Zoosyst. Evol. 97 (1) 2021, 55-82 | DOI 10.3897/zse.97.57202 zis eae Ta eee _______ap__. D> PENSOFT. Nites A NATURKUNDE BERLIN Rock island melody: A revision of the Afroedura bogerti Loveridge, 1944 group, with descriptions of four new endemic species from Angola William R. Branch?*7, Andreas Schmitz?, Javier Lobon-Rovira*, Ninda L. Baptista*>®, Telmo Antoénio®:’, Werner Conradie?® Port Elizabeth Museum, P.O. Box 13147, Humewood, 6013, Port Elizabeth, South Africa Department of Zoology, P.O. Box 77000, Nelson Mandela University, Port Elizabeth 6031, South Africa Natural History Museum of Geneva, UREC - Herpetology and Ichthyology, C.P. 6434, 1211 Geneva 6, Switzerland CIBIO Centro de Investigacdo em Biodiversidade e Recursos Genéticos, Universidade do Porto, Rua Padre Armando Quintas, Campus de Vairdo, 4485-661 Vairdo, Portugal Faculdade de Ciéncias, Universidade do Porto, 4169-007 Porto, Portugal Instituto Superior de Ciéncias da Educagdo da Huila (ISCED), Rua Sarmento Rodrigues s/n, Lubango, Angola BR WN Faculty of Natural Resources and Spatial Sciences, Namibia University of Science and Technology, Private Bag 13388, Windhoek, Namibia CON DD oO School of Natural Resource Management, George Campus, Nelson Mandela University, George 6530, South Africa http://zoobank. org/A125EC8 1-050E-449A-8904-27674F 5265 EF Corresponding author: Werner Conradie (werner@bayworld.co.za) Academic editor: Johannes Penner # Received 3 August 2020 Accepted 7 December 2020 @ Published 12 January 2021 Abstract Four new species of flat geckos in the Afroedura bogerti Loveridge, 1944 group are described from south-western and west-central Angola. The description of these new species significantly restricts the distribution range of typical A. bogerti, a morphologically very similar species, from which they differ genetically by 5.9-12% divergence for the mitochondrial /6S ribosomal RNA gene. Morpho- logically and genetically, Angolan Afroedura are divided into two main groups: a mostly south-western coastal group and a west-cen- tral inland/highland group. These two groups are further divisible into three and two subgroups respectively, all geographically isolat- ed, differing by a combination of the following features: colouration, average adult size, number of mid-body scale rows, number of scale rows on dorsal and ventral surface of each tail verticil and if nostril scales are in contact or not. All five Angolan species are mor- phologically distinguishable and in agreement with the molecular results. An updated dichotomous key to the Afroedura transvaalica group is provided. The new discovery adds to a growing number of endemic Pro-Namib reptiles described from Angola in recent years. Resumo Aqui sao descritas quatro novas espécies de osga-achatada, do grupo Afroedura Loveridge, 1944, do sudoeste e centro-oeste de Angola. A descricéo destas novas espécies reduziu significativamente a area de distribui¢do da A. bogerti tipica, uma espécie uma espécie morfologicamente muito semelhante as primeiras, da qual estas tém entre 5.9 e 12% de diferenga genética para o gene mito- condrial /6S rRNA. Do ponto de vista morfoldgico e genético, as Afroedura de Angola dividem-se em dois grupos principais: 0 do sudoeste, maioritariamente das planicies costeiras e do norte, e o do planalto interior. Estes grupos podem ainda ser divididos em dois e trés subgrupos, respectivamente, todos isolados geograficamente, e estes diferem entre si na combinacao das seguintes caracteristi- cas: coloracao, tamanho médio dos adultos, numero de fileiras de escamas na secc¢ao mediana do corpo, numero de escamas dorsais e ventrais por cada anel caudal de escamas, escamas nasais em contacto ou nao. E possivel distinguir morfologicamente as cinco espécies de Angola, de forma concordante com os resultados da genética. Aqui apresentamos uma chave dicotomica actualizada para 0 grupo Afroedura transvaalica. Esta nova descoberta junta-se ao crescente numero de répteis endémicos do Pro-Namibe descritos em Angola nos ultimos anos. t+ Deceased Copyright William R. Branch etal. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 56 Key Words William R. Branch et al.: New species of Afroedura for Angola Biodiversity hotspot, cryptic species, endemism, Gekkonidae, Reptilia Palavras-chave Endemismo, espécies cripticas, Gekkonidae, hotspot de biodiversidade, Reptilia Introduction The current reptile diversity in Angola comprises about 300 species (Marques et al. 2018; Branch et al. 2019a; and subsequent descriptions listed below). New species are constantly being added (Conradie et al. 2012, 2020; Stan- ley et al. 2016; Marques et al. 2019a, b, 2020; Branch et al. 2019b; Ceriaco et al. 2020a, b, c; Hallerman et al. 2020) and more new species are expected in the country (Branch 2016; Branch and Tolley 2017; Branch et al. 2017a, b; But- ler et al. 2019). As Angola is rebuilding after decades of civil war, more remote regions are becoming accessible to researchers, with the promise of additional cryptic species. The Gekkonidae in Angola comprises eight gen- era, Afroedura (one species), Afrogecko (one species), Chondrodactylus (three species), Hemidactylus (eight species), Kolekanos (one species), Lygodactylus (five species), Pachydactylus (11 species) and Rhoptropus (seven species) (see Marques et al. 2018, 2020; Branch et al. 2019a; Ceriaco et al. 2020a, c). A total of 15 species of geckos are considered to be endemic to the country. This includes the monotypic Kolekanos plumicaudus (Haacke, 2008), the recently re-discovered Afrogecko an- sorgii (Vaz Pinto et al. 2019), six species of Hemidactylus (Ceriaco et al. 2020a, c), three species of Lygodactylus (Marques et al. 2020), three species of Rhoptropus and Pachydactylus angolensis (Marques et al. 2018; Branch et al. 2019a). Afroedura bogerti is currently regarded as a near-endemic species as a specimen was also found in northern Namibia (Branch 1998; Griffin 2003). Loveridge (1944) transferred African Oedura to the new genus Afroedura and described A. karroica bogerti from Angola after examining specimens collected during the 1925 Vernay Angolan Expedition. He designated this material as the type for the new genus and regarded it as part of the ‘halli/karroica’ group. Later, Onderstall (1984) investigated relationships between Afroedura species after examining material from south-western Angola collect- ed by Wulf Haacke in the 1970s. He placed A. k. boger- ti in the ‘transvaalica’ group, alongside A. transvaalica (Hewitt, 1925) and A. loveridgei (Broadley, 1963) and ele- vated A. bogerti to full species status, based on differences in numbers of paired scansors per digit and possession of a verticillate tail. Such findings were supported by recent phylogenetic studies, confirming that A. bogerti is part of the ‘transvaalica’ group (Jacobsen et al. 2014; Makhubo et zse.pensoft.net al. 2015; Branch et al. 2017a), with other species occurring to the east in Mozambique, Zimbabwe and South Africa. This relationship of the A. bogerti group to the “transvaal- ica group is of zoogeographic interest as it 1s separated from other members of the group by nearly 2000 km and occupies more arid and/or mesic environments. Up to now, Afroedura bogerti was the only species of flat gecko known from Angola (Marques et al. 2018; Branch et al. 2017b). In a recent phylogenetic study, in which a new species from the ‘transvaalica’ group was described from central Mozambique (Afroedura goron- gosa), cryptic diversity within Angolan A. bogerti pop- ulations