Cyrtodactylus pseudoquadrivirgatus RÖSLER, NGUYEN, VU, NGO & ZIEGLER, 2008
Can you confirm these amateur observations of Cyrtodactylus pseudoquadrivirgatus?
|Higher Taxa||Gekkonidae, Gekkota, Sauria, Squamata (lizards: geckos)|
|Synonym||Cyrtodactylus pseudoquadrivirgatus RÖSLER, NGUYEN, VU, NGO & ZIEGLER 2008|
Cyrtodactylus cf. pseudoquadrivirgatus — JESTRZEMSKI et al. 2013
Cyrtodactylus pseudoquadrivirgatus — NGUYEN et al. 2021
|Distribution||Vietnam (Thua Thien-Hue Province)|
Type locality: A Luoi, Thua Thien-Hue Province, central Vietnam, at an elevation of 800 m above sea level.
|Types||Holotype: ZFMK 83895, adult female, (Fig. 1–2a– d), leg. Vu Ngoc Thanh, 20 July 2005.|
|Diagnosis||Diagnosis. A medium-sized Cyrtodactylus that can be distinguished from all congeners from the Oriental and Australian regions on the basis of the following combination of characters: 1) SVL up to 84 mm (TL > SVL); 2) head depressed, distinct from body, snout with me dial concavity; 3) 37–55 interorbital scales; 4) body rounded in cross-section, belly lat, not wider than head; 5) lateral fold narrow, with enlarged lateral tubercles; 6) tubercles present on dorsum of head, body, forearm, hind limbs and tail; 7) 16–24 rows of dorsal tubercles; 8) 41–57 longitudinal rows of ventral scales at midbody; 9) rounded tail in cross-section, tail base not enlarged; 10) 5–9 precloacal pores in angular series and 1–12 enlarged precloacal scales; 11) absence of enlarged femoral scales, lack of femoral pores and a precloacal groove; 12) absence of transversely enlarged subcaudals; 13) 9–13 lamellae under first digit, 10–13 lamellae under irst toe, and 16–25 lamellae under 4th toe; 14) body dorsum light brown with dark brown pattern (head mottled, dorsum mottled, striped or with bands); 15) neck band medially interrupted; 16) limbs striped or mottled, tail with dark and light bands (Rösler et al. 2008).|
Comparisons.– Cyrtodactylus pseudoquadrivirgatus sp. nov. differs from the Vietnamese species C. badenensis Nguyen, Orlov and Darevsky, 2006, in the absence of transversely enlarged subcaudals, in having higher ventral counts and lower numbers of precloacal pores; from C. condorensis (Smith, 1921) in the absence of transversely enlarged subcaudals and femoral scales, as well as in having higher ventral scale counts; from C. cryptus Heidrich, Rösler, Vu, Böhme and Ziegler, 2007, in having a lower number of precloacal pores, an interrupted nuchal band and narrow dorsal bands or stripes; from C. intermedius in the absence of transversely enlarged subcaudals and femoral scales, as well as in having narrow dorsal bands or stripes; from C. irregularis (Smith, 1921), in lacking enlarged femoral scales and having an interrupted nuchal band; from C. nigriocularis Nguyen, Orlov and Darevsky, 2006, in the absence of transversely enlarged subcaudals and having higher number of precloacal pores and narrow dorsal bands or stripes; from C. paradoxus (Darevsky and Szczerbak, 1997), in the absence of transversely enlarged subcaudals, higher ventral scale counts and a higher number of precloacal pores; from C. phongnhakebangensis Ziegler, Rösler, Herrmann and Vu 2003, in lacking transversely enlarged subcaudals and lower number of precloacal pores and narrow dorsal bands or stripes (after Smith, 1935; Darevsky and Szczerbak, 1997; Ziegler et al., 2002; Nguyen et al., 2005).
C. pseudoquadrivirgatus sp. nov. can be distinguished from the externally similar C. quadrivirgatus Taylor, 1962, by its higher SVL and lack of enlarged femoral scales (see below).
