Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar’s rainforest Author Vences, Miguel m.vences@tu-braunschweig.de Author Multzsch, Malte m.multzsch@tu-braunschweig.de Author Gippner, Sven Author Miralles, Aurélien Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire Naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, Paris, France Author Crottini, Angelica 0000-0002-8505-3050 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485 - 661 Vairão, Portugal & Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, 4099 - 002 Porto, Portugal & BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485 - 661 Vairão, Portugal acrottini @ cibio. up. pt; https: // orcid. org / 0000 - 0002 - 8505 - 3050 acrottini@cibio.up.pt Author Gehring, Philip-Sebastian 0000-0002-8266-2184 Faculty of Biology / Biologiedidaktik, University Bielefeld, Universitätsstrasse 25, 33615 Bielefeld, Germany sebastiangehring @ web. de; https: // orcid. org / 0000 - 0002 - 8266 - 2184 sebastiangehring@web.de Author Rakotoarison, Andolalao 0000-0003-2620-440X Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar andomailaka@gmail.com Author Ratsoavina, Fanomezana M. Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar Author Glaw, Frank glaw@snsb.de Author Scherz, Mark D. mark.scherz@gmail.com text Zootaxa 2022 2022-09-01 5179 1 1 61 http://dx.doi.org/10.11646/zootaxa.5179.1.1 journal article 135489 10.11646/zootaxa.5179.1.1 fab2423d-f82d-4543-8720-86cd568c18b6 1175-5326 7040745 70366A84-EBDE-427D-B525-09E5A2D81EB5 Lygodactylus hapei sp. nov. Lygodactylus sp. 26: Gippner et al. (2021) . Holotype . ZSM 298 /2018, female, collected at Djangoa (Djohahely) in the Sambirano Region in north-western Madagascar , approximately at geographical coordinates S13.7993 , E48.3361 , 20 m a.s.l. ( Fig. 15 ), by unspecified local collectors. Diagnosis. Lygodactylus hapei sp. nov. corresponds to a genetically highly distinct lineage from a poorly known site in north-western Madagascar , and forms a clade with L. tantsaha (described above) and L. sp. 24, both from Montagne d’Ambre in the North. Considering this lineage as a new species is justified by its very deep genetic divergence of over 14% to all other Domerguella (16.3–16.4% to L. tantsaha ), differences in scale counts, and a distinct longitudinally striped pattern on the throat not known from any other Domerguella . The new species belongs to subclade A2 within Domerguella as defined herein. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, and absence of a claw on the first finger. Within Domerguella , the new species is only known from one locality in the Sambirano region in northern Madagascar , and differs from the other nominal species of Domerguella by the presence of a longitudinally striped pattern on the throat, and additionally from the species occurring in northern Madagascar as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>185 vs. <170), and by a pattern of dorsolateral stripes (vs. unstriped in the only known specimen (vs. scapular semi-ocellus usually present, and striped pattern unknown); from L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape without elongated limbs (relative hindlimb length 0.43 vs.>0.55); from L. madagascariensis , L. petteri , L. salvi , and L. tantsaha by smaller longitudinal dorsal scale count (179 vs.>188) and smaller longitudinal ventral scale count (87 vs.>96). The new species appears to be very similar to L. roellae , a species from subclade A3, in scale counts and color pattern, but it may differ by smaller body size (SVL 26.3 vs. 35.9–36.0). The new species is genetically highly distinct from all species in subclade A5, based on concordant differentiation in mitochondrial genes (with deep divergence in 16S to all other species:>14%) and the unlinked loci CMOS and RAG-1. In addition it appears to differ by the absence of a spine at the tail base, which is weakly recognizable also in the females of all subclade A5 species except L. fritzi . Furthermore, the longitudinal dorsal scale count is smaller than in all known individuals of this subclade. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. We dedicate this species to Hans-Peter “HaPe” Berghof, in recognition of his contributions to the knowledge of Madagascar geckos, especially Phelsuma . The name is a patronym (i.e., a noun in the genitive case). Description of the holotype . Adult female, in good state of preservation, tail regenerated, fourth toe on the left hind limb is removed as source of tissue for molecular analysis ( Fig. 15 ). SVL 26.3 mm , TAL 27.4 mm ; for other measurements see Table 1 . Head slender with long neck, body broader than head. The distance from the tip of the snout to the anterior border of the eye ( 3.5 mm ) is greater than the interorbital distance anteriorly ( 3.2 mm ), and greater than the distance between the eye and ear opening. Snout covered with granular scales equally sized compared to the rest of the dorsum. Nostril surrounded by three scales: rostral, first supralabial, and two supranasal. Mental scale undivided; no contact between posterior projection of mental scale and first infralabial; two symmetrical postmental scales with five postpostmental scales; seven infralabial scales; eight supralabial scales; three internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular, and unkeeled scales of similar size to those on trunk, the scales on limbs are not distinctly larger; 179 dorsal scales longitudinally along the body; 87 ventral scales between mental and cloaca; venter with large homogeneous smooth scales; first finger present but very small, not bearing a claw; three pairs of subdigital lamellae on the fourth toe; no dorsolateral tubercle; tail without whorls; no obvious lateral spines at the base of the tail. TABLE 2. List of localities of Lygodactylus samples included in this study, with geographical coordinates and elevation. Numbers of decimals in coordinates reflect precision of collecting site information; values with 4–5 digits refer to coordinates taken with GPS devices, other coordinates were usually estimated or inferred from maps. Note that GPS coordinates often refer to campsites and specimens were collected at walking distance from these. * Exact location of Senbendrana unknown but close to Toamasina (type locality of L. miops ); ** record of L. miops from Moramanga relies on types of two junior synonyms ( septemtuberculatus , spinulifer ).