was documented (Branch et al. 2017a) and this was elaborated on in more detail in a subsequent study (Branch et al. 2017b). Here we take the opportunity to investigate the Angolan A. bogerti group in detail and de- scribe four new endemic species. Materials and methods Sampling For the last decade (2009-2019), material of Afroedura bogerti was collected from south-western and west-central Angola (Fig. 1; see Branch et al. 2017a). Specimens were euthanised by oral application of benzocaine gel. After euthanasia, some geckos were fixed in 10% formalin and others in 95% ethanol, after which they were transferred to 70% ethanol for long-term storage at the Port Elizabeth Museum (PEM), South Africa, Museum ftir Naturkunde, Berlin, Germany (ZMB), Instituto Superior de Ciéncias da Educa¢éo da Huila (ISCED, material listed here as NB), Huila, Angola and Fundacaéo Kissama (FKH), Luanda, Angola. Liver tissue or tail tips of newly-collected mate- rial was sampled and stored in 95-99% ethanol prior to fixation for further genetic analysis. Molecular analyses We used molecular methods to support the identified morphological groupings. Recent studies on the genus Afroedura (e.g. Branch et al. 2017a) have shown that the fast evolving /6S rRNA gene, with its hypervariable regions, perfectly resolves the different species within Zoosyst. Evol. 97 (1) 2021, 55-82 2620 mas.l @ Afroedura donveae sp. nov. @ Afroedura vazpintorum sp. nov. © Afroedura praedicta sp. nov. @ Afroedura wulfhaackei sp. nov. @ Afroedura bogerti 16°S 57 ' Cuanza Sul Cunene 225 300km Figure 1. Digital elevation map showing the predicted distribution of the five newly-recognised species of Afroedura occurring in Angola (data for A. wulfhaackei sp. nov. also contain the morphologically-indistinguishable Afroedura sp. 5—7 clades). Confirmed locality records are indicated by solid circles, with type localities indicated by stars. Coloured polygons indicate potential distribu- tion areas with more than 90% of climatic habitat suitability. this genus and even provides insights into the intraspe- cific structure of the different monophyletic lineages. Therefore, for our molecular comparisons, we combined newly-sequenced mitochondrial /6S data for specimens from different Afroedura populations in Angola and com- plemented these with previously published (Branch et al. 2017a) sequences from GenBank. Total genomic DNA was extracted from tissue samples using the E.Z.N.A. Tissue DNA Kit (VWR/Omega bio-tek) and the Qiagen DNeasy Tissue Kit, following the manufacturer’s proto- cols. A portion of the mitochondrial genome (/6S rRNA gene) was PCR amplified and sequenced, following the methods described in Schmitz et al. (2005). All sequences have been deposited in GenBank (Table 1). DNA sequences were aligned using the original chro- matograph data in the programme BioEdit (Hall 1999) using ClustalX (Thompson et al. 1997) and the resulting alignment corrected manually. We sequenced 589 bp of the /6S rRNA gene, including some fast-evolving, indel-rich loop regions. We deleted 6 bp that could not be unambiguously aligned, resulting in a total sequence length of 583 bp for each voucher. We used Bayesian Inference (MrBayes v.3.26; Ronquist et al. 2012) and Maximum Likelihood (RAXML v.7.0.4; Stamatakis et al. 2006), using the rapid hill climbing algorithm and the GTR+G model of nucleotide substitution, following Stamatakis et al. (2006), to assess phylogenetic relation- ships. The best-fit model of sequence evolution for the Bayesian analysis was the TIM2+I+G model with values of I = 0.5140 and G = 0.7180, selected using j;Modeltest v.2.1.7 (Darriba et al. 2012) using the Bayesian Informa- tion Criterion (BIC). Bootstrap analyses (BS) with 1000 pseudoreplicates were used to evaluate relative branch support in the ML analysis. We regarded tree topologies with bootstrap val- ues of 70% or greater as supported (Huelsenbeck and Hillis 1993). Bayesian analyses were run for 10 million generations using four chains, sampling every 1000 gener- ations, with the first 25% of trees discarded as burn-in. Sta- tionarity, convergence and mixing of the parameters (ESS values) for the Bayes runs were checked in Tracer v.1.7.1 (Rambaut et al. 2018). Clades with posterior probabilities (PP) = 0.95 were considered strongly supported. In total, sequences from 47 Afroedura specimens from Angola zse.pensoft.net 58 William R. Branch et al.: New species of Afroedura for Angola Table 1. Afroedura specimens (field and museum numbers), generalised localities (see systematic accounts for full details) and GenBank accession numbers of vouchers used in this study. Species Locality Sample Number Museum Number GenBank Number A. donveae sp. nov. Omauha Lodge KTHO9-196 PEM R17936 LM993776 KTHO9-197 PEM R17937 LM993777 P9-284 NA MW354008 P9-285 NA MW354009 A. praedicta sp. nov. Serra da Neve NB 853 ZMB 91607 MW354010 NB 854 ISCED-NB 854 MW354011 NB 855 ISCED-NB 855 MW354012 A. vazpintorum sp. nov. 50 km east Namibe on main tar road to Leba ANG 289 PEM R21595 MF565454 Meva Beach “30” PEM R22488 MF565455 10.4 km south of Rio Mucungo on tar road to Bentiaba samp 57/E260.13 PEM R24203 MF565458 samp 58/E260.14 PEM R24204 MF565457 samp 39/E260.12 NA MF565459 1 km east of Farm Mucungo AG 137 PEM R24114 MF565460 AG 138 PEM R24115 MF565463 AG 141 PEM R24118 MF565462 20 km south Bentiaba samp 62/E260.15 PEM R24219 MF565456 52 km north on tar road on road to Lucira ANG 311 PEM R21596 MF565461 Mariquita P9-154 NA MW354018 approx. 18 km E Lucira NB 834 ISCED-NB 834 MW354019 NB 835 ISCED-NB 835 MW354020 Carivo P8-20 NA MW354016 P8-19 NA MW354015 Bimbe, Esta¢do Zootecnica NB 743 ISCED-NB 743 MW354017 NB 745 ISCED-NB 745 MW354013 NB 746 ISCED-NB 746 MW354014 A. bogerti Farm Namba samp 23/E260.1 PEM R24184 MF565467 samp 24/E260.2 PEM R24185 MF565468 samp 25/E260.3 PEM R24186 MF565466 400 m north of Mission de Namba grounds samp 27/E260.4 PEM R24187 MF565465 samp 28/E260.5 NA MF565464 Namba JLRZCOO15 NA MW354021 JLRZCOO016 NA MW354022 A. wulfhaackei sp. nov. 5 km west of Maka-Mombolo samp 71/E260.17 PEM R24232 MF565477 samp 72/E260.18 PEM R24233 MF565478 samp 73/E260.19 PEM R24234 MF565479 Maka-Mombolo, north-east of Balombo samp 70/E260.16 PEM R24236 MF565476 Morro do Moco, camp near Canjonde NB 817 ISCED-NB 817 MW354024 NB 818 ISCED-NB 818 MW354025 NB 819 ISCED-NB 819 MW354026 A. sp. 5 Farm Victoria-Verdun, 2 km S of Mt. Sandula samp 33/E260.8 PEM R24191 MF565471 samp 34/E260.9 PEM R24192 MF565469 samp 31/E260.6 NA MF565470 Sandula P9-141 NA MW354023 A. sp. 6 5 km southwest of Lepi samp 37/E260.11 PEM R24201 MF565472 A. sp. 7 Candumbo Rocks Memorial samp 35/E260.10 PEM R24200 MF565473 WC-4037 PEM R22490 MF565474 WC-4038 PEM R22491 MF565475 were included in the phylogenetic analysis (Table 1) to corroborate our morphological analyses. Afroedura love- ridgei (MF565446), shown by Branch et al. (2017a) to be a sister taxon, but still well outside the Angolan clade, was chosen as outgroup. Pairwise comparisons of uncorrect- ed sequence divergences (p-distance) were computed in MEGA X (Kumar et al. 2018; Table 2). Morphology We examined material in the collections of the Port Eliza- beth Museum (PEM), Ditsong National Museum of Natu- ral History (formerly Transvaal Museum ~ TM), Museum fir Naturkunde, Berlin, Germany (ZMB) and Instituto zse.pensoft.net Superior de Ciéncias da Educagao da Huila (ISCED), Huila, Angola (see Material examined under systematic accounts). High quality photographs of the type specimen of Afroedura bogerti were also