Celestus hewardi ( Gray 1845 )

Schools, Molly & Hedges, Blair, 2024, A new forest lizard fauna from Caribbean islands (Squamata, Diploglossidae, Celestinae), Zootaxa 5554 (1), pp. 1-306 : 63-68

Celestus hewardi ( Gray 1845)

Red-spotted Forest Lizard

(Fig. 22)

Celestus hewardii — Gray, 1845:118. Replacement name for Tiliqua striata Gray (1839:293) , a junior homonym of Celestus striatus Gray (1839:288) . Holotype: BMNH 1946.8.7.99.

Diploglossus (Celestus) variegatus — Peters, 1874:370. Holotype: ZMB 8029 , “unbestimmter Herkunft, gekauft” (of unknown origin, bought).

Celestus impressus — Cope, 1868:127. Lectotype: ANSP 9225 .

Diploglossus hewardii — Boulenger, 1885:291.

Celestus occiduus hewardii —Grant, 1940:106.

Celestus hewardi — Schwartz & Henderson, 1991:374.

Celestus hewardii — Hedges et al., 2019:17.

Celestus hewardii — Schools & Hedges, 2021:220.

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

Material examined (n=23). JAMAICA. Manchester. BMNH 1938.4 .13.6, C. Grant, Mandeville ; BMNH 1946.8 .7.99 (holotype, collected by R. Heward in Jamaica with no precise locality) ; BMNH 1845.12.27.6–8 (one of the series, specimen number unmarked); USNM 108234 , 108238 , Mandeville, 5 May 1939 ; MCZ R-45173, R-45177, Chapman Grant, Mandeville, 11 May 1937 ; MCZ R-127906–7, Thomas A. Jenssen, Mandeville, 16 June 1970 ; USNM 102651 , Kensworth, February 1937 ; USNM 108231 , Mandeville, 13 April 1937 . Saint Ann. USNM 251919 , 1.5 mi NE of Orange Valley , 4 September 1976 . Saint James. USNM 108329 , 108330–5 , 5 mi W of Montego Bay , 25 March 1938 . Trelawny. KU 226546 , Duncans, 13 July 1961 ; USNM 251918 , 1.1 mi NW of Windsor , 4 July 1971 . Westmoreland. KU 226549–50 , 6.9 mi SSE Darliston , 11 July 1967 .

Diagnosis. Celestus hewardi has (1) a dorsal pattern of mottled/bands, (2) head markings absent, (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 129–171 mm, (6) ventral scale rows, 113–137, (7) midbody scale rows, 43–59, (8) total lamellae on one hand, 50–61, (9) total strigae on ten scales, 164–315, (10) relative length of all digits on one hindlimb, 24.1– 30.6 %, (11) relative distance between the angled subocular and mouth, 0.744–1.40 %, (12) relative eye length, 2.98–4.05 %, (13) relative forelimb length, 22.2–24.6 %, (14) relative ear width, 1.40–1.82 %, (15) relative rostral height, 1.50–1.76 %, (16) relative head length, 16.8–21.5 %, (17) relative mental width, 1.75–1.81 %, (18) relative postmental width, 2.84–3.44 %, (19) relative cloacal width, 8.81–9.89 %, (20) relative prefrontal width, 4.18–4.80 %, (21) relative largest supraocular width, 2.43–2.96 %, (22) relative longest finger length, 5.03–5.66 %, (23) relative distance between the ear and eye, 6.72–8.73 %, (24) relative head width, 68.4–77.1 %, (25) relative frontal width, 57.3–75.3 %, (26) relative nasal height, 1.21–1.24 %, (27) relative angled subocular height, 0.918–1.30 %, (28) relative distance between the eye and naris, 5.00–5.60 %, (29) relative canthal iii length, 1.70–2.12 %, (30) relative angled subocular width, 1.63–2.23 %, and (31) relative nasal length, 1.56–1.88 %. The species stem time is 3.59 Ma and the species crown time is unavailable (Fig. 4).

