 versus 145), the ventral scale rows (112–116 versus 101–109), the midbody scale rows (46–48 versus 41–43), the total lamellae on one hand (52–54 versus 59–66), and the relative distance between angled subocular and mouth (1.39–1.66 versus 0.710).

Description of holotype. BMNH 1946.8.3.82. An adult; SVL 254 mm; tail laterally compressed, 150 mm (59.1% SVL); axilla-to-groin distance 141 mm (55.5% SVL); forelimb length 67.9 mm (26.7% SVL); hindlimb length 88.2 mm (34.7% SVL); head length 48.6 mm (19.1% SVL); head width 39.1 mm (15.4% SVL); head width 80.5% head length; diameter of orbit 9.41 mm (3.70% SVL); horizontal diameter of ear opening 3.64 mm (1.43% SVL); vertical diameter of ear opening 6.46 mm (2.54% SVL); length of all toes on one foot 71.0 mm (28.0% SVL); shortest distance between angled subocular and lip 3.52 mm (1.39% SVL); shortest distance between the ocular and auricular openings 20.4 mm (8.03% SVL); longest finger length 13.9 mm (5.47% SVL); largest supraocular width 6.70 mm (2.64% SVL); cloacal width 28.4 mm (11.2% SVL); mental width 4.75 mm (1.87% SVL); postmental width 9.68 mm (3.81% SVL); prefrontal width 10.0 mm (3.94% SVL); frontal width 78.4% frontal length; nasal height 3.00 mm (1.18% SVL); angled subocular height 2.98 mm (1.17% SVL); shortest distance between the eye and naris 15.3 mm (6.02% SVL); canthal iii width 5.05 mm (1.99% SVL); angled subocular width 6.53 mm (2.57% SVL); nasal width 4.44 mm (1.75% SVL); rostral 1.53X 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 fused to them, posteriorly touching the fused to interoccipital, which is wider than long; parietal separated from supraoculars by 1 st temporal and frontoparietal (left)/1 st and 2 nd temporals and frontoparietal (right); nasal single; nostril above suture between 1 st and 2 nd supralabials (left)/(right); 1 postnasal (left)/(right); 3 loreals (left)/(right); 1 st loreal small, higher than wide (left)/(right); 2 nd loreal large, irregular, fused with 3 rd loreal at the very bottom (left), higher than long, in contact with posterior internasals, prefrontal/frontonasal complex, anterior most ocular, canthal iii, 1 st loreal, 3 rd loreal, and supralabials 3–4 (left)/large, irregular, higher than wide, in contact with posterior internasals, prefrontal/frontonasal complex, median ocular 1, canthal iii, subcanthal ( Savage and Lips 2008), 1 st and 3 rd loreals, and supralabials 3–5 (right); 3 rd loreal irregular, excluded from contact with supraocular by canthal iii (left)/rectangular, excluded from contact with canthal iii by subcanthal (right); final loreal posteriorly bordering the upper and lower preoculars (left)/(right); canthal iii wider than high (left)/(right), contacting 1 st median ocular, anterior supraciliary, upper preocular, and 2 nd and 3 rd loreals (left)/1 st median ocular, anterior supraciliary, upper preocular, subcanthal, and the loreals 2–3 (right); 11 (left)/10 (right) median oculars, 1 st and 2 nd contacting the prefrontal (left)/(right); 2 upper preoculars (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); 10 supralabials (left)/(right), 7 to level below center of eye (left)/(right); 9 (left)/10 (right) infralabials, 7 to level below center of eye (left)/(right); mental small, followed by a single, slightly 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 1–5 scales; 109 transverse rows of dorsal scales from interoccipital to base of tail; 116 transverse rows of ventral scales from mental to vent; 48 scales around midbody; 5 digits; finger lengths 3>4>2>5>1; 13 (3 are fused) (left)/13 (2 are fused) (right) lamellae under longest finger; 54 total lamellae on one hand; toe lengths 4>3>5>2>1; 24 lamellae under longest toe (left)/(right); dorsal body and caudal scales striate, some with a faint median keel; smooth ventral scales; 398 total strigae counted on ten scales.