examined. As with many other gecko genera, Afroedura is mor- phologically conservative, with few discrete differences. The following characters (detailed in Jacobsen 1992) were assessed: 1) presence or absence of internasal gran- ules between the nasal scales; 2) number of postmental scales; 3) number of scales in contact in a straight line be- tween the anterior corners of eyes across the crown of the head; 4) number of scales between upper edge of earhole and rear margin of eye, counted along the shortest dis- tance between them; 5) number of scales between nostril and front edge of orbit, excluding postnasal; 6) number Zoosyst. Evol. 97 (1) 2021, 55-82 of enlarged supralabials to the angle of the jaw at mid-or- bital position; 7) number of enlarged infralabials to the angle of the jaw at mid-orbital position; 8) number of mid-body scale rows (MSR), counted at the widest part of the trunk; 9) number of scale rows on dorsal surface per tail whorl (counted 3-6 verticils, posterior to the clo- aca); 10) number of scales rows on ventral surface per tail whorl (counted 3-6 verticils, posterior to the cloaca); and 11) number of precloacal pores in males. The following measurements were taken in millime- tres (mm) using a digital calliper (accuracy of 0.1 mm) with the aid of a Nikon SMZ1270 dissecting microscope: 1) snout-vent length (SVL — from the tip of the snout to the cloaca with the gecko flattened on its back), 2) tail length (TL, only original tails were measured); 3) head length (HL — tip of snout to retroarticular process of jaw); 4) head width (HW — widest point of head, approximately at the level of eyes); 5) snout length (SL — tip of snout to front of orbit); 6) eye diameter (ED —measured in hori- zontal orientation); 7) ear to eye length (EE — top edge of earhole to back of eye); 8) ear opening (EO — great- est length); and 9) internostril distance (IN — shortest distance between nostrils). All head measurements were taken on the right side of the head. Measurements and scale counts were carried out most- ly by WRB and WC incorporated newly-collected mate- rial to supplement the final dataset. Only adult material was used for additional comparative morphological anal- ysis. Specimens were considered adult (sexually mature) at SVL > 38 mm (following Makhubo et al. 2015). Box- plots were created for all measurements and scale counts. Only the following measurements SVL, HL, HW, ED, EE and IN, number of scales rows on the dorsal surface per tail whorl, precloacal pores and MSR were included in morphological comparisons. The other measurements and scale counts were found not to be informative for further comparative analysis. These morphological data were test- ed for significant differences amongst species pairs using a Tukey multiple comparison of means test. Differences were considered significant at P < 0.05. All above-men- tioned quantitative morphological comparisons were con- ducted using R v.3.4.1 (R Development Core Team 2014). Potential distribution model In order to identify potential distribution areas for Ango- lan Afroedura, an bioclimatic niche model was conducted using Maxent (Yang et al. 2013). The sampling area in- cluded a buffer of two degrees (~250 km) from the most peripheral observations of Afroedura species. Nineteen bioclimatic variables were obtained in WorldClim dataset (Fick and Hijmans 2017; http://www.worldclim.org/) at a spatial resolution of 30 arc-second (~ | km/7). For those variables, we ran a correlation model to eliminate collin- earity between variables in the sampled area and within sample points (Candau and Flemin 2005) and variables, with correlation coefficient > 0.7, were selected in order 59 to capture the bioclimatic range over the distribution of the species (Enriquez-Urzelai et al. 2019). Therefore, the variables included for the analysis were: mean di- urnal temperature range (BIO1); maximum temperature of the warmest month (BIOS), minimum temperature of the coldest month (BIO6); annual precipitation (BIO12); precipitation seasonality (BIO15), precipitation of wet- test quarter (BIO16). Given the small sample size for some species, we ran a cross-validation model using all the samples but leaving one out in each run, and hinge features with the regularisation parameters set to 2.5, to produce smoother response curves and reduce overfitting (Bittencourt-Silva et al. 2016; Briscoe et al. 2016). The final maps were generated selecting areas with more than 90% of habitat suitability for each species. Results Molecular analyses Molecular analyses confirm that all Angolan Afroedura form a monophyletic clade, distinct from other members of the ‘transvaalica’ group (Branch et al. 2017a). The An- golan clade is resolved into eight genetically well-support- ed clades (Fig. 2), most of which show consistently large genetic differences between them (5.9—-12%, Table 2) to a level that can be considered representative of species-level divergence (Table 2). While these values are lower than observed between other species of the ‘transvaalica’ group (Branch et al. 2017a), they are still surprisingly high for a local monophyletic radiation and indicative of recent and ongoing speciation in the genus Afroedura in Angola. Two sister clades occur exclusively in the coastal re- gion (except for the first subclade, see below) with a third related clade (Afroedura sp. 2) occurring a little further inland in the Serra da Neve mountain range (Fig. 1). The southernmost taxon within the identified coastal region (Afroedura sp. 1) is currently restricted to the arid Namib Desert region of south-western Angola and clusters with the species from Serra da Neve. Both clades are genetical- ly well differentiated and show minimal internal genetic variation (> 0.8%, Table 2). The sister taxon to those two taxa (Afroedura sp. 3) is much more widely distributed across the semi-arid ‘Pro-Namib’ habitats of the coastal littoral north of Mocamedes (= Namibe) (Fig. 1). Congru- ent with its much larger distribution, it also contains some clear, but consistent, genetic substructure (Fig. 2, Table 2). There are two well-supported major subclades within this taxon and the second, larger subclade shows some additional substructure. The first subclade contains two specimens from Bimbe (NB 745-6) on the Humpata pla- teau which are genetically quite distant from the rest of the coastal clade (~ 4.1%, Table 2). However, a third sam- ple from the same locality (NB 743) groups well within the coastal clade. Follow-up analyses will be needed to resolve the status of this population which may prove to be not conspecific with Afroedura sp. 3. zse.pensoft.net 60 William R. Branch et al.: New species of Afroedura for Angola Afroedura donveae sp. nov. P9-284 Afroedura donveae sp.nov. P9-285 Sp.1 Afroedura donveae sp. nov. LM993776 Afroedura donveae sp. nov, LM993777 0.97 Afroedura praedicta sp, nov, NBO853 1 400 Afroedura praedicta sp. nov.Ng0855 Sp. 2 100 Afroedura praedicta sp. nov. NB0854 1 Afroedura vazpintorum sp. nov, NBO745 100 Afroedura vazpintorum sp. nov. NBO746 41 Afroedura vazpiniorum sp. nov. MF 565455 96 1 Afroedura vazpintorum sp. nov. P8-19 100 Afroedura vazpintorum sp. nov. P8-20 Afroedura vazpintorum sp. nov, MF565454 65 Afroedura vazpintorum sp. nov. NBO743 Afroedura vazpiniorum sp. nov. P9-154 1 Afroedura vazpintorum sp. nov. NBOS24 100 Afroedura vazpintorum sp. nov. NBO835 Afroedura vazpintorum sp. nov. MF565456 0.98 84 Afroedura bogerti MF 565464 Afroedura bogerti MF 565465 Afroedura bogerti MF565466 