Celestus crusculus ( Garman 1887 )

Schools, Molly & Hedges, Blair, 2024, A new forest lizard fauna from Caribbean islands (Squamata, Diploglossidae, Celestinae), Zootaxa 5554 (1), pp. 1-306 : 49-54

Celestus crusculus ( Garman 1887)

Common Jamaican Forest Lizard

(Fig. 16–17)

Diploglossus crusculus Garman, 1887:22 . Holotype: MCZ R-6051, collected by Samuel Walton Garman at Kingston, Saint

Andrew Parish , Jamaica, on 22 December 1878 (17.97, -76.78). Diploglossus bakeri — Boulenger, 1900:13. Holotype: BMNH 1946.8 .29.38, presented by Mr. C. H. Baker to the Corporation

Museum of Leicester from Jamaica. Celestus impressus — Barbour, 1922:669. Celestus impressus — Barbour, 1935:123. Celestus impressus — Barbour, 1937:139. Celestus crusculus cundalli Grant, 1940a:157 . Holotype: MCZ R-45163, collected by Chapman Grant in Mandeville between

13–15 February 1937 (18.04, -77.51). Celestus crusculus crusculus — Grant, 1940a:102. Celestus crusculus cundalli — Grant, 1940a:103. Diploglossus crusculus crusculus — Greer, 1967:94. Celestus crusculus — Schwartz & Henderson, 1991:369. Celestus crusculus cundalli — Schwartz & Henderson, 1991:370. Celestus crusculus crusculus — Hedges et al., 2019:17. Celestus crusculus cundalli — Hedges et al., 2019:17. Celestus crusculus — Schools & Hedges, 2021:220. Celestus crusculus cundalli — Schools & Hedges, 2021:220. Celestus crusculus — Landestoy et al., 2022:204.

Material examined (n=26). JAMAICA. Manchester. MCZ R-45163 , MCZ R-45166–7, Chapman Grant, Mandeville , 13–15 February 1937 ; USNM 108220–2 , Mandeville, 22 March 1938 . Saint Andrew. BMNH 1940.3 .11.69, Ivan Sanderson, Constant Spring ; BMNH 1954.1 .2.37, Garth Underwood, “ St. Andrews ” (no specific locality) ; BMNH 1965.129 , 1965.135 , 1965.139 , Mona ; BMNH 1970.1726 , J. Rankin, Red Hills ; BMNH 1970.1727 , Garth Underwood, Papine ; MCZ R-6051, Samuel Walton Garman, Kingston, 22 December 1878 . Saint Ann. BMNH 1970.1714 , Garth Underwood, St. Ann, Runaway Bay (0 m). Saint Catherine . USNM 328186 , S. Blair Hedges and Carla Ann Hass, 5.6 km SW of Braeton (in Hellshire Hills, at Hellshire Beach ), 15 August 1987 (17.897835, -76.894068; 7 m). Trelawny . ANSP 38504 , Carla Ann Hass, S. Blair Hedges, Kimberlyn Nelson, and Stephen Schaeffer, 0.3 km W Duncans (jct with Silver Sands access road), 3 March 1990 (18.47105, -77.53887; 101 m) ; USNM 328159–60 , 328161–7 , Carla Ann Hass, S. Blair Hedges, Kimberlyn Nelson, and Stephen Schaeffer, 0.3 km W Duncans (jct with Silver Sands access road) .

FIGURE 16. (A–F) Celestus crusculus ( MCZ R-6051, holotype), SVL 75.4 mm.