In lacking enlarged subcaudals, C. pseudoquadrivirgatus sp. nov. differs from C. aaroni Günther and Rösler, 2003; C. aequalis Bauer, 2003; C. annandalei Bauer, 2003; C. chanhomeae Bauer, Sumontha and Pauwels, 2003; C. cracens Batuwita and Bahir, 2005; C. consobrinoides (Annandale, 1905); C. consobrinus (Peters, 1871); C. darmandvillei (Weber, 1890); C. deveti (Brongersma, 1948), which has enlarged but medially divided subcaudals; C. elok Dring, 1979; C. edwardtaylori Batuwita and Bahir, 2005; C. feae (Boulenger, 1893); C. fraenatus (Günther, 1864); C. ingeri Hikida, 1990; C. jarujini Ulber, 1993; C. khasiensis (Jerdon, 1870); C. louisiadensis (De Vis, 1892); C. malayanus (De Rooij, 1915); C. malcolmsmithi (Constable, 1949); C. mimikanus (Boulenger, 1914); C. murua Kraus and Allison 2006; C. oldhami (Theobald, 1876); C. peguensis (Boulenger, 1893); C. ramboda Batuwita and Bahir, 2005; C. redimiculus King, 1962; C. russelli Bauer, 2003; C. soba Batuwita and Bahir, 2005; C. subsolanus Batuwita and Bahir, 2005; C. sumonthai Bauer, Pauwels and Chanhome, 2002; C. tuberculatus (Lucas and Frost, 1900); and C. variegatus (Blyth, 1859) (after De Rooij, 1915; Smith, 1935; Taylor, 1963; Dring, 1979; Hikida, 1990; Ulber, 1993; Bauer et al., 2002, 2003; Bauer, 2003; Batuwita and Bahir, 2005; Couper et al., 2006; Rösler et al., 2007).
In lacking a precloacal groove in both sexes, C. pseudoquadrivirgatus sp. nov. differs from C. agamensis (Bleeker, 1860); C. aurensis Grismer, 2005; C. baluensis (Mocquard, 1890); C. cavernicolus Inger and King, 1961; C. marmoratus Gray, 1831; C. papuensis (Brongersma, 1934); C. philippinicus (Steindachner, 1867); C. pulchellus Gray, 1828; C. pubisulcus Inger, 1957; C. rubidus (Blyth, 1861); C. sadleiri Wells and Wellington, 1985; C. semenanjungensis Grismer and Leong, 2005; and C. tiomanensis Das and Lim, 2000 (after Das and Lim, 2000; Grismer, 2005; Rösler et al., 2007; Tikader and Sharma, 1992; Youmans and Grismer, 2006).
In lacking preanofemoral pores, C. pseudoquadrivirgatus sp. nov. differs from C. fumosus (Müller, 1895), which has 42–52 of such pores; C. loriae (Boulenger, 1898), having 30–81; C. novaeguineae (Schlegel, 1837), having 24–34; and C. seribuatensis Youmans and Grismer 2006, having 42–45.
C. pseudoquadrivirgatus sp. nov. can be differentiated in the combination of precloacal and femoral pores from C. agusanensis (Taylor, 1915), that has 7–11 precloacal and 3–14 femoral pores; C. biordinis Brown and McCoy, 1980, having 11–14 precloacal and 15–28 femoral pores in two rows; C. brevipalmatus (Smith, 1923), having 6–7 precloacal and 9–10 femoral pores; C. gubernatoris (Annandale, 1913), having 9 precloacal and 6 femoral pores; C. interdigitalis Ulber, 1993, having 14 precloacal and 8–9 femoral pores; C. slowinskii Bauer, 2002, having 11–12 precloacal and 9–1 femoral pores; C. tigroides Bauer, Sumontha and Pauwels, 2003, having 5–7 precloacal and 8–9 femoral pores; C. wetariensis (Dunn, 1927), having 11–12 precloacal and 12–13 femoral pores (after Annandale, 1913; Dunn, 1927; Rösler et al., 2007; Ulber, 1993; Youmans and Grismer, 2006).
C. pseudoquadrivirgatus sp. nov. differs in showing a larger SVL from C. jellesmae (Boulenger, 1897); C. laevigatus (Darevsky, 1964); and C. thirakhupti Pauwels, Bauer, Sumontha and Chanhome 2004; the new species differs in being smaller (in terms of SVL) from C. sermowaiensis (De Rooij, 1915); from the aforementioned species, C. pseudoquadrivirgatus sp. nov. differs additionally in showing precloacal pores (after Boulenger, 1897; Darevsky, 1964; De Rooij, 1915; Pauwels et al., 2004).