Color (in alcohol): dorsal surface of head very dark brown, patternless; lateral surfaces of head grading from dark brown to medium brown, patternless; dorsal surfaces of the body are dark brown, interspersed with large, gray-tan patches that have several dark brown scales in them; dorsal surface of tail the same as the body, interspersed with large, gray-tan patches that have several dark brown scales in them; lateral areas have the same coloration and pattern as the dorsal areas; dorsal surfaces of the limbs are dark brown with gray-tan patches; lateral and ventral areas of the limbs same as the body; ventral surfaces of the head, body, and tail are the same as the dorsal body scales with slightly reduced colors.

Variation. The other examined specimen closely resembles the holotype in pattern and scalation. Measurements and other morphological data for the holotype and other examined material are presented in Table 1.

Distribution. No information is known on the distribution of Celestus macrolepis as the most specific location recorded for any of the specimens is “ Jamaica ” (BMNH 1961.1851).

Ecology and conservation. Based on literature reports, Celestus macrolepis is a swamp-dwelling lizard. Although confusion has persisted throughout the literature regarding the identification of C. macrolepis , ecological reports of a dark brown diploglossid (presumably C. macrolepis ) living in the Jamaican swamps confirm its placement in the Swamp Ecomorph. Browne (1789) described C. occiduus as normally being a dirty brown color, but often changing to a fine golden color. Diploglossids are not known for such dramatic color change, and he more likely was referring to both the dark brown C. macrolepis and the yellow C. occiduus . Sloane (1725) described a giant Jamaican diploglossid that lived “both in water and on land,” He also noted that upon dissection, the specimen had crabs in its stomach. Based on the brown color of the specimen that he dissected, Sloane was probably referring to a member of the species C. macrolepis , not C. occiduus . As reported in Ahrenfeldt (1954), Duméril & Bibron (1839) had also doubted that the species described by Sloane was C. occiduus . Celestus macrolepis also has a strongly laterally compressed tail, a common adaptation for swimming in vertebrates ( Gray 1845). A laterally compressed tail was also reported for C. occiduus by Boulenger (1885); however, his description included the holotype of C. macrolepis , in addition to other specimens that are identified as C. occiduus .

The IUCN Redlist ( IUCN 2023) considers the conservation status of Celestus macrolepis to be Critically Endangered (Possibly Extinct) C2a(i,ii); D. It faces a primary threat from habitat destruction from agriculture and urbanization. Secondary threats include predation from introduced mammals, including the mongoose and black rats. Studies are needed to determine the health and extent of remaining populations and threats to the survival of the species. Captive-breeding programs should be undertaken, because eradication of introduced mammalian predators is not possible on Jamaica.

FIGURE 25. (A–F) Celestus macrolepis (BMNH 1946.8.3.82, holotype), SVL 254 mm.

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

Etymology. The species name is derived from the prefix macro- (large) and lepis (scale). Gray (1845), who described the species, named it the “Large-scaled Galliwasp.”

Remarks. The species Celestus macrolepis was recently recognized as a valid species ( Schools & Hedges 2021) long after it had been synonymized with C. occiduus ( Boulenger 1885) . Although both species share a laterally compressed tail ( Schools & Hedges 2022), several key differences exist between C. macrolepis and C. occiduus . In addition to its darker coloration, C. macrolepis also has scale differences and a more robust head than C. occiduus ( Schools & Hedges 2021) . We recognize BMNH 1961.1851 as a member of C. macrolepis largely because it shares the distinct bicolored pattern as the holotype (Fig. 25).

Celestus macrolepis is included in our genetic dataset and is placed outside the group containing C. duquesneyi , C. hesperius sp. nov., C. hewardi , C. jamesbondi sp. nov., C. molesworthi , C. occiduus , and C. striatus with significant support in our ML analysis and a support value of 80% in out Bayesian analysis. Based on our timetree (Fig. 4), C. macrolepis diverged from its closest relative 4.64 Ma, consistent with typical species of vertebrates (> 0.7 Ma; Hedges et al. 2015). Celestus macrolepis sp. nov. was recognized as a distinct species in our ASAP analysis.