Spondylurus haitiae, Hedges & Conn, 2012

Hedges, S. Blair & Conn, Caitlin E., 2012, A new skink fauna from Caribbean islands (Squamata, Mabuyidae, Mabuyinae) 3288, Zootaxa 3288 (1), pp. 1-244 : 158-163

Spondylurus haitiae sp. nov.

Hispaniolan Four-lined Skink

( Figs. 55E , 64A , 65 )

Mabuia nitida — Garman, 1887:51 (originally a syntype).

Mabuya sloanii — Stejneger, 1904:608 (part).

Mabuya sloanii — Barbour, 1914:320 (part).

Mabuya sloanii — Schmidt, 1928:121 (part; removed from type series of Mabuia nitida ).

Mabuya sloanii —Barbour, 1930:105 (part).

Mabuya mabouia — Barbour, 1935:129 (part).

Mabuya mabouya sloanii — Dunn, 1936:544 (part).

Mabuya mabouia — Barbour, 1937:147 (part).

Mabuya mabouya sloanei — Schwartz & Thomas, 1975:141 (part).

Mabuya mabouya sloanei — MacLean et al., 1977:24 (part).

Mabuya mabouya sloanei — Schwartz & Henderson, 1988:151 (part).

Mabuya mabouya sloanei — Schwartz & Henderson, 1991:457 (part).

Mabuya bistriata — Powell et al., 1996:82 (part).

Mabuya sloanii — Mayer & Lazell, 2000:883 (part).

Holotype. MCZ R-3617, an adult female from "San Domingo" collected by D. F. Weinland, here restricted to Jérémie, Grand'Anse, Haiti. Date of collection inferred to be 1857–58 (see Remarks).

Paratypes (n = 4). Haiti. MCZ R-189392–395, fetuses from the holotype .

Diagnosis. Spondylurus haitiae sp. nov. is characterized by (1) maximum SVL in males, not available; (2) maximum SVL in female holotype, 85.2 mm; (3) snout width, 2.69% SVL; (4) head length, 15.8% SVL, (5) head width, 12.3% SVL; (6) ear length, 1.19% SVL; (7) toe-IV length, 9.01% SVL; (8) prefrontals, two; (9) supraoculars, four; (10) supraciliaries, four; (11) frontoparietals, one (50%), two (50%); (12) supralabial below the eye, six; (13) nuchal rows, two; (14) dorsals, 59–60; (15) ventrals, 69–72; (16) dorsals + ventrals, 129–131; (17) midbody scale rows, 30–32; (18) finger-IV lamellae, 12–13; (19) toe-IV lamellae, 16–17; (20) finger-IV + toe-IV lamellae, 29–30; (21) supranasal contact, Y (50%), N (50%); (22) prefrontal contact, Y (50%), N (50%); (23) supraocular-1/frontal contact, N; (24) parietal contact, Y; (25) pale middorsal stripe, Y; (26) dark dorsolateral stripe, Y; (27) dark lateral stripe, Y; (28) pale lateral stripe, Y; and (29) palms and soles, pale ( Tables 3–5).

Within the Genus Spondylurus , S. haitiae sp. nov. is separated from S. culebrae sp. nov., S. fulgidus , S. macleani , S. magnacruzae sp. nov., S. martinae sp. nov., S. monae sp. nov., S. monitae sp. nov., S. nitidus , S. powelli sp. nov., S. spilonotus , and S. turksae sp. nov. by having a smaller ear (ear length 1.19% SVL versus 1.23– 2.83% in those other species). It differs from S. fulgidus , S. lineolatus , S. macleani , S. nitidus , S. powelli sp. nov., S. sloanii , S. spilonotus , and S. turksae sp. nov. by having a higher number of ventral scales (69–72 versus 55–68 in those other species). From S. anegadae sp. nov., S. culebrae sp. nov., S. monae sp. nov., S. monitae sp. nov., and S. semitaeniatus , it is distinguished by having a narrower dark dorsolateral stripe (dorsolateral stripe width 2.12% SVL versus 2.24–4.64% SVL in those other species). It differs from S. culebrae sp. nov., S. fulgidus , S. macleani , S. magnacruzae sp. nov., S. monae sp. nov., S. monitae sp. nov., S. nitidus , and S. semitaeniatus by having a shorter head (head length 15.8% SVL versus 15.9–21.6% SVL in those other species). From S. magnacruzae sp. nov., S. martinae sp. nov. and S. spilonotus , it is distinguished by having wider dark dorsolateral stripes (dorsolateral stripe width 2.12% SVL versus 1.16–2.09% SVL in those other species). It is separated from S. lineolatus by having two dark dorsolateral stripes and two dark lateral stripes (versus 10 dark stripes in S. lineolatus ). It differs from S. nitidus by having a shorter toe-IV (toe-IV length 9.01% SVL versus 9.45–12.7% SVL in S. nitidus ). From S. semitaeniatus , it is separated by having a shorter head (head length 15.8% SVL versus 15.8– 19.4% SVL in S. semitaeniatus ).

