See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/303746834 Rediscovery and a new record of Hemidactylus laevis (Reptilia: Gekkonidae) from Somaliland, with notes on and resurrection of Hemidactylus fragilis Article in Zootaxa · June 2016 DOI: 10.11646/zootaxa.4117.4.5 CITATIONS READS 4 904 3 authors: Tomas Mazuch 49 PUBLICATIONS 268 CITATIONS Jiri Smid National Museum, Prague, Czech Republic 97 PUBLICATIONS 450 CITATIONS SEE PROFILE SEE PROFILE A. M. Bauer Villanova University 671 PUBLICATIONS 11,827 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: Los reptiles de Arabia como modelo para investigar el origen y mantenimiento de la biodiversidad en zonas aridas View project Integrative taxonomy of gecko fauna (Gekkonidae) in Sri Lanka View project All content following this page was uploaded by Tomas Mazuch on 03 June 2016. The user has requested enhancement of the downloaded file. Zootaxa 4117 (4): 529–542 http://www.mapress.com/j/zt/ Copyright © 2016 Magnolia Press Article ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) http://doi.org/10.11646/zootaxa.4117.4.5 http://zoobank.org/urn:lsid:zoobank.org:pub:B05704F2-CF95-4724-9398-2D8F6B01A09B Rediscovery and a new record of Hemidactylus laevis (Reptilia: Gekkonidae) from Somaliland, with notes on and resurrection of Hemidactylus fragilis TOMÁŠ MAZUCH1,4, JIŘÍ ŠMÍD2 & AARON M. BAUER3 1 Dříteč 65, 53305, Czech Republic Department of Zoology, National Museum, Cirkusová 1740, Prague, Czech Republic 3 Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA 4 Corresponding author. E-mail: [email protected] 2 Abstract Hemidactylus laevis, a gekkonid lizard originally described from the Gaan Libah Mountains in Somaliland, is recorded herein from Erigavo, Sanaag Region, in the same state, which represents the second-ever known specimen and locality for this species in more than 100 years since its description. The species is endemic to Somaliland. Data on morphology and natural history, as well as the first life photographs are provided. Based on morphological data we conclude the species to be closely related to H. laticaudatus. Additionally, H. fragilis from southern Somalia and extreme south-east Ethiopia is resurrected from the synonymy of H. frenatus based on morphological data. Key words: Distribution, Ethiopia, Hemidactylus frenatus, Hemidactylus laticaudatus, Horn of Africa, Somalia Introduction The Horn of Africa, comprising Somalia, Somaliland (a self-declared independent state internationally recognized as an autonomous region of Somalia), Ethiopia, Eritrea, Djibouti and Kenya, is one of the main diversity centres of the genus Hemidactylus Oken. Currently about 39 species are known from the region, of which most are endemic (Parker 1942; Loveridge 1947; Lanza 1990; Spawls et al. 2002; Largen and Spawls 2006; Šmíd et al. 2013, 2014, 2015; Malonza and Bauer 2014; Uetz 2015). There are two major ecological groups of Hemidactylus that can be recognized in the Horn of Africa: i) small terrestrial species with only slightly developed subdigital lamellae that shelter under rocks or logs (e.g., H. albopunctatus Loveridge, H. citernii Boulenger, H. isolepis Boulenger, H. modestus Günther, H. ophiolepis Boulenger, H. puccionii Calabresi, H. ruspolii Boulenger, H. somalicus Parker, H. squamulatus Tornier, H. tropidolepis Mocquard) and ii) large arboreal geckos with well developed subdigital lamellae (e.g., H. angulatus Hallowell, H. arnoldi Lanza, H. barodanus Boulenger, H. flaviviridis Rüppell, H. frenatus Duméril and Bibron, H. laevis Boulenger, H. laticaudatus Andersson, H. mabouia Moreau de Jonnès, H. macropholis Boulenger, H. platycephalus Peters, H. robustus Heyden, H. yerburii pauciporosus Lanza). In spite of this great diversity, only two species are known exclusively from montane habitats above 1600 m elevation—H. laevis from Somaliland and H. laticaudatus from Ethiopia. Hemidactylus laevis was described by Boulenger (1901) on the basis of a single female specimen with a snoutvent length of 39 mm and collected at Gan Lebar [= Gaan Libah; fide Parker 1942], Somaliland, at 1770 m of elevation. The Gaan Libah is one of the Goolis Mountains ranges situated in a mountainous belt that extends from eastern Ethiopia to northeastern Somalia (Fig. 1). It is covered by a relict Juniperus forest that is surrounded by a narrow evergreen zone defined as ‘montane evergreen thicket and scrub’ and is dominated by Buxus (Buxaceae), Cadia (Fabaceae), Dodonaea (Sapindaceae) and Euclea schimperi (Ebenaceae) (Pichi-Sermolli 1957; Hemming 1966). Although later Boulenger (1912) also listed H. laevis also from southern Ethiopia (Dolo) and Somalia (‘Rahanuin country’), the records were found to be misidentified H. fragilis Calabresi (Parker 1942). Except for the original morphological description virtually nothing is known about this species. Accepted by S. Carranza: 19 Apr. 2016; published: 2 Jun. 2016 