Celestus hewardi differs from most (except two) species in the genus Celestus in having a dorsal pattern of mottled/bands. From Celestus barbouri , we distinguish C. hewardi by the dorsal pattern (mottled/bands versus chevrons), the adult SVL (129–171 versus 78.4–93.6), the total lamellae on one hand (50–61 versus 36–49), the total strigae on ten scales (164–315 versus 105–136), the relative length of digits on one hindlimb (24.1–30.6 versus 18.2–23.5), the relative distance between angled subocular and mouth (0.744–1.40 versus 0.437 –0.556), the relative forelimb length (22.2–24.6 versus 15.4–19.0), the relative head length (16.8–21.5 versus 14.6–16.6), the relative cloacal width (8.81–9.89 versus 7.64–8.26), the relative longest finger length (5.03–5.66 versus 2.92–3.81), the relative nasal height (1.21–1.24 versus 0.930–1.12), and the relative distance between the eye and naris (5.00–5.60 versus 4.68–4.83). From C. capitulatus sp. nov., we distinguish C. hewardi by the dorsal pattern (mottled/bands versus irregular dots/dots in chevrons), the adult SVL (129–171 versus 62.1–81.8), the total lamellae on one hand (50–61 versus 25–38), the relative length of digits on one hindlimb (24.1–30.6 versus 17.6–22.3), the relative forelimb length (22.2–24.6 versus 14.3–18.1), the relative cloacal width (8.81–9.89 versus 7.84–8.67), the relative longest finger length (5.03–5.66 versus 3.45–3.75), and the relative frontal width (57.3–75.3 versus 78.1–81.6). From C. crusculus , we distinguish C. hewardi by the dorsal pattern (mottled/bands versus absent/flecks in series/ dots in chevrons), the adult SVL (129–171 versus 59.6–77.6), the total lamellae on one hand (50–61 versus 30–39), the relative forelimb length (22.2–24.6 versus 12.8–20.7), the relative cloacal width (8.81–9.89 versus 6.89–8.77), the relative longest finger length (5.03–5.66 versus 2.94–4.10), the relative frontal width (57.3–75.3 versus 82.6– 91.1), and the relative distance between the eye and naris (5.00–5.60 versus 4.31–4.86). From C. duquesneyi , we distinguish C. hewardi by the adult SVL (129–171 versus 62.1), the total lamellae on one hand (50–61 versus 64), the total strigae on ten scales (164–315 versus 130), the relative length of digits on one hindlimb (24.1–30.6 versus 31.4), the relative distance between angled subocular and mouth (0.744–1.40 versus 0.644), the relative eye length (2.98–4.05 versus 4.36), the relative ear width (1.40–1.82 versus 2.45), the relative rostral height (1.50–1.76 versus 2.14), the relative mental width (1.75–1.81 versus 2.35), the relative cloacal width (8.81–9.89 versus 9.98), the relative prefrontal width (4.18–4.80 versus 5.41), the relative longest finger length (5.03–5.66 versus 6.52), the relative head width (68.4–77.1 versus 64.6), the relative angled subocular height (0.918–1.30 versus 1.61), the relative width of canthal iii (1.70–2.12 versus 1.59), the relative angled subocular width (1.63–2.23 versus 2.90), and the relative nasal width (1.56–1.88 versus 2.01). From C. hesperius sp. nov., we distinguish C. hewardi by the dorsal pattern (mottled/bands versus dots in chevrons), the adult SVL (129–171 versus 54.0–62.3), the total lamellae on one hand (50–61 versus 29–34), the total strigae on ten scales (164–315 versus 95–122), the relative distance between angled subocular and mouth (0.744–1.40 versus 0.594 –0.648), the relative forelimb length (22.2–24.6 versus 18.6–21.3), the relative cloacal width (8.81–9.89 versus 7.99–8.55), the relative largest supraocular width (2.43–2.96 versus 1.91–2.22), the relative longest finger length (5.03–5.66 versus 3.50–4.04), and the relative frontal width (57.3–75.3 versus 80.5–86.1). From C. jamesbondi sp. nov., we distinguish C. hewardi by the dorsal pattern (mottled/bands versus absent/irregular dots/dots in chevrons), the adult SVL (129–171 versus 54.7–72.0), the ventral scale rows (113–137 versus 91–112), the total lamellae on one hand (50–61 versus 30–36), the relative forelimb length (22.2–24.6 versus 14.4–19.9), and the relative longest finger length (5.03–5.66 versus 3.66–4.33). From C. macrolepis , we distinguish C. hewardi by the dorsal pattern (mottled/bands versus bicolored), the adult SVL (129– 171 versus 254–316), the total strigae on ten scales (164–315 versus 398), the relative forelimb length (22.2–24.6 versus 