A revision of the Dwarf Geckos, genus Lygodactylus (Squamata: Gekkonidae) from Angola, with the description of three new species Author Marques, Mariana P. 0000-0002-1712-2632 Research Center in Biodiversity and Genetic Resources (CIBIO), InBIO, University of Porto, Rua Padre Armando Quintas 7, Vairão, 4485 - 661 Porto, Portugal & mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632 & Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Rua da Escola Politécnica, 58, 1269 - 102 Lisboa, Portugal & mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632 & lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978 & Museu de História Natural e da Ciência da Universidade do Porto, Praça Gomes Teixeira 4099 - 002 Porto, Portugal & lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978 & mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632 mpltmarques@gmail.com Author Ceríaco, Luis M. P. 0000-0002-0591-9978 Museu de História Natural e da Ciência da Universidade do Porto, Praça Gomes Teixeira 4099 - 002 Porto, Portugal & lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978 & Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Rua da Escola Politécnica, 58, 1269 - 102 Lisboa, Portugal & mpltmarques @ gmail. com; https: // orcid. org / 0000 - 0002 - 1712 - 2632 & lmceriaco @ mhnc. up. pt; https: // orcid. org / 0000 - 0002 - 0591 - 9978 lmceriaco@mhnc.up.pt Author Buehler, Matthew D. 0000-0001-5460-7770 Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085 - 1699, USA & mbuehle 3 @ gmail. com; https: // orcid. org / 0000 - 0001 - 5460 - 7770 & adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119 & Department of Biological Sciences & Museum of Natural History, Auburn University, Auburn, Alabama, 36849, USA & mbuehle 3 @ gmail. com; https: // orcid. org / 0000 - 0001 - 5460 - 7770 mbuehle3@gmail.com Author Bandei- Ra, Suzana A. 0000-0001-5460-7770 Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085 - 1699, USA & mbuehle 3 @ gmail. com; https: // orcid. org / 0000 - 0001 - 5460 - 7770 & adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119 & Instituto Nacional da Biodiversidade e Áreas de Conservação, Ministério do Ambiente de Angola, Centralidade do Kilamba, Rua 26 de Fevereiro, Quarteirão Nimi Ya Lukemi, edifício Q 11, 3 º andar, Luanda, Angola. & adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119 mbuehle3@gmail.com Author Janota, Joyce M. 0000-0002-7496-9119 Instituto Nacional da Biodiversidade e Áreas de Conservação, Ministério do Ambiente de Angola, Centralidade do Kilamba, Rua 26 de Fevereiro, Quarteirão Nimi Ya Lukemi, edifício Q 11, 3 º andar, Luanda, Angola. & adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119 adez.bandeira@gmail.com Author Bauer, Aaron M. 0000-0001-5460-7770 Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085 - 1699, USA & mbuehle 3 @ gmail. com; https: // orcid. org / 0000 - 0001 - 5460 - 7770 & adez. bandeira @ gmail. com; https: // orcid. org / 0000 - 0002 - 7496 - 9119 mbuehle3@gmail.com text Zootaxa 2020 2020-09-23 4853 3 301 352 journal article 8437 10.11646/zootaxa.4853.3.1 9ed37c70-0d85-41bf-809f-58349df9a4a7 1175-5326 4410727 802D1C38-1186-4187-8289-5AC125242802 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 ) 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). 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 ).