4 Afroedura bogerti MF565468 100 Afroedura bogerti MF565467 ogil Afroedura bogeni JURZC0015 a Afroedura bogerti JURZC0016 Afroedura sp. 5 MF565469 1 Afroedura sp. 5 MFS65470 99 Afroedura sp. 5 MF565471 0.84 Afroedura sp. 5 P$-141 70 Afroedura sp, 6 MF565472 Afroedura wulfhaackei sp. nov. MF 565476 Afroedura wulfhaackei sp. nov. MF565479 Afroedura wulfhaackei sp. nov. MF 565477 0.99 Afroedura wulfhaackei sp. nov. MF 565478 91 Afroedura wulfhaackei sp. nov. NBO817 i Afroedura wulfhaackei sp. nov. NBOB19 100 Afroedura wulfhaackei sp. nov. NBO818 Afroedura sp. 7 MF565474 0.87 Afroedura sp. 7 MF565475 96 Afroedura sp. 7 MF585473 Afroedura loveridgei MF565446 0.03 Sp. 3 Afroedura vazpintorum sp. nov. MF 565457 1 Afroedura vazpintorum sp. nov. MF 565459 78 Afroedura vazpintorum sp. nov. MF565460 Afroedura vazpintorum sp. nov, MF565463. Afroedura vazpintorum sp. nov. MF565461 Afroedura vazpintorum sp. nov. MF565462 Afroedura vazpintorum sp. nov, MF565458 [sp 6] Figure 2. Phylogenetic tree topology for the /6S gene (identical for Bayes and ML) using Afroedura loveridgei as outgroup. Support values for posterior probability and bootstraps are indicated by circles (ML: > 70% / PP: > 0.95). Table 2. Summary of intra- and interclade uncorrected pairwise genetic distances for specimens of Afroedura clades compared to A. loveridgei for 16S rRNA. Intraclade/intraspecific distances are marked in bold. Intraclade Species/Clade distances | A. loveridgei | A. donveae sp. nov. A. praedicta sp. nov. A. loveridgei A. donveae sp. nov. Interclade/interspecific distances A. vazpintorum | A. bogerti | A.sp.5 |A.sp.6| A. sp. 7 | A. wulfhaackei sensu stricto Sp. nov. A. praedicta sp. nov. A. vazpintorum Sp. NOV. A. bogerti sensu 0.1757 0.1083 0.0908 0.1124 stricto A. sp. 5 0.0000 0.1588 0.1021 0.0988 0.1071 0.0634 - A. sp. 6 0.1667 0.1037 0.0976 0.1074 0.0650 0.0353 A. sp. 7 0.0017 0.0422 | 0.0378 - A. wulfhaackei 0.0033 0.0399 | 0.0327 - Sp. Nov. Five further, strongly supported, monophyletic clades occur above the Angolan escarpment and within the re- gion of the putative type locality. The Namba population (here referred to as A. bogerti sensu stricto) is genet- ically well-differentiated from four southern highland clades (Afroedura sp. 4—7) and shows negligible internal genetic variation (> 0.15%, Table 2). The four remaining southern highland clades (Afroedura sp. 4—7), each of which is strongly supported, are genetically closely re- lated (Table 2). This complex is very similar to the mono- phyletic clade Afroedura sp. 3 in having a comparative- ly-wide distribution (Fig. 1) and containing a surprising, but consistent, genetic substructure. The geographic and zse.pensoft.net taxonomic implications of these groupings require fur- ther analysis. The clades of Afroedura sp. 4—7 all show very low internal variation versus higher inter-clade variation. Genetic distances between Afroedura sp. 4—7 are about, on average, half as great as the shortest dis- tances identified between the major clades identified in Angola (3.3—4.2% vs. 5.9-12.0%; Table 2). Morphology Results for the morphological analysis are summarised in Table 3 and Figure 3 and are discussed in more detail in Zoosyst. Evol. 97 (1) 2021, 55-82 61 SVL Head Length Head Width 65 60 55 50 45 40 Eye Diameter MSR Ventral Verticiles 90 85 80 75 70 65 2 2 z z < 2 2 2 2 z 2 é 2 2 z Figure 3. Summary boxplots (top whisker — maximum value; lower whisker — minimum value; dark horizontal line — median; box — 1*and 3" quartile) comparing morphological features amongst the four species of Afroedura separated by sex: Afroedura bogerti (dark green —Ab; n= 9), Afroedura wulfhaackei sp. nov. (including the morphologically-indistinguishable Afroedura sp. 5—7 clades; blue — Aw; n=40), Afroedura praedicta sp. nov. (light green — Ap; n=5), Afroedura donveae sp. nov. (red —Ad; n= 17), Afroedura vazpintorum sp. nov. (orange — Av; n = 48). Table 3. Summary of morphological data for Afroedura bogerti, A. wulfhaackei sp. nov. (including the members of the morpholog- ically-indistinguishable Afroedura sp. 5—7 clades), A. donveae sp. nov., A. vazpintorum sp. nov. and A. praedicta sp. nov. Values are given as a range with mean values in parenthesis. M = male, F = female, n = sample size. Character A. bogerti A. wulfhaackei A. donveae sp. nov. A. vazpintorum A. praedicta sp. nov. (n=9) sp. nov. (n = 40) (n =17) sp. nov. (n = 48) (n = 5) Snout vent length (max) M50 mm M 58 mm M 59mm M 58 mm M 52 mm F 54 mm F 59mm F65 mm F 59mm ron Aggian Head Length/Head Width 1 34 O209 1A 0,14 BRN uc 6 2h ire Orbs 1.3+0,14 Snout Length/Eye Distance 1.6 + 0.34 1.9 + 0.36 2:0 $0.19 1.8 + 0.29 1.7+0.19 Snout Length/Eye-Ear Distance 1.2 + 0.07 Te? 5 6. Leah 30) Pa pare i1 7 Tels Or09 Precloacal pores (males) 8 9-12 (9.7) 11-12 (11.5) 9-11 (10.2) 8 (8.0) Ventral rows per tail verticil 4 (4.0) 4—5 (4.0) 5-6 (5.5) 5-7 (5.0) 4 (4.0) Dorsal rows per tail verticil 5 (5.0) 5-6 (5.1) 6-7 (6.6) 6-7 (6.1) 5 (5.0) Scales below 4* toe 6-9 (6.9) 6-9 (7.4) 6-8 (7.7) 6-10 (8.0) 9-11 (9.6) Mid-body scale rows 69-77 (73.5) 76-88 (79.3) 64-78 (72.8) 73-86 (80.3) 73-78 (74.8) Scales between eyes 11-14 (12.4) 11-16 (13.7) 11-14 (11.0) 11-15 (13.1) 12-15 (13.5) Scales: nostril to eye 8-12 (9.9) 8-11 (8.5) 8-11 (9.3) 7-11 (9.1) 9-10 (10.2) Scales: ear to eye 14-16 (15.4) 15-18 (16.0) 11-14 (11.9) 13-17 (15.6) 13-16 (14.8) Anterior nasals in contact 33% 68% 100% 100% 100% Suptalabials 8-10 (8.4) 8-11 (8.2) 8-10 (9.0) 8-10 (8.8) 8-10 (9.2) Infralabials 8-9 (8.3) 8-9 (8.4) 8-11 (9.3) 8-9 (9.1) 8-9 (8.5) the species descriptions below. The Angolan material can be broadly divided into two main morphological groups: 1) south-western (mostly) coastal group: Afroedura sp. 1 and Afroedura sp. 3, defined by higher precloacal pores in males (10—12), higher numbers of scale rows on ventral and dorsal surface per tail verticil (5 and 6, respectively), nasals always in contact and ventre immaculate, without pigmentation; 2) west-central inland/highland group: comprising species Afroedura sp. 4-7 and A. bogerti sensu stricto, defined by lower precloacal pores in males (8-10), lower numbers of scale rows on ventral and dor- sal surface per tail verticil (4 and 5, respectively), nasals zse.pensoft.net 62 William R. Branch et al.: New species of Afroedura for Angola Table 4. Tukey multiple comparisons of means of the four species combinations of Angolan Afroedura (values for A. wulfhaackei sp. nov. include the members of the morphologically-indistinguishable Afroedura sp. 5—7 clades). Bold values indicate significant pairwise comparisons. SVL — snout-vent length; HL — head length; HW — head width; ED — eye-diameter; EE — ear-eye length; IN — internostril distance; MSR — mid-body scale rows. Combinations SVL HL HW ED A. bogerti — A. donveae A. vazpintorum — A. donveae A. wulthaackei — A. donveae A. praedicta — A. donveae A. vazpintorum — A. bogerti A. wulfhaackei — A. bogerti A. praedicta — A. bogerti A. wulfhaackel — A. vazpintorum A. praedicta — A. vazpintorum A. praedicta — A. wulfhaackei in contact (68% of specimens) or separated by smaller granules (33% of specimens) and ventre pigmented with fine black specks. Morphologically, we could not sepa- rate members of the Afroedura sp. 4—7 clades, which are supported by close phylogenetic relationships. For that reason, we grouped the morphological data of the four clades together. The south-western coastal group can be further divided into a southern and northern group, based on average adult SVL (57.6 vs. 51.3 mm) and