529 Hemidactylus laticaudatus was described by Andersson (1910) from ‘Harrar, Abyssinia [=Harar, Ethiopia]’ from elevations about 1800–2000 m a.s.l. It has subsequently been recorded from Gondar and its vicinity, Axum, Adwa, Lalibela and the vicinity of Neghelli from elevations of 1600–2500 m (Calabresi 1925; Largen and Spawls 2006; Mazuch unpubl.; Fig. 1). The species is confined to montane grassland habitats on the central plateau of Ethiopia and adjacent Eritrea. Hemidactylus fossatii Scortecci, a doubtful taxon from Saganeiti, Eritrea, has variously been considered a synonym (Largen 1997; Largen and Spawls 2006, 2010) or a subspecies of H. laticaudatus (Loveridge 1947; Wermuth 1965; Rösler 2000; Kluge 2001; De Lisle et al. 2013). For the purpose of this publication, and until a revision of this group is completed, we here regard both taxa under the name H. laticaudatus. Unlike the two high-altitude species, H. fragilis is known from the lowlands of Somalia and Ethiopia. Its type locality, ‘Bur Meldac [=Meel Daaq]’, lies at ca. 219 m elevation. The only other records are from Dolo, Ethiopia, and Rahanuin Region, Somalia, and were reported by Boulenger (1912) and later redetermined by Parker (1942). The species was synonymized with H. frenatus by Loveridge (1947) and most authors treated it as such (e.g., Loveridge 1957; Wermuth 1965; Bauer 1994; Rösler 2000; Kluge 2001; De Lisle et al. 2013; Uetz 2015). Only Lanza (1983, 1990) regarded it as a valid species. In this paper we describe a second specimen of H. laevis from a new locality and discuss the morphology of the species. Based on morphological data we also clarify the taxonomic status of H. fragilis. Material and methods Field survey. A herpetological survey was carried out around Hotel Sanaag in the city of Erigavo (10.622°N, 47.367°E, 1786 m a.s.l.; Fig. 2D,E), Sanaag Region, Somaliland, on 21–22 January 2015 and at Gaan Libah (9.869°N, 44.834°E, 1719 m a.s.l.), Togdheer Region, Somaliland, on 19–20 January 2015. One specimen of H. laevis was collected and one observed in Erigavo, while three hours of night searching (19h00–22h00) at Gaan Libah yielded no additional records. Morphological characters. The new voucher of H. laevis was compared with the holotype of H. laevis and with other Hemidactylus species occurring in the Horn of Africa. Also, we examined the holotype and the two other specimens previously determined as H. fragilis (Parker 1942) and twelve specimens of H. frenatus from the study area and other regions. All studied material is deposited in the following institutions: NHMUK (formerly BMNH)—Natural History Museum, London, UK; MCZ—Museum of Comparative Zoology, Cambridge, Massachusetts, USA; MSNM—Museo Civico di Storia Naturale, Milan, Italy; MSNG—Museo Civico di Storia Naturale ‘Giacomo Doria’, Genova, Italy; MZUF—Museo di Storia Naturale ‘La Specola’, Florence, Italy; NMP—National Museum Prague, Czech Republic; NMW—Naturhistorisches Museum, Wien, Austria; TMHC— Tomáš Mazuch Herpetological Collection, Dříteč, Czech Republic. High-resolution photographs of the target taxa were deposited and are available for download at MorphoBank (Project 2139; http://www.morphobank.org; a total of 276 media files). A list of all examined specimens is given in Appendix 1. Metric data were recorded using Filetta digital callipers to the nearest 0.1 mm. Meristic data were counted using a dissecting microscope. The following features were recorded: total length (ToL); snout-vent length (SVL), measured from tip of snout to vent; head length (HL), measured from tip of snout to retroarticular process of jaw; head width (HW), measured at the widest part of the head; head depth (HD), maximum depth of head; left eye diameter (E), measured horizontally; maximum tympanum diameter (EAR); axilla to groin distance (AG), measured from posterior insertion of front limb to anterior insertion of hind limb; snout to eye distance (SnE), measured between anteriormost point of eye and snout; eye to tympanum distance (EyEa); distance from posterior part of eye to anterior part of tympanum; maximum tympanum diameter (EAR); tail length (TL), measured from vent to tip of original tail; tail width at base (TailBase), measured at the slimmest point; tail width at the level of 2nd whorl (TailWidth); length of largest tubercle of the trunk (TubL); number of supra- and infralabials (left/right); number of interoculars, counted at the mid-point of eyes; contact of uppermost nasals; number of infralabials in contact with anterior postmentals; position of anterior postmentals; number of longitudinal rows of enlarged dorsal tubercles; number of ventrals contained in the eye diameter at midbody (VentEye), counted transversely; number of lamellae under all digits of both fore- and hind limbs, including unpaired proximal ones (presented as: undivided + divided + entire apical). 