26.1–26.7), the relative mental width (1.75–1.81 versus 1.87), the relative postmental width (2.84–3.44 versus 3.81), the relative cloacal width (8.81–9.89 versus 11.2), the relative prefrontal width (4.18–4.80 versus 3.94), the relative head width (68.4–77.1 versus 80.5), the relative frontal width (57.3–75.3 versus 78.4), the relative nasal height (1.21–1.24 versus 1.18), the relative distance between the eye and naris (5.00–5.60 versus 6.02), and the relative angled subocular width (1.63–2.23 versus 2.57). From C. macrotus , we distinguish C. hewardi by the adult SVL (129–171 versus 60.0–86.1), the ventral scale rows (113–137 versus 87–93), the total lamellae on one hand (50–61 versus 39–40), the total strigae on ten scales (164–315 versus 64–115), the relative prefrontal width (4.18–4.80 versus 4.87–5.55), the relative longest finger length (5.03–5.66 versus 6.43–6.67), the relative angled subocular width (1.63–2.23 versus 2.77–2.83), and the relative nasal width (1.56–1.88 versus 2.08–2.33). From C. microblepharis , we distinguish C. hewardi by the dorsal pattern (mottled/bands versus chevrons), the adult SVL (129–171 versus 96.4), the ventral scale rows (113–137 versus 109), the total lamellae on one hand (50–61 versus 30), the relative length of digits on one hindlimb (24.1–30.6 versus 16.6), the relative eye length (2.98–4.05 versus 1.83), the relative forelimb length (22.2–24.6 versus 14.2), the relative ear width (1.40–1.82 versus 0.446), the relative head length (16.8–21.5 versus 14.7), the relative mental width (1.75–1.81 versus 1.44), the relative postmental width (2.84–3.44 versus 2.47), the relative cloacal width (8.81–9.89 versus 8.02), the relative largest supraocular width (2.43–2.96 versus 2.06), the relative longest finger length (5.03–5.66 versus 3.11), the relative nasal height (1.21–1.24 versus 0.726), the relative angled subocular height (0.918–1.30 versus 0.778), the relative distance between the eye and naris (5.00–5.60 versus 4.79), the relative angled subocular width (1.63–2.23 versus 2.90), and the relative nasal width (1.56–1.88 versus 1.11). From C. molesworthi , we distinguish C. hewardi by the dorsal pattern (mottled/bands versus dots in chevrons), the adult SVL (129–171 versus 78.1–103), the total lamellae on one hand (50–61 versus 32–44), the total strigae on ten scales (164–315 versus 138–159), and the relative frontal width (57.3–75.3 versus 75.9–95.5). From C. occiduus , we distinguish C. hewardi by the dorsal pattern (mottled/ bands versus absent), the adult SVL (129–171 versus 269–367), the total strigae on ten scales (164–315 versus 374), the relative ear width (1.40–1.82 versus 0.948–1.39), the relative mental width (1.75–1.81 versus 1.86), the relative postmental width (2.84–3.44 versus 3.57), the relative distance between the ear and eye (6.72–8.73 versus 8.98– 10.9), the relative nasal height (1.21–1.24 versus 1.16), the relative distance between the eye and naris (5.00–5.60 versus 6.51), and the relative angled subocular width (1.63–2.23 versus 2.52). From C. oligolepis sp. nov., we distinguish C. hewardi by the dorsal pattern (mottled/bands versus dots in chevrons), the head markings (absent versus present), the ventral scale rows (113–137 versus 98), the midbody scale rows (43–59 versus 35), and the total lamellae on one hand (50–61 versus 30). From C. striatus , we distinguish C. hewardi by the dorsal pattern (mottled/ bands versus absent/chevrons), the ventral scale rows (113–137 versus 101–109), the relative length of digits on one hindlimb (24.1–30.6 versus 37.8), the relative distance between angled subocular and mouth (0.744–1.40 versus 0.710), the relative forelimb length (22.2–24.6 versus 26.1), the relative ear width (1.40–1.82 versus 1.30), the relative rostral height (1.50–1.76 versus 1.94), the relative cloacal width (8.81–9.89 versus 7.93), the relative prefrontal width (4.18–4.80 versus 5.68), the relative longest finger length (5.03–5.66 versus 7.48), the relative distance between the ear and eye (6.72–8.73 versus 9.00), the relative head width (68.4–77.1 versus 82.1), the relative frontal width (57.3–75.3 versus 76.5), the relative nasal height (1.21–1.24 versus 1.08), the relative distance between the eye and naris (5.00–5.60 versus 6.16), and the relative angled subocular width (1.63–2.23 versus 2.29).