overall co- louration (see below). An isolated inland population on the Humpata plateau agrees with the northern lowland coastal group morphologically, with the exception of the darker pigmented ventre with an immaculate throat. The two west-central inland/highland groups are morpholog- ically very similar and differ only in the average num- ber of mid-body scale rows (73 vs. 79). An apparently isolated population on Serra da Neve (Afroedura sp. 2) agrees morphologically with this inland highland group in most aspects, but differs in that the nasals are always in contact (versus 33-68% of the time). The Tukey multi- ple comparisons of means showed significance differenc- es amongst most pairwise comparisons and support the above morphological separation (Table 4). Species descriptions In summary, both genetics and morphology (particularly diagnostic features, such as the nasal condition [separated or in contact], colouration [ventral pigmentation], snout vent length, number of mid-body scale rows, number of pre-cloacal pores, number of dorsal and ventral scale rows per tail verticil), as well as geographical segregation, suggest that most of the major genetic clades recovered should be considered as separate species. We, therefore, take the opportunity here to provide an updated descrip- tion for A. bogerti and describe the remaining clades as separate species. We apply the general lineage-based spe- cies concept, treating all populations that represent inde- pendent historical lineages supported by multiple differ- ent lines of evidence as listed above as separate species zse.pensoft.net P=0.004 P =0.273 P=0.771 P=0.262 P=0.298 P=0.031 P<0.001 P<0.001 P<0.001 P<0.001 P=0.671 P<0.001 P<0.001 P=0.035 P<0.001 P=0.264 P=0.004 P=0.642 P<0.001 P<0.001 P<0.001 P=0.015 P=0.396 P=0840 P=0.951 P=0.139 P<0.001 P<0.001 P=0.932 P=0.927 P=0.424 P=0.026 P=0.992 P=1.000 P< 0.001 P=0.942 P=0.926 P=0.767 P=0.020 P=1.000 P=0.868 P =1.000 P=1.000 P=1.000 P=1.000 P=0.187 P=1.000 P=0.993 P= 1.000 P=1.000 P=0.066 P=0.839 P=1.000 P=0.769 P=0.312 P<0.001 P=0.980 P=0.912 P=0.462 P=1.000 P=0.777 P=0.868 P< 0.001 P=0.984 P=0.967 P=0.782 P=1.000 P=0.987 P=0.195 P=1.000 EE IN Preanal MSR Pores Dorsal verticil scale rows P < 0.001 P < 0.001 P < 0.000 P < 0.001 P < 0.001 P=0.983 P = 1.000 P < 0.001 P < 0.001 P=0.972 Ventral verticil scale rows P=0.014 P =0.988 P=0.103 P<0.001 P=0.012 P<0.001 P<0.001 P =0.730 P=0.168 P<0.001 P=0.381 P<0.001 P=1.000 P=0.956 P=0.676 P=0.647 P=0.007 P=0.005 P=0.048 P=0.043 (de Queiroz 1998). Except for the nominal form, no his- torical names are available for Angolan material (Uetz et al. 2020). Afroedura bogerti Loveridge, 1944 Namba or Bogert’s Flat Gecko Osga-achadata da Namba, ou osga-achatada de Bogert Figures 5A, 6; Table 3 Note. The Namba Flat Gecko, Afroedura bogerti was de- scribed using a single male specimen collected by Harry and Alan Chapman in 1925 during the Vernay Expedition to Angola (Loveridge 1944; Branch et al. 2017b). Ac- cording to Loveridge (1944), the specimen was collected at ‘Namba (Mombolo)’, even though Bogert (1940), in his list of Angolan localities, mentioned only ‘Mom- bolo’ and gave rough co-ordinates for it. The collectors were the sons of a famous South African farmer, Wil- liam Chapman, who had hosted the scientific members of the Vernay Expedition on his properties. Within the Mombolo Region, William Chapman and his family had built or lived in three farms, Sandula (which he named Monte Victoria-Verdun and which later became Monte Verde), Chipepe (= Quipepe) and Namba (where a re- ligious mission was subsequently established — Missao da Namba). Branch et al. (2017b) provided a detailed systematic review of A. bogerti in Angola and, noting the uncertainties regarding the exact localities of Chap- man’s material ascribed to Namba/Mombolo, restricted the type locality of the nominal form of A. bogerti to a polygon encompassing his three farms and the village of Maka-Mombolo. However, this created a taxonom- ical problem, as the specimens from Chapman’s farms of Sandula and Namba represent two genetically-distinct clades. To resolve this issue, we re-instated the original type locality reference of ‘Namba’ and here restrict the type locality to the northern areas of Mombolo in the vi- cinity of Missaéo da Namba and, by default, the material from William Chapman’s farm of Sandula represents an undescribed species. Zoosyst. Evol. 97 (1) 2021, 55-82 Figure 4. Typical habitat of Angolan Afroedura. A. Namba area (A. bogerti),; B. Mt Sandula (Afroedura sp. 5); C. Serra da Neve 63 wa] (A. praedicta sp. nov.),; D. Omauha Lodge (A. donveae sp. nov.); E. Farm Mucungo (A. vazpintorum sp. nov.); F. Bimbe (A. vazpin- torum sp. nov.). Photos: A. Javier Lobon-Rovira; B, D, E. William R. Branch; C. Pedro Vaz Pinto; F. Ninda L. Baptista. Synonym. Afroedura karroica bogerti — Loveridge 1944:1; Afroedura bogerti — Onderstall 1984:506, Jacob- sen et al. 2014:467 (part), Branch et al. 2017b:157 (part), Marques et al. 2018: 177 (part), Branch et al. 2019a: 287 (part); Afroedura bogerti (clade 3) — Branch et al. 2017a:146—-147. Holotype. AMNH 47841, adult male, collected from Namba (Mombolo) (approx. -11.91417, 14.82083, 1827 m as.l.), Cuanza-Sul Province, Angola, by Harry and Allan (= Alan) Chapman, between September and November 1925 during the Vernay Expedition to Angola. Additional material examined. Females. PEM R24185*, collected from granite bedrock and out- crops 200 m south of old farm paddock, Farm Nam- ba (-11.91417, 14.82083, 1827 m a.s.l.), Cuanza-Sul Province, Angola, by William R. Branch, Pedro Vaz Pinto and Ninda L. Baptista on 3 November 2016; PEM R24187*, collected from rocks 400 m north of Misséo da Namba grounds (-11.91528, 14.84556, 1786 m a.s.l), Cuanza-Sul Province, Angola, by Wil- liam R. Branch, Pedro Vaz Pinto and Ninda L. Baptista on 4 November 2016; TM 46631-—34, collected from Numba (= Namba) (approx. -11.91722, 14.84417, 1808 m as.l.), Cuanza-Sul Province, Angola, by Wulf Haacke on 29 June 1974. Juveniles: PEM R24184*, PEM R24186*, collected from granite bedrock and outcrops 200 m south of old farm paddock, Farm Nam- ba (-11.91417, 14.82083, 1827 m a.s.l.), Cuanza-Sul Province, Angola, by William R. Branch, Pedro Vaz Pinto and Ninda L. Baptista on 3 November 2016. *ge- netically confirmed. Additional non-vouchered genetic material. JL- RZCO0015-6, collected from Namba (-11.88132, 14.76218, 1752 maz.s.l), Cuanza-Sul Province, Angola, by Pedro Vaz Pinto and Javier Lobon-Rovira on 11 February 2020. zse.pensoft.net 64 William R. Branch et al.: New species of Afroedura for Angola PEM R24232); C. Afroedura praedicta sp. nov. (holotype NB 854); D. Afroedura donveae sp. nov. (holotype PEM R17937); E. Afroedura vazpintorum sp. nov. (holotype PEM R24118); F. Afroedura vazpintorum sp. nov. (NB 0745); Photos: A, B, D, E. Wil- liam R. Branch; C, F. Pedro Vaz Pinto. General description. (description based on the adult material listed above). Head and body dorsoventrally compressed; SVL 46.4—53.5 (mean 50.0) mm, HL 10.9- 12.6 (mean 11.6) mm, HW 7.8—11.2 (mean 10.6) mm, broadest at posterior level of eye and 1.1—1.4 (mean 1.3) times longer than wide. Eyes large (2.14.3 [mean 2.8] mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, elongate at up- per anterior margins, upper posterior scales with small upward pointing spines. Snout rounded, 4.5—5.0 (mean 4.3) mm long, slightly longer than distance between eye and ear openings (3.6—-4.3 [mean 3.6] mm). Scales on top of snout smooth, rounded, scales