530 · Zootaxa 4117 (4) © 2016 Magnolia Press MAZUCH ET AL. FIGURE 1. (A) Map of Somaliland and the adjoining part of Somalia showing the distribution of Hemidactylus laevis (red circle—type locality, Gaan Libah; blue circle—new record, Erigavo); (B) Map of the Horn of Africa showing the distribution of H. laticaudatus (green squares)—1: Harar (type locality); 2: Lalibela; 3: Gondar, Azezo, Fiume Caha; 4: Axum; 5: Adwa; 6: 32 km E of Neghelli; 7: Asmara; 8: Saganeiti (type locality of H. fossatii) and H. fragilis (yellow triangles)—1: Bur Meldac (type locality); 2: “Rahanuin; 3: Dolo. Records of H. laticaudatus are from Scortecci (1928), Largen (1997), Largen & Spawls (2006, 2010), and own unpublished data. Results The new record of Hemidactylus laevis Boulenger, 1901 An adult specimen of approximately 80 mm total length with completely regenerated tail was observed on 21 REDISCOVERY OF HEMIDACTYLUS LAEVIS Zootaxa 4117 (4) © 2016 Magnolia Press · 531 January 2015 at 18h30 (Fig. 2C). A subadult female was collected on 22 January 2015 at 20h15 (NMP 75148, MorphoBank M374513–M374561; Figs. 2A,B and 3. Detailed description of the new vouchered specimen follows. Description. Head rather depressed (HL/HD = 2.2); snout subacuminate, forehead slightly concave, largest scales on snout in area between eye and nares; ear opening small and round; nostril in contact with rostral, first supralabial, one enlarged internasal (post-rostral) and two postnasals; two smaller internasal scales, longitudinally arranged, between post-rostrals; small interstitial scale in upper part of rostral scale; rostrum broader than high; supralabials 10/10; infralabials 9/9; mental medium-sized, triangular; two pairs of postmentals, anterior pair larger and in broad contact along median line, posterior pair not touching each other and separated by three gulars; anterior postmentals in narrow contact with second infralabials; interorbitals 30; scales on head small, granular; those on posterior part of the head with small, round, smooth, scattered tubercles, smaller than those on the body; back covered with small juxtaposed granules intermixed with a numerous small, round, flat smooth tubercles in 10–12 regular longitudinal rows (6–8 between the forelimbs, 8 between hindlimbs); upper flanks only covered by these enlarged tubercles, tubercles absent ventrolaterally; ventral scales large, flat, smooth, imbricate, approximately 7 contained in eye diameter at the midbelly; trunk moderately depressed; body and limbs rather short; neck short and rather indistinct; forelimbs and postero-dorsal side of thigh with imbricate granules without enlarged tubercles; dorsal side of tibia with scattered large tubercles; digits free with short distal joints, strongly dilated; lamellae beneath the toes from 1st to 5th: 4 (1+2+1), 6 (1+4+1), 7 (1+5+1), 7 (1+5+1), right 7 (3+3+1) / left 8 (3+4+1); beneath the fingers: right 5 (2+2+1) / left 5 (3+1+1), 5 (1+3+1), 6 (1+4+1), right 6 (0+5+1) / left 6 (1+4+1), 6 (2+3+1); original tail 1.11 times longer than SVL; constricted at its base, slightly swollen, verticillate and carrot-shaped; dorsally and laterally covered with small granular juxtaposed scales arranged in whorls around the tail; about 6–7 transverse rows in a whorl; below with imbricate scales, median series transversely enlarged in two rows per whorl. Measurements (in mm). ToL 68.6; SVL 32.5; AG 13.4; TL 36.1; HL 10.4; HW 6.7; HD 4.6; EyEa 3.3; SnE 4.1; E 2.3; EAR 0.4; TubL 0.4; TailBase 3.5; TailWidth 4.2. Coloration in life. (Fig. 2A,B). Pale greyish brown above with four irregular transverse bands with sinuous margins; 4–5 dull spots on flanks; W-shaped dark marking on occiput and two dark spots just behind eyes; a pair of dark longitudinal markings over the sacrum; a dark stripe from the nostril, through the eye, becoming more diffuse posteriorly to the level of the ear and confluent with the first body transverse band; a dark spot on posterior part of snout; tail yellowish dorsally, with 9 dull dark bands; underside uniform whitish, tail yellowish. Natural history. The specimens were found to be active at night on walls of buildings. The unvouchered adult was very wary and almost immediately hid in the roof of the building (Fig. 2C). The temperature was 17.5 °C and humidity 22 %. The city receives on average 300–400 mm of precipitation annually (Hemming 1966; Muchiri 2007). The vegetation of the Erigavo town is rather poor with scattered trees typical of degraded towns of East Africa (Eucalyptus sp. (Myrtaceae), Schinus molle (Anacardiaceae), Acacia etbaica (Fabaceae); Fig. 2E). No congeners were observed to live syntopically with H. laevis although our survey possibilities were limited due to security reasons. Comparisons. The new specimen differs in only a few characters from the holotype (condition in parentheses): (1) anterior postmentals in contact with second supralabials (not in contact); (2) anterior postmentals in wide mutual contact (not in contact, separated by one granule; Fig. 4E); (3) higher number of labials; (4) lower number of digital lamellae. For a detailed morphological comparison of the two specimens see Table 1. Hemidactylus laevis may be distinguished from all other species of the region by the following combination of features: (1) dorsum, nape and posterior part of head covered with small granules, intermixed with enlarged flat smooth tubercles; (2) swollen tail; (3) tail strongly verticillate, covered by small granular scales without enlarged tubercles. More specifically, below follows a list of characters by which H. laevis can be distinguished from other species from the Horn of Africa. Hemidactylus laticaudatus is larger (SVL up to 62 mm), has more depressed habitus, enlarged tubercles on posterodorsal side of thighs, large tubercles on tail, and more pointed head with more sunken eyes. Hemidactylus albopunctatus, H. curlei Parker, H. flaviviridis, H. funaiolii Lanza, H. isolepis, H. klauberi Scortecci, H. megalops Parker, H. modestus, H. ophiolepis and H. ophiolepoides Lanza have homogenous dorsal scales and weakly developed subdigital lamellae. Hemidactylus barbouri Loveridge, H. squamulatus and H. tropidolepis have heterogeneous dorsal scales without tubercles, some or all scales are keeled. Hemidactylus angulatus, H. arnoldi, H. awashensis Šmíd, Moravec, Kratochvíl, Nasher, Mazuch, Gvoždík and Carranza, 532 · Zootaxa 4117 (4) © 2016 Magnolia Press MAZUCH ET AL. FIGURE 2. (A) Live female of Hemidactylus laevis (NMP 75148) from Erigavo, Somaliland; (B) detail of the head of the same specimen; (C) unvouchered adult specimen from the same locality; (D) habitat of both specimens (Hotel Sanaag); (E) immediate surroundings of the Hotel Sanaag; (F) typical landscape around Erigavo (5 km NW of the town); (G) Daalloh forest in Cal Madow Mountains, supposed primary habitat of H. laevis. This type of montane forest (ca. 1600–2100 m a.s.l.) is very similar to the forest from upper parts of Gaan Libah (ca. 1550–1700 m a.s.l.), the type locality of H. laevis. REDISCOVERY OF HEMIDACTYLUS LAEVIS Zootaxa 4117 (4) © 2016 Magnolia Press · 533 FIGURE 3. The collected specimen of Hemidactylus laevis (NMP 75148). General body habitus—dorsal (A), ventral (B) and lateral view (C); dorsal (D), ventral (E), and lateral (F, G) view of the head; (H) dorsal part of body; (I) lateral view of the tail, note the absence of enlarged tubercles; (J) left hand; (K) right hand; (L) left foot; (M) right foot. Scale refers to the general body habitus pictures only. 534 · Zootaxa 4117 (4) © 2016 Magnolia Press MAZUCH ET AL. FIGURE 4. The holotype of Hemidactylus laevis (BMNH 1946.8.25.42). General body habitus—dorsal (A), ventral (B) and lateral view (C); dorsal (D), ventral (E), and lateral (F, G) view of the head; (H) dorsal part of body; dorsal (I) and ventral (J) side of regenerated tail; (K) right foot. Scale refers to the general body habitus pictures only. REDISCOVERY OF HEMIDACTYLUS LAEVIS Zootaxa 4117 (4) © 2016 Magnolia Press · 535 H. barbierii Sindaco, Razzetti, Ziliani, Wasonga, Carugati and Fasola, H. bavazzanoi Lanza, H. citernii, H. granchii Lanza, H. macropholis, H. ruspolii, H. sinaitus Boulenger, H. taylori Parker and H. yerburii pauciporosus have large, strongly keeled dorsal tubercles. Hemidactylus somalicus and H. puccionii have tail without basal constriction, subdigital lamellae very small proximally, scarcely larger than the granules on the sole, and first supralabial excluded from nostril. Hemidactylus barodanus and H. jubensis Boulenger are larger (SVL up to 70–78 mm), have enlarged tubercles on dorsum, posterodorsal side of thighs and on tail. Hemidactylus robustus has enlarged tubercles on the posterodorsal side of thighs and on tail, dorsum with 13–18 longitudinal rows of round, weakly keeled and posteriorly pointed tubercles, higher number of subdigital lamellae, and different coloration. Hemidactylus mabouia, H. mercatorius Gray, H. mrimaensis Malonza and Bauer, H. platycephalus and H. smithi Boulenger have conical tail without basal constriction and with enlarged keeled tubercles. Hemidactylus frenatus and H. fragilis have conical tail without basal constriction and with enlarged keeled tubercles (Fig. 5E), enlarged, smooth, and flat tubercles usually only on posterior part of body, or at least not extending onto nape (H. frenatus); H. frenatus is usually longitudinally striped or patternless, never transversely banded. TABLE 1. Comparison of metric and meristic characters of the two known individuals of H. laevis. Metric data for the holotype after Boulenger (1901). BMNH 1946.8.25.42 Holotype, ♀ NMP 75148 ♀ SVL (mm) 39 32.5 Head length (mm) 12 10.4 Head width (mm) 8 6.7 Supralabials 9/10 10/10 Infralabials 8/8 9/9 Interorbitals 32 30 Median nasals in contact No No 1st postmentals in contact No Yes Number of nasals 3 3 Rows of dorsal tubercles 10–12 10–12 Number