Copeoglossum aurae, 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 : 72-79

Copeoglossum aurae sp. nov.

Greater Windward Skink

( Figs. 23A , 24A , 25)

Mabuia agilis — Boulenger, 1887:191 (part).

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

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

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

Mabuya aenea — Underwood, 1963:83 (part).

Mabuya mabouya mabouya —Peters & Donoso-Barros, 1970:200 (part).

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

Mabuya mabouya mabouya — MacLean et al., 1977:40–41 (part).

Mabuya mabouya mabouya — Schwartz & Henderson, 1988:150 (part).

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

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

Mabuya bistriata — Murphy, 1997:150 (part).

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

Mabuya mabouya —Miralles, 2005:49 (part?).

Mabuya nigropunctata —Miralles et al., 2005:833 (part).

Mabuya nigropunctata —Miralles et al., 2009:609 (part).

Mabuya mabouya — Henderson & Powell, 2009:292 (part).

Holotype. MCZ R-185624, an adult male, collected in Kingston , St. George Province, St. Vincent, in April 1886 by W.B. Richardson.

Paratypes (n = 32). Grenada. BMNH 86.6 .29.19, G. Murray, "Island of Grenada," 1886; MCZ R-79744, James Lazell, Tempe , 22 June 1964 . Grenadines. KU 242047–48 , Albert Schwartz, Park, Bequia Island , 18 March 1961 ; KU 242051, Albert Schwartz, Grand Bay, Mustique Island , 15 December 1961 ; MCZ R-79097 and R-79099, C. MacIntosh, Carriacou, 1963; MCZ R-79100, George Gorman, Bequia Island , 30 April 1964 ; USNM 79131 , Paul Bartsch, Petit Martinique , 1929 . St. Vincent. BMNH 90.11.25.16–17, F. D. Goodman, no specific locality, 1890; CAS 39439, Blake Expedition, Kingston , 18 February 1879 ; MCZ R-6040, Samuel W. Garmin, Kingston , 18 February 1879 ; MCZ R-6041, R-185623, R-185625–26, W. B. Richardson, Kingston , April 1886 . Trinidad. AMNH R101327 , George Campbell, Chaguaramas , February 1943 ; CAS 39483, C. S. Cazabon, no specific locality, 1879; CAS 231775 , R. Lawson et al. Manzanilla Beach , 18 July 2004 ; CM 6565 , M. Graham Netting, Manzanilla Beach , 22 September 1927 ; KU 242013, Albert Schwartz, Nariva, 4 May 1963 ; MCZ R-6039, Samuel Garmin (no additional information available); MCZ R-8994, R. Thaxter , Port of Spain, April 1913 ; MCZ R-79818, James D. Lazell, Aripe Savannah , 14 July 1964 ; UMMZ 79919 and 239606, no specific locality, collector, or date; MCZ R-100481, H. Boos, Huevos Island , 3 May 1966 . Tobago. KU 242009–11 , Albert Schwartz, 4 miles N Mt. St. George , 12 May 1963 ; MCZ R-185622, Garth L. Underwood, Scarborough , 5 September 1956 .

Diagnosis. Copeoglossum aurae sp. nov. is characterized by (1) maximum SVL in males, 98.5 mm; (2) maximum SVL in females, 109 mm; (3) snout width, 2.64–3.63% SVL; (4) head length, 16.3–20.7% SVL; (5) head width, 12.5–16.1% SVL; (6) ear length, 1.02–2.19% SVL; (7) toe-IV length, 10.1–12.7% SVL; (8) prefrontals, two; (9) supraoculars, three (3%), four (97%); (10) supraciliaries, four (3%), five (66%), six (31%); (11) frontoparietals, two; (12) supralabial below the eye, six (91%), seven (9%); (13) nuchal rows, one; (14) dorsals, 50–59; (15) ventrals, 54–65; (16) dorsals + ventrals, 106–117; (17) midbody scale rows, 28–32; (18) finger-IV lamellae, 12–14; (19) toe-IV lamellae, 15–18; (20) finger-IV + toe-IV lamellae, 29–32; (21) supranasal contact, N; (22) prefrontal contact, N; (23) supraocular-1/frontal contact, Y (16%), N (84%); (24) parietal contact, Y (3%), N (97%); (25) pale middorsal stripe, N; (26) dark dorsolateral stripe, N; (27) dark lateral stripe, Y; (28) pale lateral stripe, Y; and (29) palms and soles, dark ( Table 2).