C. pseudoquadrivirgatus sp. nov. differs by its larger SVL (83.3 mm versus 68.5 mm) and a higher number of dorsal tubercle rows (16–24 versus 11) from C. adleri Das, 1997; by its larger SVL (83.3 mm versus 66 mm) and lack of a U-shaped nuchal band from C. angularis (Smith, 1921); by enlarged scales behind the precloacal pores and a medially divided nuchal band from C. annulatus (Taylor, 1915); by a higher ventral scale count (41–57 versus 32–37) and a lower number of precloacal pores (5–9 versus 10–28) from C. ayeyarwadyensis Bauer, 2003; by a smaller SVL (83.3 mm versus 88 mm) and a lower number of dorsal tubercle rows (16–24 versus 27) from C. brevidactylus Bauer, 2002; by a lower number of dorsal tubercle rows (16– 24 versus 25) and more ventrals (41–57 versus 30) from C. buchardi David, Teynié and Ohler 2004; by a larger SVL (83.3 mm versus 79.1 mm) and more ventrals (41–57 versus 37) from C. chrysopylos Bauer, 2003; by a smaller SVL (83.3 mm versus 112 mm) and a lower lamellae number below the fourth toe (16–25 versus 24–26) from C. derongo Brown and Parker, 1973; by a larger SVL (83.3 mm versus 63 mm) and fewer precloacal pores (5–9 versus 16–29) from C. gansi Bauer, 2003; by a smaller SVL (83.3 mm versus 163 mm) and fewer subdigital lamellae below the fourth toe (16–25 versus 28–35) from C. irianjayaensis Rösler, 2000; by a smaller SVL (83.3 mm versus 85 mm), fewer ventral scales (41–57 versus 60–64) and fewer precloacal pores (5–9 versus 13) from C. lateralis (Werner, 1896); by a smaller SVL (83.3 mm versus 104.5 mm) and enlarged scales behind the precloacal pores from C. matsuii Hikida, 1990; by a smaller SVL (83.3 mm versus 93 mm) and more ventral scales (41–57 versus 30–34) from C. papilionoides Ulber and Grossmann, 1991; by a larger SVL (83.3 mm versus 77 mm) and a different dorsal colouration and pattern (buff with blackish brown blotches or stripes versus dark brown above, with small, irregularly-shaped white spots) from C. sworderi (Smith, 1925); by a larger SVL (83.3 mm versus 64 mm) and fewer precloacal pores (5–9 versus 12) from C. wakeorum Bauer, 2003; by a smaller SVL (83.3 mm versus 96.2 mm) and a lower number of subdigital lamellae below the fourth toe (16–25 versus 25–30) from C. yoshii Hikida, 1990 (after Brown and Parker, 1973; Das, 1997; Rösler et al., 2007; Smith, 1925; Ulber and Grossmann, 1991; Youmans and Grismer, 2006).
The new species differs from the species of the subgenus Geckoella sensu Rösler (2000b) in having a larger SVL and in having precloacal pores, which are lacking in C. (Geckoella) albofasciatus (Boulenger, 1885); C. (Geckoella) collegalensis (Beddome, 1870); C. (Geckoella) deccanensis (Günther, 1864); C. (Geckoella) jeyporensis (Beddome, 1877); C. (Geckoella) nebulosus (Beddome, 1870); and C. (Geckoella) yakhuna (Deraniyagala, 1945); the new species differs from C. (Geckoella) triedrus (Günther, 1864) in having a larger SVL (83.3 mm versus 62 mm) and in the absence of femoral pores (after Deraniyagala, 1953; Smith, 1935; Tikader and Sharma, 1992).
Comparisons with Cyrtodactylus irregularis and C. quadrivirgatus.– Based on the character analysis presented above, Cyrtodactylus pseudoquadrivirgatus sp. nov. can be distinguished from most Cyrtodactylus species by the combination of size and scalation features. Congeners that are more difficult to separate from C. pseudoquadrivirgatus sp. nov. are treated in more detail in the following para, including the Vietnamese species C. irregularis and C. quadrivirgatus from Thailand and Malaysia.