Diagnosis. Celestus crusculus has (1) a dorsal pattern of absent/flecks in series/dots in chevrons, (2) head markings absent/present, (3) markings in the longitudinal paramedian area present, (4) dots arranged in bars in the lateral band absent, (5) an adult SVL of 59.6–77.6 mm, (6) ventral scale rows, 98–114, (7) midbody scale rows, 37–44, (8) total lamellae on one hand, 30–39, (9) total strigae on ten scales, 106–194, (10) relative length of all digits on one hindlimb, 18.7–24.7 %, (11) relative distance between the angled subocular and mouth, 0.339 –0.884 %, (12) relative eye length, 2.93–3.61 %, (13) relative forelimb length, 12.8–20.7 %, (14) relative ear width, 0.716– 2.00 %, (15) relative rostral height, 1.62–2.04 %, (16) relative head length, 15.5–20.3 %, (17) relative mental width, 1.37–2.31 %, (18) relative postmental width, 2.73–3.37 %, (19) relative cloacal width, 6.89–8.77 %, (20) relative prefrontal width, 3.93–4.67 %, (21) relative largest supraocular width, 1.97–2.65 %, (22) relative longest finger length, 2.94–4.10 %, (23) relative distance between the ear and eye, 6.07–8.61 %, (24) relative head width, 72.1–76.4 %, (25) relative frontal width, 82.6–91.1 %, (26) relative nasal height, 0.925–1.37 %, (27) relative angled subocular height, 0.953–1.21 %, (28) relative distance between the eye and naris, 4.31–4.86 %, (29) relative canthal iii length, 1.59–2.07 %, (30) relative angled subocular width, 2.03–2.43 %, and (31) relative nasal length, 1.27–1.60 %. The species stem time is 4.73 Ma and the species crown time is 1.75 Ma (Fig. 4).