of ventrals in eye diameter 7 7 Tubercles on poster-dorsal side of thighs Toe lamellae No No st 4–5 4 nd 6 6 rd 3 7 7 4th 8–9 7 5th 8 7–8 1 2 Resurrection of Hemidactylus fragilis Calabresi, 1915 We examined all specimens previously identified as H. fragilis, which are: (1) the holotype (MZUF 707; adult male; Somalia, Bur Meldac [=Meel Daaq]); (2) MSNG 53212 (adult female; Ethiopia, Dolo); (3) MSNG 53213 (adult female; Somalia, Rahanuin). Their distribution is summarized in Fig. 1B. Hemidactylus fragilis has the following combination of morphological characters: (1) small size with maximum recorded SVL 31.5 mm for the male and 37.2 mm for females (Fig. 5B,C,D); (2) dorsum with subimbricate scales (Fig. 5H) with a few enlarged, smooth, flat tubercles on flanks of posterior half of body, usually in one or two rows on each side; (3) no enlarged tubercles on postero-dorsal side of thighs; (4) subcaudals slightly enlarged only on first three whorls; (5) six enlarged tubercles on each whorl of the tail (Fig. 5E); (6) tail conical without basal constriction (Fig. 5D); (7) 26 preano-femoral pores in the male; (8) three lamellae under the 1st, five under the 3rd, seven under the 4th toe; (9) 7–8 infralabials and 8–9 supralabials; (10) snout forming a convex line between eye and nostril in lateral view; (11) 536 · Zootaxa 4117 (4) © 2016 Magnolia Press MAZUCH ET AL. anterior postmentals in narrow median contact (Fig. 5G); (12) anterior postmentals only in contact with first infralabials (Fig. 5G); (13) length to width ratio of anterior postmentals 1.35–1.66; (14) 8–9 gulars in the first row behind postmentals (Fig. 5G); (15) four dark transverse bands on dorsum between axilla and groin, one on neck and one on occiput; distinct dark line from nostrils through eyes and ear openings to shoulder region and continuing diffusely to inguinal region; three dark bands on dorsum of anterior half of the tail (Fig. 5B,C,D,E). FIGURE 5. (A) A topotype adult male of Hemidactylus laticaudatus from Harar, Ethiopia; (B) The holotype of Hemidactylus fragilis (MZUF 707) from Bur Meldac, Somalia; (C) adult female of H. fragilis (MSNG 53212) from Dolo Odo, Ethiopia, with regenerated tail; (D) female of H. fragilis (MSNG 53213) from Rahanuin, Somalia, with partially regenerated tail; (E) detail of the tail of the previous specimen, note the enlarged tubercles on the original part of tail; (F) detail of mental region of H. frenatus (MZUF 27607) from Mogadishu, Somalia; (G) detail of mental region of H. fragilis (MSNG 53213) from Rahanuin, Somalia; (H) detail of dorsolateral sub-imbricate scales of H. fragilis (MSNG 53213) from Rahanuin region, Somalia. Hemidactylus fragilis can be distinguished from H. frenatus, with which it has been synonymized, by the following characters (condition in H. frenatus in parentheses): smaller size with max. SVL 37.2 mm (versus 50–60 mm); subimbricate dorsal scales (versus juxtaposed); few scattered enlarged tubercles on caudal part of flanks in REDISCOVERY OF HEMIDACTYLUS LAEVIS Zootaxa 4117 (4) © 2016 Magnolia Press · 537 one or two rows on each side (Fig. 5C) (versus enlarged dorsal tubercles in irregular rows, usually well developed posteriorly); larger dorsal scales, 11–12 in eye diameter (versus smaller, 15–17 in eye diameter); less elongated anterior postmentals, length to width ratio 1.35–1.66 (versus 1.62–2.6; Fig. 5F,G); 8–9 gular scales in the first row behind postmentals (versus 12–15; Fig. 5F,G); color pattern composed of dark transverse bands on dorsum (versus pattern lacking transverse bands, usually longitudinally striped or patternless; Fig. 5B,C,D,E). Morphological differences between the two species are summarized in Table 2. Based on this evidence we resurrect H. fragilis from the synonymy of H. frenatus. TABLE 2. Comparison of diagnostic characters of H. laevis, H. laticaudatus, H. fragilis and H. frenatus. Most distinctive characters between H. fragilis and H. frenatus are in bold. (* counted longitudinally; ** measured in the middle of each scale). H. laevis (n=2) H. laticaudatus (n=22) H. fragilis (n=3) H. frenatus (n=12) Maximum SVL (mm) 39 62.0 37.2 60.0 Enlarged tubercles on dorsum Developed, but smaller and in fewer regular rows Well developed, higher number of regular rows Few scattered on flanks in second half of body, in one or two rows on each side Rather irregular rows, usually well developed in second half body Dorsal lepidosis Juxtaposed Juxtaposed Sub-imbricate Juxtaposed Number of dorsals in the eye diameter (*) 11–12 9–11 11–12 15–17 Tubercles on postero-dorsal side No of thighs Yes No No Enlarged tubercles on tail No Yes Yes Yes Tail with basal constriction Yes Yes No No Length to width ratio of anterior 1.54–1.77 postmentals (**) 1.35–1.69 1.35–1.66 1.62–2.6 Number of gulars in first row behind postmentals 8–12 8–12 8–9 12–15 Dark transverse bands on dorsum Yes Yes Yes No Hemidactylus fragilis differs as