Locality	Species	Latitude	Longitude	Elevation (m)
Ankarana	L. rarus , L. expectatus	S12.95	E49.12	120
Nosy Be	L. madagascariensis	S13.41	E48.33	100
Tsaratanana (Manarikoba: Antsahamanara)	L. salvi	S14.0450	E48.7853	1000
Tsaratanana (Manarikoba: Andampy)	L. madagascariensis	S14.0422	E48.7617	730
Manongarivo	L. madagascariensis	S13.9755	E48.4266	690
Maromiandra	L. madagascariensis	S13.9965	E48.2177	283
Andrafainkona	L. madagascariensis	S13.7133	E49.4911	780
Montagne d’Ambre	L. madagascariensis , L. petteri , L. sp. 24	S12.52	E49.18	1000–1100
Montagne d’Ambre (low elevation)	L. madagascariensis	S12.4903	E49.1716	687
Montagne d’Ambre (west slope)	L. tantsaha	S12.585	E49.116	815–820
Senbendrana (=Sahembendrana) *	L. miops	NA	NA	NA
Moramanga **	L. miops , L. guibei	S18.92	E48.22	900
Manantantely	L. miops	S24.983	E46.917	ca. 20
Andohahela	L. miops	S24.76	46.85	ca. 250
N Andohahela	L. miops	NA	NA	NA
Sainte Luce	L. miops	S24.7667	E47.1833	30
Sainte Luce	L. miops	S24.7701	E47.1707	14
Sampanandrano	L. miops	S24.1399	E47.0742	530
Tsitongambarika:	L. miops	S24.6950	E46.9776	ca. 720
Tsitongambarika: Andranomaizina	L. miops	S24.5838	E47.1474	42
Tsitongambarika: Ivohibe	L. miops	S24.5612	E47.1924	424
Ranomafana	L. miops	S21.26	E47.46	ca. 900
Ambohitsara	L. miops	S21.3571	E47.8153	850
Mahakajy	L. miops	S21.2792	E47.5304	580
Anosibe Anala	L. miops	S19.43	E48.22	ca. 800
Vohimana	L. miops	S18.9208	E48.5158	770
Sahafina	L. miops	S18.8106	E48.9803	60
Betampona: Betakonana	L. miops	S17.9141	E49.2167	356

......continued on the next page TABLE 2. (Continued)

Locality	Species	Latitude	Longitude	Elevation (m)
Betampona: Maintimbato	L. miops	S17.8938	E49.2251	274
Betampona: Piste Fotsimavo	L. miops	S17.9231	E49.2087	205
Betampona: Sahabefoza	L. miops	S17.9127	E49.2107	481
Betampona: Sahaindrana	L. miops	S17.8968	E49.1996	344
Betampona: Sahambendrana	L. miops	S17.9014	E49.2110	458‒558
Betampona: Vohitsivalana	L. miops	S17.8850	E49.2034	481
Ambodivoahangy (Makira)	L. miops	S15.2899	E49.6203	100–300
Andasibe	L. guibei , L. sp. 19	S18.9333	E48.4166	900
Vohidrazana	L. guibei	S18.95	E48.50	700–800
Anjozorobe	L. guibei	S18.40	E47.87	1250
Mahasoa	L. guibei	S17.2977	E48.7020	1030
Angozongahy (Makira west slope)	L. guibei	S15.4370	E49.1167	1010
Ambodikakazo	L. salvi	S14.2131	E48.9052	1310
Ampotsidy	L. roellae , L. winki	S14.42–14.43	E48.71–48.72	1315–1405
Andrevorevo	L. roellae , L. winki	S14.3464	E49.1028	1720
Djangoa	L. hapei	S13.7993	E48.3361	20
Bemanevika	L. winki	S14.4306	E48.6018	1470
Marojejy	L. ulli	S14.4376	E49.7755	480
Ankanin’ny Nofy	L. fritzi	S18.6058	E49.2138	0
Vohibola	L. fritzi	S18.5897	E49.2307	10
Tsingy de Bemaraha	L. hodikazo	S18.7844	E44.8603	430
Antanambe	L. sp. 20	S16.4299	E49.7846	ca. 320

FIGURE 19. Photo of Lygodactylus hapei sp. nov. , holotype ZSM 298/2018 from Djangoa (Djohahely), in life. Photo by H.-P. Berghof. Based on available photograph ( Fig. 19 ), the holotype in life displayed a broad brown stripe on the back with a brighter center running along the spine reaching from the snout to the base of the tail. Along the brighter center, irregularly scattered black spots are present. Flanks are yellowish brown with irregular small dark spots. A distinct black stripe is running from the snout through the eye to the shoulder ending in a black marking somewhat reminiscent of a scapular semi-ocellus, but positioned more posteriorly. Above this, a second whitish and broader stripe is present, reaching from the eye to the shoulder. Dorsally brown with a diffuse pattern consisting of dark and light spots, venter whitish. Flanks brighten towards venter with a diffuse ocelli-like pattern. Brown color on head with distinct border on supralabials to whitish venter. Six black stripes radially arranged around the eye. Tail slightly brighter than dorsum with pairs of black and white spots running posteriorly along the caudal spine ( Fig. 19 ). During preservation in ethanol, the specimen darkened and patterns faded. Preserved specimen displays dark irregular spots on whitish gular region expanding to the anterior ventral torso. Variation. Only a single individual of this species (the holotype ) is known. Natural history. The only known specimen was photographed millimeters from the posterior end of a planthopper larva ( Fig. 19 ), and was presumably consuming honeydew excreted by the insect, as is known from other gecko species ( Fölling et al. 2001 ). Distribution. L. hapei is only known from its type locality, Djohahely.