We distinguish Celestus crusculus from the other species of Celestus based on a complex of traits. From Celestus barbouri , we distinguish C. crusculus by the dorsal pattern (absent/flecks in series/dots in chevrons versus chevrons), the longitudinal paramedian lines (present versus absent), the adult SVL (59.6–77.6 versus 78.4–93.6), the ventral scale rows (98–114 versus 118–151), the midbody scale rows (37–44 versus 47–56), and the relative frontal width (82.6–91.1 versus 65.6–82.1). From C. capitulatus sp. nov., we distinguish C. crusculus by the relative frontal width (82.6–91.1 versus 78.1–81.6). From C. duquesneyi , we distinguish C. crusculus by the dorsal pattern (absent/flecks in series/dots in chevrons versus bands), the longitudinal paramedian lines (present versus absent), the midbody scale rows (37–44 versus 48), the total lamellae on one hand (30–39 versus 64), the relative length of digits on one hindlimb (18.7–24.7 versus 31.4), the relative eye length (2.93–3.61 versus 4.36), the relative forelimb length (12.8–20.7 versus 24.4), the relative ear width (0.716–2.00 versus 2.45), the relative rostral height (1.62–2.04 versus 2.14), the relative head length (15.5–20.3 versus 21.6), the relative mental width (1.37–2.31 versus 2.35), the relative cloacal width (6.89–8.77 versus 9.98), the relative prefrontal width (3.93–4.67 versus 5.41), the relative largest supraocular width (1.97–2.65 versus 2.66), the relative longest finger length (2.94–4.10 versus 6.52), the relative frontal width (82.6–91.1 versus 75.2), the relative angled subocular height (0.953–1.21 versus 1.61), the relative distance between the eye and naris (4.31–4.86 versus 5.46), the relative angled subocular width (2.03–2.43 versus 2.90), and the relative nasal width (1.27–1.60 versus 2.01). From C. hesperius sp. nov., we distinguish C. crusculus by the relative head width (72.1–76.4 versus 76.5–79.8). From C. hewardi , we distinguish C. crusculus by the dorsal pattern (absent/flecks in series/dots in chevrons versus mottled/bands), the adult SVL (59.6–77.6 versus 129–171), the total lamellae on one hand (30–39 versus 50–61), the relative forelimb length (12.8–20.7 versus 22.2–24.6), the relative cloacal width (6.89–8.77 versus 8.81–9.89), the relative longest finger length (2.94– 4.10 versus 5.03–5.66), the relative frontal width (82.6–91.1 versus 57.3–75.3), and the relative distance between the eye and naris (4.31–4.86 versus 5.00–5.60). From C. jamesbondi sp. nov., we distinguish C. crusculus by the relative frontal width (82.6–91.1 versus 70.5–77.6). From C. macrolepis , we distinguish C. crusculus by the dorsal pattern (absent/flecks in series/dots in chevrons versus bicolored), the longitudinal paramedian lines (present versus absent), the adult SVL (59.6–77.6 versus 254–316), the midbody scale rows (37–44 versus 46–48), the total lamellae on one hand (30–39 versus 52–54), the total strigae on ten scales (106–194 versus 398), the relative length of digits on one hindlimb (18.7–24.7 versus 27.5–28.0), the relative distance between angled subocular and mouth (0.339 –0.884 versus 1.39–1.66), the relative eye length (2.93–3.61 versus 3.63–3.70), the relative forelimb length (12.8–20.7 versus 26.1–26.7), the relative postmental width (2.73–3.37 versus 3.81), the relative cloacal width (6.89–8.77 versus 11.2), the relative longest finger length (2.94–4.10 versus 5.47–5.51), the relative frontal width (82.6–91.1 versus 78.4), the relative distance between the eye and naris (4.31–4.86 versus 6.02), the relative angled subocular width (2.03–2.43 versus 2.57), and the relative nasal width (1.27–1.60 versus 1.75). From C. macrotus , we distinguish C. crusculus by the dorsal pattern (absent/flecks in series/dots in chevrons versus chevrons/bands), the dots arranged in bars in the lateral areas (absent versus present), the ventral scale rows (98–114 versus 87–93), the relative length of digits on one hindlimb (18.7–24.7 versus 30.2–31.2), the relative eye length (2.93–3.61 versus 3.79–5.17), the relative forelimb length (12.8–20.7 versus 22.4–25.0), the relative prefrontal width (3.93–4.67 versus 4.87–5.55), the relative largest supraocular width (1.97–2.65 versus 2.96–4.03), the relative longest finger length (2.94–4.10 versus 6.43–6.67), the relative frontal width (82.6–91.1 versus 57.6–66.1), the relative distance between the eye and naris (4.31–4.86 versus 5.48–5.60), the relative angled subocular width (2.03–2.43 versus 2.77–2.83), and the relative nasal width (1.27–1.60 versus 2.08–2.33). From C. microblepharis , we distinguish C. crusculus by the dorsal pattern (absent/flecks in series/dots in chevrons versus chevrons), the longitudinal paramedian lines (present versus absent), the adult SVL (59.6–77.6 versus 96.4), the relative length of digits on one hindlimb (18.7– 24.7 versus 16.6), the relative eye length (2.93–3.61 versus 1.83), the relative ear width (0.716–2.00 versus 0.446), the relative head length (15.5–20.3 versus 14.7), the relative postmental width (2.73–3.37 versus 2.47), the relative nasal height (0.925–1.37 versus 0.726), the relative angled subocular height (0.953–1.21 versus 0.778), the relative angled subocular width (2.03–2.43 versus 2.90), and the relative nasal width (1.27–1.60 versus 1.11). From C. molesworthi , we distinguish C. crusculus by the adult SVL (59.6–77.6 versus 78.1–103), the relative longest finger length (2.94–4.10 versus 4.28–5.19), and the relative distance between the eye and naris (4.31–4.86 versus 5.32– 5.50). From C. occiduus , we distinguish C. crusculus by the longitudinal paramedian lines (present versus absent), the adult SVL (59.6–77.6 versus 269–367), the midbody scale rows (37–44 versus 46–56), the total lamellae on one hand (30–39 versus 50–66), the total strigae on ten scales (106–194 versus 374), the relative distance between angled subocular and mouth (0.339 –0.884 versus 1.26–1.27), the relative forelimb length (12.8–20.7 versus 23.5– 23.9), the relative head length (15.5–20.3 versus 20.4–20.6), the relative postmental width (2.73–3.37 versus 3.57), the relative cloacal width (6.89–8.77 versus 9.00), the relative prefrontal width (3.93–4.67 versus 4.76), the relative longest finger length (2.94–4.10 versus 4.77–5.46), the relative distance between the ear and eye (6.07–8.61 versus 8.98–10.9), the relative frontal width (82.6–91.1 versus 63.8), the relative angled subocular height (0.953–1.21 versus 1.30), the relative distance between the eye and naris (4.31–4.86 versus 6.51), the relative angled subocular width (2.03–2.43 versus 2.52), and the relative nasal width (1.27–1.60 versus 1.83). From C. oligolepis sp. nov., we distinguish C. crusculus by the midbody scale rows (37–44 versus 35). From C. striatus , we distinguish C. crusculus by the adult SVL (59.6–77.6 versus 145), the total lamellae on one hand (30–39 versus 59–66), the total strigae on ten scales (106–194 versus 279), the relative length of digits on one hindlimb (18.7–24.7 versus 37.8), the relative eye length (2.93–3.61 versus 3.85), the relative forelimb length (12.8–20.7 versus 26.1), the relative prefrontal width (3.93–4.67 versus 5.68), the relative longest finger length (2.94–4.10 versus 7.48), the relative distance between the ear and eye (6.07–8.61 versus 9.00), the relative frontal width (82.6–91.1 versus 76.5), the relative distance between the eye and naris (4.31–4.86 versus 6.16), and the relative width of canthal iii (1.59–2.07 versus 2.12).