Besides those non-overlapping differences, there are frequency differences separating Spondylurus haitiae sp. nov. from other species. From S. anegadae sp. nov. it differs by having a smaller ear (ear length 1.19% SVL versus 1.23–2.10% in 88% of specimens belonging to S. anegadae sp. nov.). It differs from S. caicosae sp. nov. by having a higher number of dorsals + ventrals (dorsals + ventrals 129–131 versus 113–127 in 95% of specimens belonging to S. caicosae sp. nov.) and by having a smaller ear (ear length 1.19% SVL versus 1.26–2.18% SVL in 92% of specimens belonging to S. caicosae sp. nov.). Besides the three diagnostic characters noted above, which separate S. haitiae sp. nov. from S. nitidus (the taxon with which it has been confused), it also has a greater number of dorsals + ventrals: 129–131 versus 117–127 in 12 specimens of S. nitidus (except for one specimen with 129).

Description of holotype ( Figs. 64A , 65 ). An adult female in poor state of preservation posteriorly (soft), with ventral and lateral midbody injuries, abdominal slit, and disappearance of pattern (scale clearing) in posterior half of body. SVL 85.2 mm; tail length 28.4 mm (regenerated); HL 13.5 mm; HW 10.5 mm; SW 2.29 mm; EL 1.01 mm; and toe-IV length 7.68 mm; ear-opening small in size and round; toe length in the following order: I <V <II <III <IV.

Head scalation. Rostral wider than high, contacting first supralabials, nasals and supranasals. Paired supranasals not in median contact (but close), contacting anteriormost loreal. Frontonasal pentagonal and lanceolate, wider than long, laterally in contact with anterior loreal scale. A pair of quadrilateral prefrontals, separated medially, and in contact with frontonasal, both anterior and posterior loreals, first supraoculars, and frontal. Frontal heptagonal and lanceolate, in contact with the second supraoculars and paired frontoparietals. Frontoparietals also in contact with parietals and interparietal. Interparietal tetragonal and lanceolate, separated from nuchals by parietals; parietal eye distinct. Parietals in contact with upper secondary and tertiary temporal scales. Four supraoculars, the second one being the largest. Four supraciliaries, the second the longest. Nostril in posterior part of the nasal. A small postnasal, bordered by supranasal, anterior loreal and first supralabial. Anterior moderately enlarged scales behind eye on the right and five on the left comprising the postoculars; similar to temporal scales but smaller. One primary temporal, two secondary temporals, and three tertiary temporals; all imbricate, smooth, cycloid, not distinctly delimited from the scales on the nape and the sides of the neck. Seven infralabials. Mental scale wider than long, posterior margin straight. Postmental scale and two pairs of adjoining chin shields in contact with anterior infralabials. First two pairs of chin shields in contact medially; third pair separated by a smaller cycloid scale.

Body and limb scalation. Two rows of nuchal scales, both paired. Other scales on nape similar to dorsals. On lateral sides of neck, scales slightly smaller. Dorsal scales cycloid, imbricate, smooth, 60 in a longitudinal row; ventrals similar to dorsals; 72 in a longitudinal row; 32 scales around midbody. No distinct boundaries between dorsals, laterals and ventrals. Scales on tail and limbs similar to dorsals, except smaller on limbs and on regenerated portion of tail. Palmar and plantar regions with small tubercles, subequal in size and delimited by a surrounding region of flatter scales. Subdigital lamellae smooth, single, 13 under finger-IV and 16 under toe-IV. Four preanals larger than adjacent ventral scales. Enlarged median subcaudal scales on regenerated portion of tail.