Description of syntype. BMNH 1946.8.7.99.A juvenile; SVL 117 mm; tail nearly cylindrical, broken, 78.0 mm (66.7% SVL); axilla-to-groin distance 65.5 mm (56.0% SVL); forelimb length 27.5 mm (23.5% SVL); hindlimb length 41.4 mm (35.4% SVL); head length 21.2 mm (18.1% SVL); head width 15.5 mm (13.3% SVL); head width 73.4% head length; diameter of orbit 4.66 mm (3.98% SVL); horizontal diameter of ear opening 2.72 mm (2.32% SVL); vertical diameter of ear opening 3.13 mm (2.68% SVL); length of all toes on one foot 35.8 mm (30.6% SVL); shortest distance between angled subocular and lip 0.99 mm (0.846% SVL); shortest distance between the ocular and auricular openings 8.04 mm (6.87% SVL); longest finger length 6.83 mm (5.84% SVL); largest supraocular width 3.41 mm (2.91% SVL); cloacal width 4.98 mm (4.26% SVL); mental width 0.91 mm (0.778% SVL); prefrontal width 5.65 mm (4.83% SVL); frontal width 70.1% frontal length; nasal height 1.51 mm (1.29% SVL); angled subocular height 1.47 mm (1.26% SVL); shortest distance between the eye and naris 5.99 mm (5.12% SVL); canthal iii width 2.40 mm (2.05% SVL); angled subocular width 2.12 mm (1.81% SVL); nasal width 1.48 mm (1.26% SVL); rostral 1.66X as wide as high, missing, 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; two frontonasals (absent divided prefrontal), concave posterior margin, 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 and frontoparietal (left)/1 st temporal 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, frontonasal, canthal iii, 2 nd loreal, and 3 rd –4 th supralabials (left)/postnasal, posterior internasal, frontonasal, canthal iii, 2 nd loreal, and 4 th –5 th supralabials (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); 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, 1 st and 2 nd loreals, and the frontonasal (left)/(right); 11 median oculars (left)/(right), 1 st contacting the prefrontal (left)/(right); 2 upper preoculars (left)/(right); an irregular anterior supraciliary (left)/(right); 6 lateral oculars (left)/(right); 6 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 (left)/7 (right) to level below center of eye; 10 (left)/11 (right) infralabials, 7 (left)/8 (right) to level below center of eye; mental small, followed by a single, larger postmental; 5 pairs of enlarged chin shields, followed by 1 pair of reduced chin shields; 1 st pair in contact with one another, 2 nd pair in contact with one another anteriorly, posteriorly separated by one scale; 3 rd –6 th pairs separated by 1–6 scales; 124 transverse rows of dorsal scales from interoccipital to base of tail; 131 transverse rows of ventral scales from mental to vent; 50 scales around midbody; 5 digits; finger lengths 3>4>2>5>1; 14 (left)/15 (right) lamellae under longest finger; 60 total lamellae on one hand; toe lengths 4>3>5>2>1; 20 (left)/(right) lamellae under longest toe; dorsal body and caudal scales striate with a median keel; smooth ventral scales; 162 total strigae counted on ten scales.