Cyrtodactylus irregularis was described as a subspecies of Gymnodactylus peguensis (currently C. peguensis) by Smith (1921) from the central Vietnamese type locality, “Camly, 3500 feet, Langbian Plateau, S. Annam”. The holotype measured 79 mm SVL and the paratype 80 mm SVL. Characteristic features include small, irregular blotches, in combination with 8–9 supralabials, 8–9 sublabials and 5–7 precloacal pores (the tail was lacking). Subsequently, irregularis was treated by Smith (1935) as a full species, and the following features were listed: 11 supralabials; 9 sublabials; body and limbs with small granular scales, and larger conical or subtriangular tubercles; lateral folds with distinctly enlarged scales; 41–46 ventral scale rows; 5–7 precloacal pores in males, arranged in angular series; a group of enlarged precloacal scales posterior to the precloacal pores; 7 or 8 enlarged femoral scales, separated from precloacal pores by small, smooth scales; femoral scales are pitted, and in one specimen one scale perforated; tail scales not known. A third specimen was provisionally allocated by Smith (1935: BMNH 19126.96.36.199) to C. irregularis, because of a somewhat different scalation (i.e., 33 ventral scales) and an unusual pattern (i.e., nuchal band broken in the middle). The tail of this additional specimen was described as complete and “covered with small, flat, juxtaposed scales and series of enlarged tubercles above, with larger imbricate scales below, those on the median line being larger than the other”. According to Darevsky and Szczerbak (1997), C. irregularis is the sole Vietnamese species lacking transversely enlarged subcaudal scales: “Underside of tail with numerous roundish scales” (see also Youmans and Grismer, 2006).
Cyrtodactylus irregularis arguably represents the least known Vietnamese bent-toed gecko, as further records or specimens are not known. Another Vietnamese species with small, imbricate scales was described recently as C. cryptus (Heidrich et al., 2007). Based on the features known from the irregularis type series and from the specimen “Brit. Mus. 19188.8.131.52”, Ziegler et al. (2004) published another report of C. irregularis, an adult female (ZFMK 80080) from Bach Ma, Thua Thien-Hue, Vietnam. They noted that the pattern of the specimen was somewhat different from the original description, although the neck and back pattern corresponded well with the specimen “Brit. Mus. 19184.108.40.206”, tentatively identified as C. irregularis by Smith (1935). In addition, Ziegler et al. (2004) recognized, that the scalation features of ZFMK 80080 “largely correspond with the characters given in Smith (1935), except for the lack of enlarged femoral scales”, and concluded “Apparently, Cyrtodactylus irregularis is a rarely collected and therefore (is a) barely known benttoed gecko species”. With the series of Cyrtodactylus now available from central Vietnam, it is obvious, that the variations recognized by Ziegler et al. (2004) for their single specimen (which include, among others, the lack of enlarged femoral scales and the variation shown in the dorsum pattern) constitute diagnostic features of C. pseudoquadrivirgatus sp. nov. The specimen ZFMK 80080 (C. irregularis sensu Ziegler et al., 2004), thus, is conspecific with C. pseudoquadrivirgatus sp. nov. However, due to the poor state of preservation of this specimen, we refrain from adding it to the type series.