Within the Genus Copeoglossum , C. aurae sp. nov. differs from C. arajara ( Rebouças-Spieker 1981) in having more finger-IV + toe-IV lamellae (29–32 versus 26 in C. arajara ), more supraciliaries (5–6 in 97%, four in 3% of C. aurae sp. nov. versus four, or rarely three or five, in C. arajara ), and dark palms and soles (pale in C. arajara ). Copeoglossum aurae sp. nov. differs from C. nigropunctatum in having a narrower frontonasal scale (frontonasal width/length 0.97–1.28 versus 1.30–2.03 in C. nigropunctatum ; Fig. 26 ), a shorter supraocular-2 scale (supraocular-2/supraocular-1 length ratio 0.38–0.69 in C. aurae sp. nov. versus 0.68–1.23 in C. nigropunctatum ; Fig. 27 ), and supranasals not in contact (in contact in 94% of C. nigropunctatum ). It differs from C. margaritae sp. nov. in having a lower number of ventral scales (54–65 versus 66), a shorter supraocular-2 scale (supraocular-2/ supraocular-1 length ratio 0.38–0.69 in C. aurae sp. nov. versus 0.75–0.88 in C. margaritae sp. nov.) ( Fig. 27 ), a smaller body size (109 versus 120 mm maximum SVL ), and a longer toe-IV (length 10.1–12.7% SVL versus 9.06%), although the relatively shorter toe-IV in C. margaritae sp. nov. may be related to its large size. Copeoglossum aurae sp. nov. also lacks spotting on the chin (present in C. margaritae sp. nov.). Copeoglossum aurae sp. nov. differs from C. redondae sp. nov. in having no contact between the first paired chin shields and infralabials (broad contact in C. redondae sp. nov.), a smaller ear (ear length 1.02–2.19% SVL versus 2.42% SVL in C. redondae sp. nov.), and a longer toe-IV (10.1–12.7% SVL versus 9.43% in C. redondae sp. nov.). Preserved C. aurae sp. nov. appear to be paler in coloration than C. nigropunctatum and C. margaritae sp. nov., noted also by Ugueto and Rivas (2010), but that was a character difficult to quantify or classify, especially given the effects of preservation and age on coloration.

Copeoglossum aurae sp. nov. also differs in many ways from a sympatric species, Marisora aurulae sp. nov., described below, in a different genus. One scale character that may be used to separate them readily is parietal contact (not in contact, or rarely just touching, in C. aurae sp. nov.; in contact in M. aurulae sp. nov.). Another involves chin scales in contact with infralabials between postmental and first sublabial. There are none in C. aurae sp. nov. and 2–4 in M. aurulae sp. nov. ( Fig. 3 ). In coloration, C. aurae sp. nov. nearly always has a distinctly spotted dorsum and dark lateral stripes that extend to the hindlimbs and onto the tail ( Fig. 24A ). Marisora aurulae

Description of holotype (Fig. 25A–C). An adult male in good state of preservation, without injuries and with two abdominal slits. SVL 90.7 mm; tail length 64.6 mm (regenerated and broken); HL 17.2 mm; HW 12.1 mm; SW 2.92 mm; EL 1.86 mm; and toe-IV length 10.7 mm; ear-opening large in size and oval; 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, contacting anteriormost loreal. Frontonasal diamond-shaped, 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, in contact with the second supraoculars and paired frontoparietals. Frontoparietals also in contact with parietals and interparietal. Interparietal tetragonal and lanceolate, fused with right parietal, not 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. Six supraciliaries, five approximately equal in length and the sixth smaller than the others. Nostril in posterior part of the nasal. Postnasal bordered by supranasal, anterior loreal and first and second supralabials. Anterior loreal rectangular and posterior loreal squarish with posterodorsal projection on latter. One upper preocular and two lower preoculars. Eight supralabials, the sixth being the widest and forming the lower border of the eyelid. Five moderately enlarged scales behind eye 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. Eight infralabials. Mental scale wider than long, posterior margin curved toward the tip of the snout. Postmental scale and zero 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. One row of paired nuchal scales. Other scales on nape similar to dorsals. On lateral sides of neck, scales slightly smaller. Dorsal scales cycloid, imbricate, smooth, 53 in a longitudinal row; ventrals similar to dorsals; 58 in a longitudinal row; 29 scales around midbody. No distinct boundaries between dorsals, laterals and ventrals. Scales on tail and limbs similar to dorsals, except smaller on limbs. On regenerated portion of tail, one enlarged row each of middorsal and midventral scales and 3–4 lateral rows on each side similar to dorsals and ventrals. Palmar and plantar regions with small tubercles, subequal in size and delimited by a surrounding region of flatter scales. Subdigital lamellae smooth, single, 14 under finger-IV and 18 under toe-IV. Five preanals larger than adjacent ventral scales. Enlarged median subcaudal scales on regenerated portion of tail.