follows from the species that can be found or are expected to occur in the Horn of Africa: Hemidactylus angulatus, H. arnoldi, H. awashensis, H. barbierii, H. barodanus, H. bavazzanoi, H. citernii, H. granchii, H. jubensis, H. laevis, H. laticaudatus, H. mabouia, H. macropholis, H. mercatorius, H. mrimaensis, H. platycephalus, H. puccionii, H. robustus, H. ruspolii, H. sinaitus, H. smithi, H. taylori, and H. yerburii pauciporosus have enlarged dorsal tubercles on the whole dorsum, nape and head. Hemidactylus somalicus does not have depressed body habitus, has very small proximal subdigital lamellae, scarcely larger than the granules on the sole, tail without enlarged tubercles, and first supralabial excluded from nostril. Hemidactylus albopunctatus, H. funaiolii, H. isolepis, H. klauberi, H. megalops, H. modestus, H. ophiolepis, and H. ophiolepoides have homogenous dorsal scales and weakly developed subdigital lamellae, head short and not depressed, and tail without enlarged tubercles. Hemidactylus barbouri, H. squamulatus, and H. tropidolepis have heterogeneous dorsal scales without tubercles, some or all scales are keeled, well developed enlarged tubercles on nape and occiput, and head and body not depressed. Hemidactylus flaviviridis is larger (maximum SVL about 95 mm), has swollen, carrot-shaped tail clearly constricted at its base, and dorsal scales small, granular and juxtaposed. Hemidactylus curlei has swollen, carrot-shaped tail clearly constricted at its base, scales on tail large, sub-imbricate and without enlarged tubercles, no enlarged tubercles in sacral region, and males with 4 precloacal pores. 538 · Zootaxa 4117 (4) © 2016 Magnolia Press MAZUCH ET AL. Discussion Rediscovery of H. laevis. In this study we confirm the presence of H. laevis in Erigavo, Sanaag Region, Somaliland, which represents the second known locality of the species. Both localities are situated at altitudes between 1700–1800 m and are characterized by typically montane habitat. Since similar habitats with comparable environmental conditions are also present in other neighbouring mountain ranges like the Wagar, Surud, Cal Madow, and Miskaat Mts. (Fig. 1), we assume H. laevis to be more widespread in the region. The new record of H. laevis lies in the middle of the town of Erigavo with predominantly stone buildings and depauperate vegetation. The town is surrounded by a flat gypseous plateau covered by mixed grassy and shrubby vegetation with occasional trees, mainly Acacia etbaica (Fig. 2F), which we believe to be its primary habitat in this area. Approximately 14 km north of Erigavo, at Daalloh, the habitat is hilly and wetter and supports a Juniperus type of forest (Fig. 2G). Further research is necessary to resolve the occurrence of H. laevis in this type of habitat, more similar to that on top and northern slopes of Gaan Libah. The months of December and January when fieldwork took place are part of the main dry season (‘Jilal’), which lasts from November to March, and are also the coldest months of the year with mean daily temperatures of 14.3–15.0 °C (Taylor in Parker 1942; Hemming 1966; Muchiri 2007). Based on our own observations of the activity patterns of other reptile species in the same general area in months with higher mean daily temperatures (April to September), we expect H. laevis to be more active at this time of the year. Boulenger (1901) described H. laevis as having upper parts of body covered with small uniform granules. However, our examination of the holotype (BMNH 1946.8.25.42) revealed that it has, like H. laticaudatus, the dorsum covered with heterogeneous scales consisting of both small granules and larger, rounded, flat tubercles (Fig. 4H, 6A,B). Based on their morphological resemblance and montane habitat preferences we regard H. laevis closely related to H. laticaudatus. The new specimen of H. laevis differs from the holotype (BMNH 1946.8.25.42) in the arrangement of anterior postmentals (wide contact versus separated by one scale; Figs. 3E and 4E). However, as indicated by the examination of the arrangement of the postmentals of 22 individuals of H. laticaudatus across its wide range in Ethiopia and Eritrea (Appendix 1), this character is too variable to be used as diagnostic within this group of Hemidactylus. FIGURE 6. Comparison of the holotype (A,B) and the newly collected specimen (C,D) of H. laevis. Enlarged tubercles are highlighted in red, red line marks the vertebral line. REDISCOVERY OF HEMIDACTYLUS LAEVIS Zootaxa 4117 (4) © 2016 Magnolia Press · 539 Hemidactylus laevis is mentioned by some authors also from southern Somalia (Boulenger 1912; Loveridge 1947; Laurent and Gans 1965). Detailed examination of Boulenger’s (1912) specimens led us to confirm that, in fact, they belong to H. fragilis, as redetermined by Parker (1942). Laurent and Gans (1965) listed two specimens (MCZ 70525–70526) of H. laevis from 3 and 5 km north of