Description of holotype. MCZ R-6051. An adult; SVL 75.4 mm; tail nearly cylindrical, broken off at base and tip, tip regenerated, 71.5 mm (94.8% SVL); axilla-to-groin distance 44.9 mm (59.5% SVL); forelimb length 13.8 mm (18.3% SVL); hindlimb length 21.1 mm (28.0% SVL); head length 14.0 mm (18.6% SVL); head width 10.7 mm (14.2% SVL); head width 76.4% head length; diameter of orbit 2.67 mm (3.54% SVL); horizontal diameter of ear opening 0.54 mm (0.716% SVL); vertical diameter of ear opening 1.37 mm (1.82% SVL); length of all toes on one foot 14.1 mm (18.7% SVL); shortest distance between angled subocular and lip 0.39 mm (0.517% SVL); shortest distance between the ocular and auricular openings 6.49 mm (8.61% SVL); longest finger length 2.99 mm (3.97% SVL); largest supraocular width 1.83 mm (2.43% SVL); cloacal width 6.49 mm (8.61% SVL); mental width 1.03 mm (1.37% SVL); postmental width 2.21 mm (2.93% SVL); prefrontal width 3.52 mm (4.67% SVL); frontal width 91.1% frontal length; nasal height 0.92 mm (1.22% SVL); angled subocular height 0.87 mm (1.15% SVL); shortest distance between the eye and naris 3.43 mm (4.55% SVL); canthal iii width 1.21 mm (1.60% SVL); angled subocular width 1.61 mm (2.14% SVL); nasal width 0.97 mm (1.29% SVL); rostral 1.70X 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 slightly concave posterior margin, much wider than long, bordered by posterior internasals, 1 st loreals, canthal iii, 1 st 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 and 2 nd temporals 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 supralabial (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); final 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, 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); 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)/8 (right) supralabials, 6 (left)/5 (right) to level below center of eye; 10 (left)/9 (right) infralabials, 6 (left)/5–6 (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; 2 nd –4 th pairs separated by 1–3 scales; 96 transverse rows of dorsal scales from interoccipital to base of tail; 100 transverse rows of ventral scales from mental to vent; 42 scales around midbody; 5 digits; finger lengths 3>4>2>5>1; 8 (left)/7 (right) lamellae under longest finger; 30 total lamellae on one hand; toe lengths 4>3>5>2>1; 14 lamellae under longest toe on the right side; many missing, striate with a small median keel towards trunk dorsal body and caudal scales; smooth ventral scales.