Cyrtodactylus quadrivirgatus, which is phenetically similar to C. pseudoquadrivirgatus sp. nov., was described by Taylor (1962) based on an adult female holotype No. 387, from Khao Chong Forest Experiment Station, Trang Province, Thailand. Besides a topotypic paratype, the paratype series also contained specimens from several locations in Malaysia (see also Taylor, 1963). C. quadrivirgatus is known to occur from southern Thailand through Malaysia, Singapore to northern Sumatra (Manthey and Grossmann, 1997). With a maximum SVL of 83.3 mm, C. pseudoquadrivirgatus sp. nov. grows somewhat larger than C. quadrivirgatus which has a maximum SVL of 71 mm (Taylor, 1962; Manthey and Grossmann, 1997; Youmans and Grismer, 2006). A faint ventrolateral fold exists in both C. pseudoquadrivirgatus sp. nov. and C. quadrivirgatus (Taylor, 1962), although the lateral fold tubercles are more strongly developed in the new species. According to Taylor (1962), caudal tubercles are absent in C. quadrivirgatus (see also Manthey and Grossmann, 1997). However, all five Malaysian C. quadrivirgatus studied by us (CPHR 2259–60, 2262, SAMA R36783, ZFMK 86751) showed tubercles in the first tail whorl and two of them also in the fifth caudal whorl, corresponding to the data provided by Youmans and Grismer (2006). In scalation, C. pseudoquadrivirgatus sp. nov. and C. quadrivirgatus differ significantly (p < 0.001) in the number of granular scales behind upper labials to angle of mouth; internasals; scales between fifth supralabials; gular scales bordering postmentals; ventral scales; enlarged femoral scales and precloacal pores in males, as well as precloacal scales in females (see Table 3). However, the size of the femoral scales serves best for a diagnosis. Whereas in the femoral region of C. pseudoquadrivirgatus sp. nov., the anteriorly largest scales gradually pass over to the posteriorly smallest scales in both sexes, there exists a sharp separation in between these scales in C. quadrivirgatus (Figs. 2e–f). In the latter species, independent of sex, the large (femoral) scales are ca. x 3 larger than small posterior scales. Counts of 17–25 single-sided femoral scales are known for C. quadrivirgatus (Taylor, 1962; Manthey and Grossmann, 1997); the specimens studied by us have 4–21 unilateral femoral scales. Whereas the male precloacal pores are relatively large and easily discernible in C. pseudoquadrivirgatus sp. nov., the pores were hardly detectable in male specimens of C. quadrivirgatus, CPHR 2262 and ZFMK 86751. Also, the structured epithelial layer of the small pores in female C. pseudoquadrivirgatus sp. nov. was usually clearly discernible, whereas such depressions were hardly recognizable in the scales of the female C. quadrivirgatus, specimen CPHR 2259–60. However, such a difference could also be related to reproductive phase. Taylor (1962), Manthey and Grossmann (1997) and Youmans and Grismer (2006) mention 0–4 precloacal pores; the C. quadrivirgatus males studied by us show 3 and 6 precloacal pores.
Cyrtodactylus pseudoquadrivirgatus sp. nov. and C. quadrivirgatus show a similar hemipenes morphology (Fig. 4): in both species, the sulcus spermaticus is a deep groove, bordered by bulging lips; on the pedicel, the sulcus runs nearly crossways, to turn in an angle of maximum 90° in the truncus region; in the upper truncus region, the sulcus bifurcates beneath a transverselly raised skin bulge; both arched sulcus branches stretch towards concentric depressions of the apical lobes; in C. quadrivirgatus, these lobes bear serrated skin rims, that extend towards the concentric depressions; in C. pseudoquadrivirgatus sp. nov. (IEBR 2264), these apical lobes are not fully everted, but show a similar structure. However, the serrated apical lobes are medially separated by a smooth area in C. quadrivirgatus, which is not the case in C. pseudoquadrivirgatus sp. nov. In addition, C. quadrivirgatus and C. pseudoquadrivirgatus sp. nov. distinctly differ from C. phongnhakebangensis by having groove-shaped apical ornamentation, whereas the latter species shows a typical apical calyx-ornamentation, that also can be found in C. aaroni and C. mimikanus (Ziegler et al., 2002; Günther and Rösler, 2003).
Finally, C. pseudoquadrivirgatus sp. nov. and C. quadrivirgatus show an unusual dorsal pattern variability. In both species, the pattern on dorsum may consist of stripes, bands or blotches, and an uninterrupted nuchal band is lacking (Manthey and Grossmann, 1997; Chan-ard et al., 1999). However, a distinct difference can be observed in the flank pattern: whereas the flanks beneath the dorsolateral stripes and/or bands are typically unicoloured and covered with light blotches in C. quadrivirgatus, C. pseudoquadrivirgatus sp. nov. typically bears distinct dark stripes or a longitudinal series of blotches on a light ground colouration (Rösler et al. 2008).
|Etymology||The name refers to Cyrtodactylus quadrivirgatus, with which the new species may be easily confused with due to the striking similarities in external morphology, including scalation, colouration and pattern; the prefix pseudo derives from the Greek pseudes = false, pretended.|