to the side larger than central ones, with no intervening minute granules. Scales on snout slightly subequal in size to those on back of head or nape. Scales on eyelids larger than those on the crown. Circumorbital scales are separated by a row of smaller scales from the large scales on eyelid. Nostril pierced between rostral, two to three nasal scales and the 1“ supralabial; the supranasal being much larger zse.pensoft.net than the subequal postnasals and being separated from each other by one to two smaller scales. Nostrils slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to extensions into the nostril. Su- pralabials 8—10 (mean 8.4). Infralabials 8—9 (mean 8.3). A total of 11-14 (mean 12.4) scales across the crown at level of front of eyes. A total of 14-16 (mean 15.4) scales from front of ear to back of eye. Mid-body scale rows 69-77 (mean 73.5). Original tail slightly dorsoven- trally flattened and distinctly verticillate, with obvious lateral constrictions that are not that distinct to the tip of tail; each verticil comprising 5 imbricate rows of scales dorsally and 4 imbricate scale rows ventrally and with ventral scales approximately twice the size of those on the dorsal surface. Limbs well developed, hindlimbs slightly longer than forelimbs. All digits with a large pair of distal scansors, separated by a large, curved claw and followed after a large gap by a smaller pair of scan- sors; infero-median row of digital scales enlarged trans- versely, particularly towards the scansors, where the ter- Zoosyst. Evol. 97 (1) 2021, 55-82 65 Figure 6. Holotype of Afroedura bogerti (AMNH 47841) from Namba (Mombolo), Cuanza-Sul Province, Angola. Photos: Luis M.P. Ceriaco. minal scale adjoining the first pair of scansors may be medially constricted, swollen and scansor-like, enlarged subdigital lamellae on 4" toe 6—9 (mean 6.9). Precloacal pores 8. Colouration. /n life (based on PEM R24187, Fig. 5A). Greyish above with six irregularly spaced darker grey crossbars from the occiput to the sacrum, central cross- bars fused to form an X-shape; head with a dark grey band across the posterior edge of crown encompassing a small central pale spot; dark grey to black bar from nostril to the anterior margins of the ear opening; a vague, thin pale canthal stripe, extends on both sides from the nasal re- gion to anterior margins of eye, continuing posteriorly of the eye to above the ear opening; upper and lower labials grey with diffuse yellow edges; lateral sides of the body with very faintly yellow-mustard scales scattered amongst more prominent darker grey scales; limbs greyish above with scattered darker grey markings with intervening faint yellow-mustard colouration; tail (regenerated) with irreg- ular grey mottling; iris golden with a black narrow elliptic pupil with crenulated edge and black reticulation; ventre uniform greyish with scattered black specks; ventral limbs with scattered black specks, more prominent than on the ventrum. /n preservative (based on PEM R24187): dorsum with six irregularly-spaced darker grey crossbars from the occiput to the sacrum with beige intervening blotches, central crossbars fused to from dark grey X-shape; ventre beige with numerous small scattered black specks on each scale, more prominent anteriorly, laterally and posteriorly. Variation. Greyish to brownish above with five to six ir- regularly-spaced darker grey-brown W-shaped crossbars from the occiput to the sacrum, sometimes fused in the middle to form X-shapes, limbs and tail with grey blotch- es; ventre uniform greyish with scattered black specks. Juveniles with more sharply-defined pattern. The origi- nal description of the holotype (Fig. 6) was based on a preserved and bleached specimen: ‘Above, greyish; back with five or six obsolescent, irregularly W-shaped brown crossbars; limbs and tail immaculate. Below, whitish, uniform’ (Loveridge 1944). Examination of images of the holotype confirm the presence of fine brown coloured specks ventrally that indicates that the ventre was not im- maculate as originally described. Natural history and habitat (Fig. 4A). An exclusive- ly rupicolous species sheltering in crevices and under flakes of exfoliating rock amongst large granite boulders during the day. Individuals were observed at night forag- ing in vertical rock faces of large granite boulders. The surrounding habitat was mostly montane grasslands and some cultivated fields, but also included stunted altitude miombo and some Afromontane forest elements in steep- er gorges and between boulders. It should be noted that the southern face of Namba Mountain contains the most extensive and well-preserved Afromontane forest patches in Angola, recognised as an important biodiversity reser- voir and regional hotspot (Mills et al. 2013). zse.pensoft.net 66 Distribution and conservation. This species complex was previously considered to be widespread in granite boulders throughout south-western Angola (Branch et al. 2017b; Marques et al. 2018). Here, we show that it 1s restricted to the Namba Region in the southern parts of Cuanza-Sul Province, Angola (Fig. 1). Namba consists of a 25 km west-east orientated mountain range that climbs to an elevation of 2,420 m a.s.l. and towers southwards above the highland plateau of Mombolo. Our new series was collected at elevations of 1,750—1,850 maz.s.l., at the base of the south-facing slopes of the Namba Mountain chain, but the species probably occurs in suitable habitat higher up and possibly well above 2,000 m a.s.l. (Fig. 1). Considering the deep genetic divergence between this species and Afroedura present at the southern edge of Mombolo and the recognition of Namba as a biodiver- sity hotspot, it seems plausible for this species to have evolved in isolation on this mountain ‘island’. Neverthe- less, A. bogerti may also extend its distribution further north into other poorly-studied mountain ranges in Cuan- za-Sul Province (Fig. 1). The species seems to be rela- tively common in the Namba Mountains, but its biology remains poorly known. Afroedura wulfhaackei sp. nov. http://zoobank.org/681544BA-AA5F-473B-90BD-B2790838BC81 Angolan Flat Gecko Osga-achatada de Angola Figures 5B, 7; Tables 3, 5 Note. The phylogenetic analysis identified four well-de- fined clades (Afroedura sp. 4—7) within the southern in- land/highland major clade (see Results). Our data show that this southern major clade clearly represents at least one undescribed species of Afroedura. Despite compar- atively-low genetic distances between the four clades recovered (Table 2), they were all retrieved separately in our phylogenetic analyses and, as our morphological examination failed to separate the four clades, we cannot assign them as separate species. Therefore, we decided to take a conservative approach and to describe only one of the clades as a new species for now and tentatively assign Afroedura sp. 5—7 to this species pending further work. Thus, we grouped the material according to these clades and selected the clade called here Afroedura sp. 4 (Fig. 2) as the name-bearing clade for our new species. Material from lower-altitude localities near Bocoio in Benguela Province groups morphologically with our new species, but needs to be tested in a phylogenetic framework. Synonym. A/roedura bogerti — Branch et al. 2017b:157 (part); Marques et al. 2018: 177 (part); Branch et al. 2019a: 287 (part); Afroedura bogerti (clade 4) — Branch et al. 2017a:147. Holotype. PEM R24234, adult male, collected 3 km north of Maka-Mombolo (-12.17056, 