Pattern and coloration. Dorsal ground color medium grayish-green with small-to-medium dark brown spots (usually 1/2 to 1/5 of a scale in size), uniformly distributed on body, base of tail, and limbs (limbs more densely spotted and with larger spots). Dark dorsolateral stripes absent. Dark lateral stripes present, brown, irregular (series of close medium and dark brown blotches), extending from loreal region onto tail. Pale middorsal stripe absent. Pale dorsolateral stripes absent. Pale lateral stripes present, greenish-white, extending from below ear to hindlimbs, bordered below by a series of discontinuous brown spots. Two thin dark brown dorsolateral stripes on regenerated part of tail. Ventral surface of body without pattern. Palmar and plantar surfaces dark brown. No information is available on color in life of the holotype.

Variation. In coloration (in preservative), most specimens resembled the holotype, except that dorsal color varies among shades of brown, gray, and green ( Table 5). The dorsum of some specimens is heavily spotted, sometimes with dark brown spots aligned dorsolaterally; others have fewer spots. Some specimens have paler areas just above the dark lateral stripes, but on others, the dorsal color is constant.

A photo of a live individual from Union Island, Grenadines is shown here (Fig. 25D). Ugueto and Rivas (2010) noted that skinks from Trinidad and Tobago are paler than those (= C. nigropunctatum ) from South America. The photo of a skink from near Arima, Trinidad ( Murphy 1997) is of C. aurae sp. nov. (as opposed to M. aurulae sp. nov.) because it shows separated parietal scales. The greenish hue seen in many preserved specimens is an artifact of preservation, as it is not seen in these live individuals.

Distribution. Copeoglossum aurae sp. nov. is distributed on Grenada, St. Vincent, the Grenadines (Bequia, Carriacou, Mustique, Petit Martinique, and Union Islands), Trinidad (including Huevos Island), and Tobago ( Fig.

11D, I–J). The Union Island record is based on image identification (Fig. 25D). A DNA sequence from an uncataloged specimen collected on the nearby Peninsula de Paria (Sucre, Venezuela), reported by Miralles and Carranza (2010), clusters with this species in our tree ( Fig. 5 ), indicating that the species also occurs on the mainland, possibly restricted to that peninsula. Previous distributional data in the literature ( Murphy 1997; Daudin & de Silva 2007) cannot be used because it confounds C. aurae sp. nov. and Marisora aurulae sp. nov.

Ecology and conservation. No ecological information is available specifically for this species. Past ecological information reported for skinks from Trinidad and Tobago, summarized in Murphy (1997), probably confounds Copeoglossum aurae sp. nov. and Marisora aurulae sp. nov. In those reports, skinks were noted as occurring in a diversity of habitats, including rainforest, forest edge, coconut trash, and cultivated and disturbed areas. In the Grenadines, skinks have been found usually on the ground "in woody underbrush and between cacti" and climbing among cacti and on tree trunks ( Daudin & de Silva 2007). According to Barbour (1937), skinks were already extirpated from the large islands of St. Vincent and Grenada by 1937, by the introduced mongoose. However, one specimen of C. aurae sp. nov. was collected in 1964 at Tempe, Grenada. Many herpetologists have visited Grenada and St. Vincent in the last four decades and no sightings of skinks have been reported. Circumstantial evidence suggests that black rats ( Rattus rattus ) are also predators, and these are on many islands. Skinks have not been extirpated from Trinidad, despite the presence of the mongoose on that island. In the past, Trinidad has had geological connections with South America and has a continental mammalian fauna that included natural predators of skinks, which may explain how they have survived ( Murphy 1997). We identified more than twice as many specimens in museums of C. aurae sp. nov. than of Marisora aurulae sp. nov., suggesting that C. aurae sp. nov. is the more abundant species of the two. Now that these two species have been identified and described, studies are needed to assess their ecological relations and further clarify their conservation status.