Mogadishu. They were, however, uncertain about the determination since both specimens differed from true H. laevis in not having the subdigital lamellae abruptly differentiated from the scales of the soles. We have seen images of both specimens and conclude that they represent H. puccionii. Hemidactylus laevis is therefore known with certainty only from two localities—Gaan Libah, Togdheer Region and Erigavo, Sanaag Region, Somaliland (Fig. 1). Resurrection of H. fragilis. By resurrecting H. fragilis from thy synonymy of H. frenatus we restrict the distribution of the latter to coastal areas of East Africa from where it is, nonetheless, very poorly documented. Spawls et al. (2004) listed it from coastal Kenya, southern Somalia, and Djibouti. Museum records verified by us confirm that H. frenatus is relatively widespread in the mesic areas of southern Somalia. The coastal character of the distribution is consistent with the propensity of the species to be easily transported via anthropogenic means, especially to port cities where cargo is unloaded (Kraus 2009). Genetic data suggest that it has been recently introduced to Africa from Southeast Asia (Carranza and Arnold 2006). The distribution of H. fragilis is for the time being confirmed from a relatively small area in southern Somalia and extreme southern Ethiopia. However, it cannot be ruled out that further study of museum specimens or fieldwork will bring new localities of this species from the region. Acknowledgements We wish to thank the following curators for allowing access to comparative material housed at their institutions: P.D. Campbell (NHMUK), G. Doria (MSNG), S. Scali (MSNM), A. Nistri (MZUF), S. Schweiger (NMW). We thank P. Hanáček, F. Kovařík, P. Novák, P. Bruyns, R. Štarha and V. Trailin for their assistance with the fieldwork. We thank S. Baha El Din, B. Branch, B. Drewes and P. Wagner for their helpful comments, P. Bruyns, P. Hanáček, T. McCoy, G. Orlando for their help with identification of plants, A. Nistri, E. Razzetti and R. Sindaco for their help with translation from Italian. The work of JŠ was supported by Ministry of Culture of the Czech Republic (DKRVO 2016/15, National Museum, 00023272). AMB was supported by National Science Foundation EF 1241885 (subaward 13-0632). References Andersson, L.G. (1910) A new species of Hemidactylus from Harrar, Abyssinia. Jahrbüchern des Nassauischen Vereins für Naturkunde in Wiesbaden, 63, 200–205. Baha El Din, S. (2006) A Guide to the Reptiles and Amphibians of Egypt. The American University in Cairo Press, Cairo & New York, 359 pp. Bauer, A. (1994) Gekkonidae (Reptilia, Sauria). Part I. Australia and Oceania. Das Tierreich, Part 109, Walter de Gruyter, Berlin, 306 pp. Boulenger, G.A. (1901) A list of the batrachians and reptiles obtained by Dr. Donaldson Smith in Somaliland in 1899. Proceedings of the Zoological Society of London, 1901, 47–49. Boulenger, G.A. (1912) Missione per la frontiera Italo—Etiopica sotto il comando del Capitano Carlo Citerni. Risultati zoologici. List of the reptiles and batrachians. Annali del Museo Civico di Storia Naturale di Genova, 3, 329–332. Calabresi, E. (1915) Contributo alia conoscenza dei Rettili della Somalia. Monitore Zoologico Italiano, 26, 234–247. Calabresi, E. (1925) Anfibi e rettili raccolti dal Signor Ugo Ignesti nell´Abissinia settentrionale. Atti della Societa Italiana di Scienze Naturali, e del Museo Civico di Storia Naturale, Milano, 64, 100–109. Carranza, S. & Arnold, E.N. (2006) Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 38, 531–545. http://dx.doi.org/10.1016/j.ympev.2005.07.012 De Lisle, H.F., Nazarov, R.A., Raw, L.R.G. & Grathwohl, J. (2013) Gekkota: A catalog of recent species. Privately published, Winnipeg, 387 pp. Hemming, C.F. (1966) The vegetation of the northern region of the Somali Republic. Proceedings of the Linnaean Society of London, 177, 173–250. http://dx.doi.org/10.1111/j.1095-8312.1966.tb00958.x 540 · Zootaxa 4117 (4) © 2016 Magnolia Press MAZUCH ET AL. Kluge, A.G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1–209. Kraus, F. (2009) Alien Reptiles and Amphibians, a Scientific Compendium, and Analysis. Springer, Dordrecht, 563 pp. Lanza, B. (1983) A list of the Somali amphibians and reptiles. Monitore Zoologico Italiano (Nuova Serie) Supplemento, 18, 193–247. Lanza, B. (1990) Amphibians and reptiles of the Somali Democratic Republic: check list and biogeography. Biogeographia, 14, 407–465. Largen, M. & Spawls, S. (2006) Lizards of Ethiopia (Reptilia Sauria): an annotated checklist, bibliography, gazetteer and identification key. Tropical Zoology, 19, 21–109. Largen, M. & Spawls, S. (2010) The Amphibians and Reptiles of Ethiopia and Eritrea. Edition Chimaira, Frankfurt am Main, 687 pp. Laurent, R.F. & Gans, C. (1965) Lizards; In: Gans, C., Laurent, R.F. & Pandit, H. (Eds.), Notes on a herpetological collection from the Somali Republic. Annales du Musée