14.88167, 1756 m zse.pensoft.net William R. Branch et al.: New species of Afroedura for Angola a.s.l.), Benguela Province, Angola, by William R. Branch and Pedro Vaz Pinto on 15 November 2016. Paratypes. PEM R24232-3, adult females, collected 3 km north of Maka-Mombolo (-12.17056, 14.88167, 1756 m a.s.l), Benguela Province, Angola, by William R. Branch and Pedro Vaz Pinto on 15 November 2016; PEM R24236, adult male, collected above Maka-Mombolo, north-east of Balombo (-12.19833, 14.86833, 1857 m a.s.l.), Benguela Province, Angola, by William R. Branch and Pedro Vaz Pinto on 15 November 2016. Additional referred material (not examined). NB 817-9, collected at Morro do Moco, camp near Canjonde (-12.42611, 15.14778, 1931 m a.s.l), Huambo Province, Angola, by Pedro Vaz Pinto on 13 November 2017 (ge- netic samples included in this study). Tentative referred additional material examined. Afroedura sp. 5 (Males): PEM R24192-4, collected at William Chapman’s Farm Victoria-Verdun (Sandula), (-12.17194, 15.02667, 1834 m a.s.l.), Cuanza-Sul Prov- ince, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 6 November 2016. Afroedu- ra sp. 5 (Females): PEM R24190-1, PEM R24195-6, PEM R24199, collected at William Chapman’s Farm Victoria- Verdun (Sandula), (-12.17194, 15.02667, 1834 m as.l.), Cuanza-Sul Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 6 No- vember 2016. Afroedura sp. 5 (Juveniles): PEM R24197— 8, collected at William Chapman’s Farm Victoria- Verdun (Sandula), (-12.17194, 15.02667, 1834 m a.s.l.), Cuan- za-Sul Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 6 November 2016. Afro- edura sp. 6 (Males): PEM R24201, collected near Rio Chicanda, 5 km southwest of Lepi (-12.90861, 15.36472, 1534 m a.s.l.), Huambo Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 8 No- vember 2016. Afroedura sp. 7 (Males): TM 45382, TM 45387-8, TM 45392, TM 45396, collected at Candumbo Rocks, 16 km west of Vila Nova = Huambo (-12.73614, 15.97442, 1760 m asl), Huambo Province, Angola, by Wulf Haacke on 11 May 1971. Afroedura sp. 7 (Fe- males): PEM R22490-1, collected 1 km west of Candum- bo on road to Boas Aguas (-12.73614, 15.97442, 1760 m a.s.1), Huambo Province, Angola, by Luke Verburgt on 11 March 2016; TM 45383, TM 45390, TM 45393-4, TM 45397, collected at Candumbo Rocks, 16 km west of Vila Nova = Huambo (-12.73614, 15.97442, 1760 m a.s.l), Huambo Province, Angola, by Wulf Haacke on 11 May 1971. Afroedura sp. 7 (Juveniles): PEM R24200, collect- ed at Candumbo Rocks Memorial, 20 km east Humana to Cuito (-12.73722, 15.97333, 1750 maz.s.1), Huambo Prov- ince, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 7 November 2016; TM 45386, TM 45389, TM 45391, TM 45395, collected at Candumbo Rocks, 16 km west of Vila Nova = Huambo (-12.73614, 15.97442, 1760 m a.s.l), Huambo Province, Angola, by Zoosyst. Evol. 97 (1) 2021, 55-82 67 Figure 7. Holotype of Afroedura wulfhaackei sp. nov. (PEM R24234) from Maka-Mombolo, Benguela Province, Angola. Photos: Werner Conradie. Wulf Haacke on 11 May 1971. Unassigned clades (Fe- males): TM 45374, collected 1 km south of Luimbale (-12.25367, 15.31694, 1591 m a.s.l.), Huambo Province, Angola, collected by Wulf Haacke on 10 May 1971; TM 45367-8, collected 10 km west of Soque (-12.34590, 15.01180, 1974 ma.s.1), Benguela Province, Angola, col- lected by Wulf Haacke on 10 May 1971; PEM R24743, collected at Morro do Pundo (-12.44389, 13.92250, 939 ma.s.l.), Benguela Province, Angola, by Pedro Vaz Pinto on 6 June 2018; TM 46587-8, TM 465890, collected 3 km west of Bocoio (-12.46605, 14.10694, 926 m a.s.1.), Benguela Province, Angola by Wulf Haacke on 25 May 1971. Unassigned clades (Males): TM 45366 collected 10 km west of Soque (-12.34590, 15.01180, 1974 maz.s.l), Benguela Province, Angola, by Wulf Haacke on 10 May 1971, TM 46589, adult male, collected 3 km west of Bo- colo (-12.46605, 14.10694, 926 ma.s.l.), Benguela Prov- ince, Angola, by Wulf Haacke on 25 May 1971. Tentative referred additional material not exam- ined. TM 45381, 45384—5, TM 45398, collected at Can- dumbo Rocks, about 25 km east of the city of Huambo (-12.73614, 15.97442, 1760 m a.s.l), Huambo Prov- ince, Angola, by Wulf Haacke on 11 May 1971 (placed in Afroedura sp. 7, based on same geographical area as genetically-assigned material); FKH 0239, collected at Monte Verde-Sandula (-12.17924, 15.03086, 2055 m a.s.l.), Cuanza-Sul Province, Angola, by Pedro Vaz Pinto on 29 May 2019 (genetic sample included in this study placed it in Afroedura sp. 5). Etymology. The new species is named in honour of Wulf Haacke, retired curator of the herpetology collection at the former Transvaal Museum (now Ditsong National Museum of Natural History). His herpetological expedi- tions to Angola in the early 1970s paved the way for this study and much of the material used in this study resulted from his expeditions. The name is constructed in the mas- culine singular genitive. Diagnosis. A member of the greater ‘transvaalica’ group in possessing two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail (Jacobsen et al. 2014). It is part of the A. bogerti-group which differs from other members of the ‘transvaalica’ group by having less than 88 mid-body scale rows (vs. 97-102 in A. gorongo- sa, 113—120 in A. loveridgei, 102-119 in A. transvaali- ca); by the rostral bordering the nostril (nostril excluded from rostral in A. /overidgei); by the anterior nasals being mostly in contact ~ 68% (separated by 1-3 granules in A. gorongosa; always in broad contact in A. loveridgei, usually in broad contact in A. transvaalica ~ 3—18%); and in having 11—16 scales between the anterior borders of the eyes (19-22 in A. gorongosa; 15-19 in A. loveridgei; zse.pensoft.net 68 15—20 in A. transvaalica) (comparative data fide Branch et al. 2017a). Afroedura wulfhaackei sp. nov. differs from other members of the A. bogerti-group by a combination of the following characters (see Tables 3, 4): 76-88 (mean 79.3) mid-body scale rows (69-77 [mean 73.5] in A. bogerti, 64—78 [mean 72.8] in A. donveae sp. nov., 73—86 [mean 80.3] in_A. vazpintorum sp. nov., 73-78 [mean 74.8] in A. praedicta sp. nov.); by the anterior nasals being mostly (~ 68% of the time) in contact (~ 33% of the time in contact in A. bogerti; always in contact in A. donveae sp. nov., A. vazpintorum sp. nov. and A. praedicta sp. nov.); each verticil comprising 4—5 (mean 4.0) ventral and 5—6 (mean 5.1) dorsal rows of scales (4 and 5 in A. bogerti and A. praedicta sp. nov.; 5—6 [mean 5.5] and 6—7 [mean 6.6] in A. donveae sp. nov.; 5—6 [mean 5.0] and 6—7 [mean 6.1] A. vazpintorum sp. nov.); ventral surfaces greyish with scattered small black spots (similar to 4. bogerti and A. praedicta sp. nov., immaculate in A. donveae sp. nov. and A. vazpintorum sp. nov.). Afroedura wulfhaackei sp. nov. differs more specifically from its sister highland species A. bogerti in having a higher number of mid-body scale counts (76—88 [mean 79.3] versus 69-77 [mean 73.5]) and differs from A. praedicta sp. nov. in that the nasals are separated by smaller granules (versus always in contact). Holotype description. Adult male; SVL 51.4 mm; tail 47.1 mm (full original tail), with a small mid-ventral incision for the removal of liver sample. Measurements and meristic characters of holotype presented in Table 5. Head and body dorsoventrally compressed; HL 11.9 mm, HW 9.0 mm, broadest at posterior level of eye and 1.32 times longer than wide. Eye large (2.3 mm wide), pu- pil vertical with indented margins; circumorbital scales small and