Based on IUCN Redlist criteria ( IUCN 2011), we assess the conservation status of Copeoglossum aurae sp. nov. as Vulnerable (VU A2ace). It faces a primary threat from the introduced mongoose, which has probably led to its extirpation on Grenada and St. Vincent. Secondary threats include habitat destruction from agriculture and urbanization, and predation from other introduced predators, including black rats. Studies are needed to determine if the species still exists on Grenada and St. Vincent, the health of any remaining populations, and threats to the survival of the species.

Reproduction. Two females (93.0 and 95.3 mm SVL) contained one and three developing young. The dates of collection for those specimens were 12 May 1963 and 18 March 1961, respectively.

Etymology. The species name ( aurae ) is a feminine genitive singular noun, from the Latin noun aura (wind), alluding to its distribution on the Windward Islands (the southern Lesser Antilles, sometimes including Trinidad and Tobago). The term "windward" is in reference to early travel across the Atlantic in sailing ships, which encountered these islands first because of wind direction. The Leeward Islands were those encountered subsequent to the Windward Islands. There has been some confusion over the centuries as to which islands should be called windward and leeward, but those in the southern Lesser Antilles ( Saint Vincent, Grenadines, Grenada) have always been called windward ( Hedges 2011). The first part of the common name (Greater Windward Skink) refers to the larger body size of this species compared with M. aurulae sp. nov. (Lesser Windward Skink), described below.

Remarks. The name Tiliqua (Mabuya) aenea ( Gray 1831a) does not to apply to this species. We have examined the two syntypes, BMNH 1946.8.15.12 and 1946.8.19.78, and they agree with populations currently placed in Copeoglossum nigropunctatum from South America. Both specimens have supranasals in broad contact (typical of C. nigropunctatum ), whereas all 33 specimens examined of C. aurae sp. nov. (from throughout its range) lack supranasal contact, something alluded to by Boulenger (1891) when he reported on two specimens of what is here called C. aurae sp. nov. (BMNH 90.11.25.16–17) from St. Vincent that did not agree with the type of M. aenea . Also the ratios of frontonasal width versus length in the two syntypes of Tiliqua aenea (1.32 and 1.65) agree with C. nigropunctatum (1.30–2.03) and not C. aurae sp. nov. (0.97–1.28) ( Fig. 26 ). Those syntypes also have 27–28 finger-IV + toe-IV digital lamellae, in agreement with modal values in C. nigropunctatum (range, 26– 32) and fewer than in specimens of C. aurae sp. nov. (29–32). Also, one (BMNH 1946.8.15.12) has prefrontal contact, and prefrontals are nearly in contact in the other specimen (BMNH 1946.8.19.78). Prefrontal contact is rare in mabuyines and not present in any C. aurae sp. nov. examined, but it occurs in C. nigropunctatum at a moderate frequency (~20% in those examined). Gray (1831a) described Tiliqua aenea from "Brazils," a name generally applied at that time, without precision, to many areas in eastern and northern South America. Later, Gray (1845) listed specimens (now considered the syntypes) "in spirits" from "W. Indies" and "St. Vincents," and "from the Rev. L. Guilding's collection." Because the BMNH register starts in 1837 ( Underwood 1993), it is not possible and animals on that island ( Howard & Howard 1985). There is no record that he collected on the mainland of South America, but he was visited by colleagues who did collect material in South America ( Howard & Howard 1985), and that may explain the origin of the syntypes of T. aenea . Confusion in localities (West Indies versus South America) has been documented for other BMNH specimens of reptiles, including skinks, collected in the early nineteenth century ( Underwood 1993; Miralles et al. 2009a). We place Tiliqua aenea Gray (1831a) in the synonymy of C. nigropunctatum Spix (1825) . However, given the great genetic diversity of the latter species (see discussion above, in Remarks for Copeoglossum ), it is likely that the name Copeoglossum aeneum will re-emerge in the future as a valid species of South American skink. Miralles et al. (2005; 2006b; 2009; 2009) and Miralles and Carranza (2010) examined specimens designated here as C. aurae sp. nov. but considered them to be members of C. nigropunctatum .