Pattern and coloration. Cochran (1941) described the pattern in this specimen when it was in better condition, so we first quote from her account: "Above bronze green, with a pale dorsolateral band extending from the supranasals over the supraciliaries to the middle of the body, where it gradually disappears; this band is bordered above by a narrow dark stripe and below by a broader dark brown band over ear and shoulder, gradually becoming lighter and disappearing on the groin; labials dark edged; a short light stripe beginning on the upper lip, passing beneath the ear, reaching the shoulder and fading out on the sides; lower parts pale bluish gray." Presently, the dorsal ground color is medium grayish-brown with dark brown spots, distributed on body and tail. Forelimbs brown with darker brown mottling on dorsal surfaces and patternless on ventral surfaces. Hindlimbs without spots or mottling. Dark dorsolateral stripes present, narrow (1.81 mm), dark brown, extending from top of head to first third of body. Dark lateral stripes present, dark brown with pale spots, extending from loreal region at least to first third of body, where no pattern remains due to poor preservation. Pale middorsal stripe present, wide (2.83 mm), grayish-brown, extending from tip of snout to at least first third of body. Pale dorsolateral stripes present, pale gray—noticeably paler than middorsal stripe, and especially so in photograph in Cochran (1941) —extending from tip of snout at least to first third of body. Pale lateral stripes present, whitish, extending from below ear at least to first third of body, bordered below by irregular brown spots. Ventral surface of body without pattern. Palmar and plantar surfaces unpigmented. There is no information on color in life of the holotype.

Variation. In coloration and scalation, the fetuses all resembled the holotype ( Tables 4–5).

Distribution. The species is known only from the type-locality of Jérémie, Haiti ( Fig. 9B ).

Ecology and conservation. There are no ecological data known for this species, and it has not been collected since the holotype was taken in 1857–58. As with declines in other species of skinks in the late 19th and early 20th centuries, this species was likely affected by the introduction of the mongoose. Jérémie is located on the Tiburon Peninsula of Haiti, and in that region (Massif de la Hotte and foothills) there are small patches of original forest, even at low elevations (where dissected limestone substrate makes agriculture difficult), and therefore it is not possible to say with certainty that this species is extinct. The persistence of the other two Hispaniolan species, Mabuya hispaniolae sp. nov. and Spondylurus lineolatus , to at least 1937 and 1985 (respectively), demonstrates that the introduction of the mongoose did not result in immediate extinction of all skink species on the island.

The FAO (2005) lists total forest area of Haiti as 4.0% and Dominican Republic as 28.4%, but these numbers are inflated because their definition of total forest includes areas with up to 90% of the trees missing (10% canopy). Primary forest area values are not listed by FAO for these countries, but where they are listed elsewhere, they average 10–20% of total forest ( Hedges 2006a). Therefore the primary forest of Haiti is likely <1% of total land area, and that of the Dominican Republic, ~5% of land area. There are national parks and protected areas in Haiti and the Dominican Republic, but deforestation takes place within park boundaries, and therefore they do not afford protection.

Based on IUCN Redlist criteria ( IUCN 2011), we assess the conservation status of Spondylurus haitiae sp. nov. to be Critically Endangered and possibly extinct (CR A2ace). It faces a primary threat from the introduced mongoose, which has probably led to its extinction. Other major threats include habitat destruction from agriculture and charcoaling, and predation from other introduced mammals, including black rats. Studies are introduced mammalian predators is not possible on large islands. All mongoose-free islets of Hispaniola need to be thoroughly surveyed to determine if this species, or the other two species of mabuyines from Hispaniola, still exist.

Reproduction. The holotype (85.2 mm SVL) contained four well-developed fetuses, designated here as paratypes. No specific date (month) of collection is available .

Etymology. The species name ( haitiae ) is a feminine genitive singular noun derived from the Amerindian (Taino) name for the entire island of Hispaniola, transliterated in English as " Haiti " or "Hayti," meaning "high mountains."

Remarks. This specimen was originally a syntype of Mabuia nitida Garman. Stejneger (1904) placed that species in the synonymy of Mabuya sloanii . Schmidt (1928) retained " Mabuya nitida " in the synonymy of M. sloanii but noticed that this syntype from Hispaniola differed from the Puerto Rican syntypes (and other specimens from Puerto Rico) sufficiently that he restricted the name " Mabuya nitida " to Puerto Rico, should the species ever be considered valid (as it is here). Garman (1887) listed the type-locality of the species as " Porto Rico and San Domingo" (MCZ R-3617 representing "San Domingo"). The name "San Domingo" has been interpreted by subsequent authors, and the MCZ, as pertaining to the city of Santo Domingo, Dominican Republic, and global databases have used the GPS coordinates for the city, as the locality. However, this place name (and " Santo Domingo ") was commonly applied to the entire island of Hispaniola during the nineteenth century, as was the name " Haiti " or "Hayti" ( Keim 1870; Barbour 1914; see historical maps in Hedges 2011). "San Domingo," to a lesser extent, was also a name used for the newly independent country now called the Dominican Republic during the late nineteenth century (other names for the country were " Dominica " and "Santo Domingo"). To distinguish the city of Santo Domingo from the country or island, the word "city" was usually used (e.g., "San Domingo city"), and therefore it is nearly certain that Garman was not referring to the city of Santo Domingo. He did not list a collector (perhaps for brevity because there were multiple syntypes and collectors), but the MCZ records the collector as "D. F. Weinland," with no date of collection or other information.