Color (in alcohol): dorsal surface of head dark tan to brown and patternless; lateral surfaces of head grading from dark tan-brown to gray; dorsal surfaces of the body are dark tan-brown with darker brown markings arranged in chevrons across the body; dorsal surface of tail paler than the body with the same pattern; lateral areas grade from dark brown to gray-tan with continuations of the dorsal pattern in the lateral band that appear mottled; dorsal surfaces of the limbs are dark brown with paler mottling and patches; lateral and ventral areas of the limbs fade to gray-tan with mottling on the forelimbs; ventral surfaces of the head, body, and tail are anteriorly gray with paler mottling, tan posteriorly.

Variation. The examined material resembles the syntype closely in pattern with most specimens exhibiting the same alternating dark and pale bands extending down the body. In some cases, these bands appear mottled ( MCZ R-45173) or more closely resemble chevrons ( MCZ R-127907). In MCZ R-45177, the entirety of the dorsum appears more mottled than banded. Markings in the longitudinal paramedian area range from absent to mottled to lineate, whereas dots in the lateral bands range from more closely resembling mottling (in KU 226549 and KU 226550 ) to well defined. Measurements and other morphological data for the syntype and other examined material are presented in Table 1 .

Distribution. Celestus hewardi is widespread from central to northern Jamaica at elevations of 0–1000 m (Fig. 12).

Ecology and conservation. Little is known of the ecology of this species. It has been sighted in rocky areas near pastures and on stone walls and piles of rotting vegetation ( Grant 1940 a, Schwartz 1971a; Schwartz & Henderson 1991). The natural habitat of the species is assumed to be the primary limestone forest adjacent to the disturbed areas where collections were made (see Materials and Methods). This species is reported to bask in the sun and is assumed to be diurnal ( Grant 1940a). When examined, the majority of stomach contents were “cow’s tongues,” a slug-like invertebrate ( Grant 1940a). One of us (SBH) spent about a year (cumulative) collecting reptiles all over Jamaica and has encountered only a single individual of this species in the late 1970s. Although it has been seen alive in recent years by resident naturalists, it remains one of the rarest reptiles in Jamaica.

The IUCN Redlist ( IUCN 2023) considers the conservation status of Celestus hewardi to be Endangered B1ab(iii,v) because “the species has an extent of occurrence of around 3,500 km 2 and occurs in one location defined by a significant widespread threat from mongoose predation. There is a continuing decline in the extent and quality of its habitat driven by multiple factors, and in the number of mature individuals inferred from both this and the continuing presence of invasive predators. The species is therefore listed as Endangered.” 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 yet possible on Jamaica.

FIGURE 22. (A–F) Celestus hewardi (BMNH 1946.8.7.99), the same specimen formerly called BMNH 36.12.3.88 and 1836.12.3.88), SVL 117 mm.

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

Etymology. This species was named after Robert Heward, who obtained one individual of the type series.

Remarks. Tiliqua striata was first used in reference to this lizard (Gray 1838); however, this name was a junior homonym of Celestus striatus , which was referenced previously in the same publication. The name C. hewardi was later introduced ( Gray 1845) to correct this oversight. The original description was based on three syntypes (BMNH 1946.8.7.88, 1946.8.7.99, and 1946.8.8.2), one of which (1946.8.7.99) is described in detail above (description of holotype section).

The type locality of Celestus hewardi is unknown, with only “ Jamaica ” being recorded in the original description. The description of C. variegatus (later synonymized with C. hewardi ) recorded the type locality as “unbestimmter Herkunft” (= “undetermined origin”). Grant (1940a) listed this species as C. occiduus hewardi based on a correspondence with Stejneger, in which they agreed that it was possible that the “original ‘galliwasp’ has been exterminated. If that is true, and if the ecological niche of our occiduus has helped in differentiating a recognizable subspecies, my guess would be that the name should be C. occiduus hewardi Gray. ” Our genetic data show that this conclusion was incorrect because C. occiduus is most closely related to C. striatus , and all three species ( C. occiduus , C. hewardi , and C. striatus ) are valid and separated by 3.5–4.3 million years (Fig. 4). Also, these three species are sympatric in the same region of Jamaica.

Grant (1951) said that he believed that Celestus impressus is a synonym of C. occiduus . However, Schwartz (1964) reported that one syntype of C. impressus , which he designated as lectotype (ANSP 9225), was C. hewardi and the other was C. crusculus . We examined those specimens and agree with Schwartz, although the damage to both specimens is substantial. One specimen of C. hewardi was collected with a blue-and-brown-striped tail, a trait of C. duquesneyi , further supporting the close genetic relationship between those two species (Fig. 3) with significant support in our ML analysis and a support value of 71% in our Bayesian analysis. Schools et al. (2022) showed the same topology as our ML analysis with a support value of 70% at the stem node in the ML analysis and a support value less than 50% in the Bayesian analysis. Based on our timetree (Fig. 4), C. hewardi diverged from C. duquesneyi 3.59 Ma, consistent with typical species of vertebrates (> 0.7 Ma; Hedges et al. 2015). Celestus hewardi was recognized as a distinct species by our ASAP analysis.