Lygodactylus angolensis Bocage, 1896

Marques, Mariana P., Ceríaco, Luis M. P., Buehler, Matthew D., Bandei- Ra, Suzana A., Janota, Joyce M. & Bauer, Aaron M., 2020, A revision of the Dwarf Geckos, genus Lygodactylus (Squamata: Gekkonidae) from Angola, with the description of three new species, Zootaxa 4853 (3), pp. 301-352 : 322-324

Lygodactylus angolensis Bocage, 1896

( Tables 4; Fig. 1–11 )

Lygodactylus angolensis ( Bocage 1896: 110, 1897: 193; Monard 1937: 52; Loveridge 1947: 207; 1957: 187; Hellmich 1957: 35; Pasteur 1965 [1964]: 56; Branch 1998: 245; Broadley & Cotterill 2004: 41; Ceríaco et al. 2016a: 57, 2016b: 56, 57, 2018b: 413, 421, 422; Marques et al. 2018: 188,189; Branch et al. 2019: 315)

Lygodactylus laurae ( Schmidt 1933: 4; Mertens 1937: 6; Barbour & Loveridge 1946: 147; Marx 1959: 464; McCoy & Richmond 1966: 154)

Lygodactylus capensis [part] ( Bocage 1895: 15)

Despite being a widespread species in Angola and surrounding countries, the species was only described at the end of the nineteenth century based on two specimens collected by the Portuguese explorer José d’Anchieta “on the walls of a corral” in Hanha, Benguela Province ( Bocage 1896). Bocage (1896) noted that some specimens from Cahata and Galanga, both in Benguela Province, that he had previously ( Bocage 1895) identified as L. capensis were in fact the newly described species as well. The main characters used by Bocage to diagnose the newly described species against L. capensis were the number of perinasal scales (= scales entering the nostril) (four in L. angolensis versus three in L. capensis ), the number of supralabials (eight to nine versus six to eight in L. capensis ) and infralabials (seven to eight versus six to seven in L. capensis ) and the number of preanal pores in males (nine versus four to seven in L. capensis ). In the type catalog of the Lisbon Museum, Bocage (1897) mistakenly considered both the specimens from Hanha and those from Cahata and Galanga as types of L. angolensis . Monard (1937) reported a putative specimen of L. angolensis from Ebanga, but the number of perinasal scales was, in the opinion of the author, different from the typical L. angolensis . Sternfeld (1911) described Lygodactylus fischeri (non Boulenger 1890) from Mozambique, a name that was later synonymized with L. angolensis by Loveridge (1947), which was the first record of the species out of Angola. Other records from Tanzania, Zambia, and South Africa (in error) were later provided by Loveridge (1933) and Pitman (1934), respectively. Schmidt (1933), based on the material collected by the Pulitzer Angolan Expedition in Chitau, Bié Province, central Angola, described Lygodactylus laurae ( Fig. 5 ). This species was also synonymized with L. angolensis by Loveridge (1947), and recent collection of new specimens very close to Chitau ( MHNC-UP /REP 495–498; CAS 266003–266009) revealed that the only species present in the region corresponds to L. angolensis . Loveridge (1947) provided several additional records for Angola and Hellmich (1957) reported three specimens from Entre Rios, Benguela Province. There are extensive records of the species distribution outside Angola (see, for example, Pasteur 1965 [1964] and Branch 1998 for estimates of the distribution of the species), but no precise locality records have been published from Angola since the late 1950s. The first new records of the species were based on specimens collected in Cangandala National Park, Malanje Province, by Ceríaco et al. (2016b; 2018b). Other recent records are from other areas of Malanje (Laúca) and Bié (14 km by dirt road from Cassumbi village) provinces (see Specimens examined below). Marques et al. (2018) considered Laurent’s (1964) records of L. capensis as L. angolensis , but subsequent examination of these specimens, which are currently housed in the Museu do Dundo, do not support this synonymization (see account for Lygodactylus tchokwe sp. nov. below).