Royal de l´Afrique Centrale (8vo), Sciences Zoologiques, Tervuren, 134, pp. 25–45. Loveridge, A. (1947) Revision of the African lizards of the family Gekkonidae. Bulletin of the Museum of Comparative Zoology, 98, 1–469. Loveridge, A. (1957) Checklist of the reptiles and amphibians of East Africa (Uganda; Kenya; Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, 117, 153–362. Malonza, P.K. & Bauer, A.M. (2014) A new species of arboreal forest-dwelling gecko (Hemidactylus: Squamata: Gekkonidae) from coastal Kenya, East Africa. Zootaxa, 3786 (2), 192–200. http://dx.doi.org/10.11646/zootaxa.3786.2.7 Mazuch, T. (2013) Amphibians and Reptiles of Somaliland and Eastern Ethiopia, based on two field trips in 2010/ 2011. Tomáš Mazuch, Dříteč, 80 pp. Muchiri, P.W. (2007) Climate of Somalia. Technical Report No W-01. FAO-SWALIM, Nairobi, 73 pp. Parker, H.W. (1932) Two collections of amphibians and reptiles from British Somaliland. Proceedings of the Zoological Society of London, 1932, 335–367. Parker, H.W. (1942) The lizards of British Somaliland. Bulletin of the Museum of Comparative Zoology, 91, 1–101. Pichi-Sermolli, R.E.G. (1957) Una carta geobotanica dell´Africa orientale (Eritrea, Etiopia, Somalia). Webbia: Journal of Plant Taxonomy and Geography, 13, 15–132. http://dx.doi.org/10.1080/00837792.1957.10669673 Rösler, H. (2000) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28–153. Scortecci, G. (1928) Una nuova specie di Hemidactylus dell’ Eritrea: Hemidactylus fossatii. Atti della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale, Milano, 67, 33–36. Scortecci, G. (1929) Una nuova specie di Hemidactylus dell’ Eritrea: Hemidactylus zolii. Atti della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale, Milano, 68, 116–120. Sindaco, R., Razzetti, E., Ziliani, U., Wasonga, V., Carugati, C. & Fasola, M. (2007) A new species of Hemidactylus from Lake Turkana, Northern Kenya (Squamata: Gekkonidae). Acta Herpetologica, 2, 37–48. Spawls, S., Howell, K., Drewes, R.C. & Ashe, J. (2002) A field guide to the reptiles of East Africa. Academic Press, London, 543 pp. Šmíd, J., Carranza, S., Kratochvíl, L., Gvoždík, V., Nasher, A.K. & Moravec, J. (2013) Out of Arabia: a complex biogeographic history of multiple vicariance and dispersal events in the gecko genus Hemidactylus (Reptilia: Gekkonidae). PLoS ONE, 8, e64018. http://dx.doi.org/10.1371/journal.pone.0064018 Šmíd, J., Mazuch, T. & Sindaco, R. (2014) An additional record of the little known gecko Hemidactylus granchii Lanza, 1978 (Reptilia: Gekkonidae) from Somalia. In: Capula, M. & Corti, C. (Eds.), Scripta Herpetologica. Studies on Amphibians and Reptiles in honour of Benedetto Lanza. Societas Herpetologica Italica, Latina, pp. 165–169. Šmíd, J., Moravec, J., Kratochvíl, L., Nasher, A.K., Mazuch, T., Gvoždík, V. & Carranza, S. (2015) Multilocus phylogeny and taxonomic revision of the Hemidactylus robustus species group (Reptilia, Gekkonidae) with descriptions of three new species from Yemen and Ethiopia. Systematics and Biodiversity, 13, 346–368. http://dx.doi.org/10.1080/14772000.2014.996264 Uetz, P. & Hošek, J. (2015) The Reptile Database. Available from: http://www.reptile-database.org (accessed 7 March 2015) Wermuth, H. (1965) Liste der rezenten Amphibien und Reptilien. Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, 1–246. REDISCOVERY OF HEMIDACTYLUS LAEVIS Zootaxa 4117 (4) © 2016 Magnolia Press · 541 APPENDIX 1. List of specimens examined for morphological comparison. H. frenatus (12 individuals)—MZUF 27607 (MorphoBank M374789–M374805) (Somalia, Mogadishu); MSNM 549 (MorphoBank M374773–M374788) (Somalia, Gelib), MCZ 169941 (syntype of H. hexaspis) (Madagascar); TMHC 2011.12.031 (Vietnam), TMHC 2013.11.448 (Vietnam), TMHC 2015.03.617–623 (Vietnam). H. fragilis (3 individuals)—MZUF 707 (MorphoBank M374562–M374581) (holotype, Somalia, Bur Meldac); MSNG 53213 (MorphoBank M374666–M374687) (Somalia, Rahanuin region); MSNG 53212 (MorphoBank M374644–M374665) (Ethiopia, Dolo) (two vouchers under the same number; the second is H. cf. isolepis). H. laevis (1 individual)—BMNH 1946.8.25.42 (MorphoBank M374599–374643) (holotype, Somalia, Gaan Libah). H. laticaudatus (22 individuals)—MSNM 958–959 (Ethiopia, Fiume Caha, below Gondar); MSNM 858–859 (Ethiopia, Gondar); MSNM 626 (holotype of H. l. fossatii) (Eritrea, Saganeiti); NMW 36826 (MorphoBank M374756–M374772) (Ethiopia, Harar); NMW 20385:1 (MorphoBank M374744–M374755) (Ethiopia, Harar); NMW 17741 (MorphoBank M374729–M374743) (Ethiopia, Harar); MSNG 31943 (Ethiopia, Harar); TMHC 2015.02.607 (MorphoBank M374688– M374711) (Ethiopia, Harar); TMHC 2015.02.608 (MorphoBank M374712–M374728) (Ethiopia, Harar); TMHC 2015.02.609–613 (Ethiopia, Harar); TMHC 2015.02.127–129 (Ethiopia, Azezo, SW of Gondar); TMHC 2015.02.130–131 (Ethiopia, Adwa), TMHC 2014.10.499 (Eritrea, Asmara). 542 · Zootaxa 4117 (4) © 2016 Magnolia Press View publication stats MAZUCH ET AL.