Copeoglossum aurae sp. nov. is broadly sympatric with another new species, Marisora aurulae sp. nov. (described below). They have remarkably similar distributions, with both occurring on St. Vincent, the Grenadines, Grenada, Trinidad, and Tobago. They are similar in appearance in that both lack dark dorsolateral stripes but have dark lateral stripes. It is not known whether they have similar microhabitats, but they have been taken together at two localities (Carriacou Island, Grenadines and Scarborough, Tobago), each on the same date and by the same collector. The first museum specimens of what is here called C. aurae sp. nov. turned up nearly 132 years ago, and those of what is here called Marisora aurulae sp. nov. first appeared about 129 years ago, and old specimens of each are from multiple islands. There is no indication that either species was introduced to the islands or among the islands, and it is unlikely they were a significant food source for native Americans (another possible explanation for distribution among islands). Thus, their current distribution on multiple islands is probably natural, transported by flotsam before the arrival of humans.

Figures

FIGURE 3. Head scalation in Neotropical skinks (Mabuyidae, Mabuyinae). Locations and names of head scales on side (A) and top (B) of head. Abbreviations are AL (anterior loreal), F (frontal), FN (frontonasal), FP (frontoparietal), IP (interparietal), LST (lower secondary temporal), M (mental), NU (nuchal), P (parietal), PF (prefrontal), PL (posterior loreal), PT (primary temporal), R (rostral), UST (upper secondary temporal), and UTT (upper tertiary temporal). Selected (non-standard) measurements on side (C) and top (D) of head. Length of eyelid window, supraciliary-2, narial opening, and supranasal scale defined by longest axis. Other scale length measurements (not shown) follow axis of body. Chin scale configuration: (E) no contact between chin shields and infralabials; (F) two

FIGURE 5 (continued on the next page)

FIGURE 11. Map showing the distribution of mabuyine skinks occurring in the Lesser Antilles and Trinidad and Tobago. See Fig. 1 for location of the islands. The islands and species are: (A) Anguilla, Dog Island, and St. Barts (Spondylurus powelli sp. nov., squares), and St. Martin (S. martinae sp. nov., circle); (B) Guadeloupe: Basse-Terre (Mabuya guadeloupae sp. nov., square), Grande-Terre (Mabuya grandisterrae sp. nov., triangle), Îlet à Cochons (Mabuya cochonae sp. nov., star), La Désirade and Terre de Bas, Petite-Terre (Mabuya desiradae sp. nov., diamonds), and Marie-Galante (Capitellum mariagalantae sp. nov., circle); (C) Martinique (Mabuya mabouya, circles; Capitellum metallicum, triangle); (D) St. Vincent, the Grenadines, and Grenada (Copeoglossum aurae sp. nov., circles; Marisora aurulae sp. nov., triangles); (E) Redonda (Copeoglossum redondae sp. nov., circle) and Montserrat (Mabuya montserratae sp. nov., square); (F) Dominica (Mabuya dominicana); (G) St. Lucia (Alinea luciae), (H) Barbados (Alinea lanceolata); (I) Trinidad (Copeoglossum aurae sp. nov., circles; Marisora aurulae sp. nov., triangle), and (J) Tobago (Copeoglossum aurae sp. nov., circles; Marisora aurulae sp. nov., triangles). Hollow symbols in A and J are literature records (specimens not examined). Where no specific localities within an island are known for a species, a symbol is placed centrally on the island: Alinea luciae, Capitellum mariagalantae, Capitellum metallicum, and Copeoglossum redondae sp. nov. The scale in (A) applies to all islands.

FIGURE 23. Head scalation in species of the Genus Copeoglossum occupying Caribbean islands (left, side view; right, dorsal view). (A) C. aurae sp. nov. (BMNH 90.11.25.16, paratype); (B) C. margaritae sp. nov. (USNM 217141, holotype); and (C) C. redondae sp. nov. (ANSP 9517, holotype).

FIGURE 24. Representative patterns in species of the Genus Copeoglossum occupying Caribbean islands (top, dorsal view; bottom, side view). (A) C. aurae sp. nov.; (B) C. margaritae sp. nov.; and (C) C. redondae sp. nov.

FIGURE 26. Graph of frontonasal width versus frontonasal length in the five species of the Genus Copeoglossum. The syntypes of Tiliqua aenea Gray (= Copeoglossum nigropunctatum) are indicated with arrows.

FIGURE 27. Graph of suture length between supraocular-2 and supraciliaries versus that between supraocular-1 and supraciliaries in four species of the Genus Copeoglossum. Copeoglossum arajara is not included because it has three supraoculars, possibly from fusion between first and second supraoculars.