David Friedrich Weinland was a German zoologist and collector of animals who spent six months in Jérémie, Haiti, during 1857–58 ( Weinland 1858). The MCZ collections record 475 specimens (various animals) associated with his name. Of the large number from Caribbean locations, nearly all are from Jérémie, Haiti, as would be expected. A few are from other specific locations in the country of Haiti, and two snails are from the Bahamas. Barbour's (1914) reference to "Haitian" material of Spondylurus sloanii is uninformative for our purpose because he preferred to use the original Taino name for the entire island ( Haiti). Cochran (1941) examined this specimen, referring to the type locality variously as " San Domingo " and " Santo Domingo " and noting a collection date of 1859, information not currently associated with the specimen (and a year after Weinland left the island). Because (1) " San Domingo " or " Santo Domingo " was the common name for the entire island at the time of Weinland and Garman, (2) there is no evidence that Weinland ever collected in the Dominican Republic, and (3) nearly all of his specimens came from the vicinity of Jérémie, Haiti, we have restricted the type-locality of this species to Jérémie, Haiti, with a collection date of 1857–58. Cochran's date of 1859 was probably the date that the specimen was received at MCZ.

Cochran (1941) concluded that MCZ R-3617 was "very similar" to specimens that we place here in another species ( M. hispaniolae sp. nov.) and genus ( Mabuya ). However her table of characters for the specimens shows the major differences in ventrals and nuchals that separate the two species. Cochran indicated three supraoculars in MCZ R-3617, but it has four, another character separating it from M. hispaniolae sp. nov. Other diagnostic characters are noted above. Two photographs of the holotype of Spondylurus haitiae sp. nov. are shown, as " Mabuya mabouya sloanii ," in Cochran (1941). With its mouth preserved in open position, and precise details of pattern matching, it is clearly the same animal that we examined, but the pattern is strikingly bold, compared with its much faded current state. Cochran (1941) also made no mention of its poor state of preservation and described pattern in the posterior half of the body, which is now soft and patternless (scales cleared). Apparently the specimen was in excellent condition after its first 80 years but suffered significant damage in its second 80 years.

A specimen in the Slater Museum (PSM 10269), labeled as Mabuya mabouya from Restauración, Dominican Republic, was examined by us and found to be Celestus costatus (Anguidae) . Also BMNH 1982.1448 from Portau-Prince Haiti, cataloged as Mabuya mabouya , is not a skink and probably a Celestus as well (Colin McCarthy, BMNH, personal communication).

Figures

FIGURE 9. Map showing the distribution of mabuyine skinks occurring on islands in the central, southern, and western Caribbean region. See Fig. 1 for location of the islands. The islands and species are: (A) Jamaica (Spondylurus fulgidus; hollow circle = specimen not examined); (B) Hispaniola (Spondylurus haitiae sp. nov., circle; Mabuya hispaniolae sp. nov., triangle; Spondylurus lineolatus, squares); (C) Turks and Caicos Islands (Spondylurus caicosae sp. nov., circles; Spondylurus turksae sp. nov., triangles; hollow triangle is Cotton Cay record (see text); (D) Margarita (Copeoglossum margaritae sp. nov.); and (E) Roatán (Marisora roatanae sp. nov.).

FIGURE 55. Representative patterns in species of the Genus Spondylurus (top, dorsal view; bottom, side view). (A) S. anegadae sp. nov.; (B) S. caicosae sp. nov.; (C) S. culebrae sp. nov.; (D) S. fulgidus; (E) S. haitiae sp. nov.; (F) S. lineolatus; (G) S. macleani; (H) S. magnacruzae sp. nov.; (I) S. martinae sp. nov.; and (J) S. monae sp. nov.

FIGURE 64. Head scalation in species of the Genus Spondylurus (left, side view; right, dorsal view). (A) S. haitiae sp. nov. (MCZ R-3617, holotype); (B) S. lineolatus (MCZ R-156938); (C) S. macleani (MCZ R-182270, paratype); and (D) S.

FIGURE 65. Spondylurus haitiae sp. nov., from Hispaniola. (A–D) MCZ R-3617, holotype, near Jérémie, Haiti. E, fetuses of