Diagnosis. Lygodactylus angolensis is small dwarf gecko with a maximum SVL of 34 mm. It can be distinguished from other members of the genus occurring in Angola and the surrounding regions by having 1) three scales surrounding the nostril (one supranasal, 1 st supralabial, one postnasal); 2) nostrils separated from the rostral and pierced above the suture between rostral and first supralabial; 3) mental followed by three postmentals; 4) first infralabial <25% overlap with postpostmental; 5) 19 to 21 rows of ventral scales; 6) between 7 and 10 precloacal pores in males; 7) median subcaudals arranged in an irregular pattern of one row of scales about ⅓ width of the tail alternating with two rows (sometimes just one) of scales of about ½ the width of the tail ( Fig. 4A ); 8) venter cream; 9) without markings on the gular region. The background coloration is usually light grey or light olive-brown. Some individuals have a dark stripe from the snout, through the eye, ending near the forearm insertion. The upper flank is marked with a series of pale spots forming a pale light line. These spots are usually edged with a dark mark above and below sometimes yielding the appearance of an ocellus ( Figs. 6–7 ).

Figures

FIGURE 1. Maximum likelihood phylogeny of Lygodactylus using ND2. Nodes supported by ML and BI are indicated by black dots. nodes only supported by the ML tree are indicated with half-filled black dots. nodes only supported by BI are indicated by half-filled red dots. and nodes with low support in both the ML and BI trees have red dots. Species of Lygodactylus occurring in Angola have a star next to them in the tree. Species within the Madgascan Lygodactylus clade include L. tolampyae, L. heterurus, L. arnoulti. Species in the Greater Drakensberg group include; L. nigropunctatus, L. graniticolus, L. ocellatus, L. soutpansbergensis. L. incognitus, L. montiscaeruli, L. waterbergensis, and L. methueni. Note that this tree should be used only as a guide to shallow relationships in the genus and to indicate relative divergence.

FIGURE 2. Comparison of the number of precloacal pores in Angolan individuals of Lygodactylus angolensis, L. baptistai sp. nov., L. nyaneka sp. nov., L. lawrencei and L. tchokwe sp. nov.

FIGURE 3. Comparison of the nostrils position inAngolan individuals of:A. L. angolensis (AMB 9216); B. Lygodactylus baptistai sp. nov. (CAS 263557); C. L. nyaneka sp. nov. (UF 187229); D. L. lawrencei (CAS 264259); E. L. tchokwe sp. nov. (MD 5312b).

FIGURE 4. Comparison of the tail scales arrangement in Angolan individuals of A. Lygodactylus angolensis (MHNC-UP/REP 533); B. L. nyaneka sp. nov. (CAS 263531); C. L. baptistai sp. nov. (CAS 263557); D. L. lawrencei (CAS 264259); E. L. tchokwe sp. nov. (MD 5312d).

FIGURE 5. Dorsal and ventral whole-body views (left) and dorsal. lateral and ventral views of the head (right) of the holotype of Lygodactylys laureae (CM 5860).

FIGURE 6. Dorsal and ventral whole-body views (left) and dorsal. lateral and ventral views of the head (right) of Lygodactylys angolensis (MHNC-UP/REP 495).

FIGURE 7. Lygodactylus angolensis in life (MHNC-UP/REP 495). Base camp surroundings in a banana plantation [-11.08391º, 16.66367º, 1240 m a.s.l.]. 14 km by dirt road from Cassumbi. Bié Province. Angola Plateau. Photo by Luis M. P. Ceríaco.

FIGURE 8. Distribution of Lygodactylus angolensis in the country. White star represents L. angolensis type locality (Hanha. Benguela Province). dark star represents L. laurae type locality (Chitau. Bié Province).

FIGURE 9. Habitats of Lygodactylus angolensis. Top: Human structures and banana platantion in Cassumbi. Bié Province. Below: Miombo woodlands. Cangandala National Park. Malanje Province. Photos by Luis M. P. Ceríaco.

FIGURE 10. Dorsal and ventral whole-body views (left) and dorsal. lateral and ventral views of the head (right) of the holotype of Lygodactylys baptistai sp. nov. (CAS 263557).

FIGURE 11. Lygodactylus baptistai sp. nov. in life (Holotype CAS 263557). Mamué. Serra da Neve. southwestern Angola Photos by Ishan Agarwal.