Color (in alcohol): mid shed, dorsal surface of head dark gold, patternless; lateral surfaces of head grading from dark brown to gray-tan with irregular brown mottling and darker brown eye masks; dorsal surfaces of the body are pale brown with two darker longitudinal paramedian lines that end before the forearms and brown dots arranged into almost complete chevrons; dorsal surface of tail the same as body with more complete chevrons; lateral areas grade from medium brown to cream; dorsal surfaces of the limbs are medium brown with some irregular brown mottling; lateral and ventral areas of the limbs fade to gray-tan, patternless; ventral surfaces of the head, body, and tail are cream, darker brown mottling begins under the head and ends near the forelimbs.

Variation. The majority of the specimens are similar to the holotype in scalation and pattern with specimens ranging in dorsal pattern from obscure mottling to chevrons. All specimens other than the holotype exhibit markings on their head scales. The longitudinal paramedian markings are most pronounced in the holotype whereas they range from being completely absent to very thin in the other specimens. Measurements and other morphological data for the holotype and other examined material are presented in Table 1.

Ecology and conservation. No ecological information is associated with the type specimen. The specimens used in our genetic analysis were found under rocks along the road. Past literature accounts of ecological data for this species conflate multiple species and therefore cannot be used. However, this species is considered to be common (SBH), has a relatively wide distribution, and is tolerant of human disturbance.

We consider the conservation status of C. crusculus sp. nov. to be Least Concern, based on IUCN Redlist criteria ( IUCN 2023). However, studies are needed to determine the health and extent of the populations and any threats to the survival of the species.

Reproduction. Past literature accounts of reproductive data ( Greer 1967) for this species conflate multiple species and therefore cannot be used.

Distribution. Celestus crusculus is distributed in and around Kingston, Jamaica, and extending through the central areas of the country to the north coast at elevations of 0–860 m (Fig. 12).

FIGURE 17. Celestus crusculus (voucher not available), in life. From 0.3 km W Duncans, Trelawny Parish, Jamaica. Photo by SBH.

Etymology. The species name crusculus is derived from the Latin “ crusculum ” meaning “little leg”, likely in reference to the species’ shorter legs.

Remarks. The description of this species lists the type locality as “ Kingston, Jamaica.” Cochran (1941) speculated that Celestus crusculus could possibly be young C. occiduus , which was incorrect because adult male and female C. crusculus exist and the two species differ in habitus and scalation. This species was confused with C. impressus by Thomas Barbour (1922, 1935, 1937), a taxon that we consider here to be a synonym of C. hewardi . Diploglossus bakeri (locality unknown other that “ Jamaica ”) was designated as a synonym for C. crusculus by Barbour (1910), who described Diploglossus bakeri as “identical” to C. crusculus . We were unable to examine the holotype of D. bakeri and therefore additional studies should examine it for the morphological characters presented herein to confirm this placement.

Both Grant (1940a) and Barbour (1922, 1935, & 1937) considered this species to be common but poorly-known. When Grant (1940a) introduced Celestus crusculus cundalli as a subspecies of C. crusculus , he noted that scale counts between the two subspecies blended together on an elevational gradient. We agree with Cousens (1956) that C. crusculus cundalli is not supported as a valid taxon with available morphological data, although a thorough genetic analysis of C. crusculus is warranted (i.e., with additional samples from throughout the distribution of that taxon).

Celestus crusculus is included in our genetic dataset and has significant support values at the crown node in our Bayesian and ML analyses. The stem node that identifies it as the closest relative of C. oligolepis sp. nov. and C. barbouri has significant support in our ML analysis and 93% in our Bayesian analysis. Based on our timetree (Fig. 4), C. crusculus diverged from its closest relative 4.73 Ma, consistent with typical species of vertebrates (> 0.7 Ma; Hedges et al. 2015). Celestus crusculus was recovered as conspecific with Celestus jamesbondi sp. nov. in our ASAP analysis.