smooth, elongate at upper anterior margin, upper three posterior scales with small upward pointing spines. Snout rounded, 4.9 mm long, slightly longer than distance between eye and ear openings (4.0 mm). Scales on top of snout smooth, rounded, scales to the edge larger than central ones, with no intervening minute granules. Scales on snout slightly subequal in size to those on the back of head or the nape. Scales on eyelids larger than those on the crown, five scales deep from circumorbital scale to crown. Circumorbital scales are separated by a row of smaller scales from the larger scales on eyelids. Nostril pierced between rostral and three nasal scales; 1“ supralabial narrowly excluded from nostril; the suprana- sal being much larger than the subequal postnasals and separated from each other by two smaller scales. Nos- trils slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to extensions into the nostril. Eight supralabials on each side, the labial margin flexing upwards at the rictus (approx. mid-orbital posi- tion), with 3—4 minute scales proximal to the flexure. Ten infralabials on either side, with a small scale proximal to the flexure. At the lip, mental slightly narrower than adjacent infralabial, only three quarters the width of ros- tral and in contact with two distinctly elongate postmen- zse.pensoft.net William R. Branch et al.: New species of Afroedura for Angola tal scales. Scales on throat much smaller than those on belly; scales touching infralabials larger. Fifteen scales across the crown at level of front of eyes; 16 scales from ear to eye; 83 scales around mid-body. Ear opening deep, oblique and roughly oval, only half as high as wide (0.3 x 0.5 mm). Scales on dorsum smooth, non-overlapping, largest at mid-body, smaller on nape and tail base. Scales on ventrum flattened, not overlapping, more-or-less ovate at mid-ventrum, twice the size of lateral granules and 1.5 times those along the backbone. Original tail slightly dor- soventrally flattened and distinctly verticillate (11 verti- cils in total), with obvious lateral constrictions that are not that distinct to tip of tail; each verticil comprising 5 imbricate rows of scales dorsally and 4 imbricate scale rows ventrally and with ventral scales approximately twice the size of those on the dorsal surface. Limbs well developed, hindlimbs slightly longer than forelimbs, mite pockets (dermal crevices inhabited by small ectoparasitic mites) only present at anterior margin of hindlimbs. All digits with a large pair of distal scansors, separated by a large, curved claw and followed by a large gap (twice the length of terminal scansor) by a smaller pair of scansors; infero-median row of digital scales enlarged transverse- ly, particularly towards the scansors, where the terminal scale adjoining the first pair of scansors may be medially constricted, swollen and scansor-like; enlarged subdigital lamellae on 4" toe 9. Precloacal pores 9. Paratype variation (see Table 5 for more measure- ments and scale counts of type series). SVL varied from 49.1-55.7 mm; head length 1.34-1.45 times the head width; snout 1.99 times the diameter of eye. Supranasals always separated by smaller granules, usually with a sin- gle large granule in contact with the rostral between the Table 5. Measurements (in mm) and scale counts for the type series of Afroedura wulfhaackei sp. nov. Catalogue number PEM R24234 PEM R24232 PEM R24233 Type Status Holotype Paratype Paratype Sex Male Female Female Snout vent length 51.4 49.1 55:7. Tail length 47.1 35.4 “B/D Tail condition Original Regenerated Regenerated Head length TS 10.9 123 Head width 9G LS 8.5 Snout length 4.9 4.5 5.1 Eye distance Ze 2.2 2.6 Eye-Ear distance 4.0 3.5 4.3 Precloacal pores (males) 9 - - Dorsal rows per tail verticil 5 - 5 Ventral rows per tail verticil 4 - 4 Scales below 4'" toe S) 8 7 Mid-body scale rows 83 82 76 Scales between eyes 15 13 14 Scales: nostril to eye 7 7 9 Scales: ear to eye 16 14 17 Anterior nasals in contact No No No Supralabials 9 7 9 Infralabials 8 9 8 Zoosyst. Evol. 97 (1) 2021, 55-82 supranasals, followed by 1—2 smaller granules in lateral contact; the first upper labial and rostral always enters the nostril and the width of the rostral at the lip margin is always wider than that of the mental; 2-3 postmental scales; supralabials 7—9, infralabials 8—9; scales between anterior edges of eyes 13—14; scales between nostril and anterior edge of orbit 7-9; scales between anterior edge of ear and rear margin of orbit 14-17; scales around mid- body 76-82; subdigital lamellae on 4 toe 7-8; dorsal scales per tail verticil 5; ventral scales per tail verticil 4. Precloacal pores 9. Additional material variation. SVL 36.4—59.6 mm; original tail length 36.4—-57.8 mm, 0.94 times SVL; head length 1.19-1.75 times head width; snout 1.91 times diameter of eye. The supranasals in contact in 22 spec- imens and separated by granules in eight specimens, usu- ally with a single large granule in contact with the rostral between the supranasals, followed by 1—2 smaller gran- ules in lateral contact; the first upper labial and rostral always enters the nostril and the width of the rostral at the lip margin is always wider than that of the mental; 2—4 postmental scales; supralabials 8—9; infralabials 8—9; scales between anterior edge of eye 12-16; scales be- tween nostril and anterior edge of orbit 7-11; scales be- tween anterior edge of ear and rear margin of orbit 15—18; scales around mid-body 75-88; subdigital lamellae on 4" toe 7—9; dorsal scales per tail verticil 5 (TM 45366, TM 46588 and PEM R24743 with 6); ventral scales per tail verticil 4 (TM 46588 with 5). Precloacal pores 9-12. Colouration. /n /ife (paratype PEM R24232, Fig. 5B): Greyish above with five irregularly-spaced darker cross- bars from the occiput to the sacrum, each crossbar con- sisting posteriorly out of three to four black scales wide forming a W-shape; anterior to W-shape are 8-10 scales deep with a mix of dark grey and mustard colours; each dark crossbar separated by light grey to beige blotches; head with irregular dark grey-mustard blotches on the crown with intervening light grey colouration; dark mus- tard to dark grey bar from nostril to the anterior margins of the ear opening; a vague, thin pale grey canthal stripe, extends on both sides from the nasal region to anterior margins of eye; upper and lower labials grey with diffuse mustard edges; lateral sides of the body with a mix of dark grey and yellow-mustard colouration; limbs greyish above with scattered darker grey markings with interven- ing yellow-mustard colouration; tail (regenerated) with irregular grey-mustard mottling; iris golden with a black narrow elliptic pupil with crenulated edge and black re- ticulation with light grey intervening blotches; ventrum uniform greyish with scattered black specks; ventral limbs with scattered black specks, more prominent than on the ventrum. /n preservative (holotype PEM R24234, Fig. 7): Dorsum with five irregularly-spaced dark grey W-shaped crossbars from the occiput to the sacrum with beige inter- vening blotches; ventrum is beige with numerous small scattered black specks on each scale, more prominent pos- 69 teriorly. Variation: Greyish to brownish above with five to six irregularly-spaced darker grey-brown W-shaped cross- bars from the occiput to the sacrum, limbs and tail with grey blotches; ventrum uniform greyish with scattered black specks. Juveniles have sharper patterns and colours. Natural history and habitat (Fig. 4B). An exclusively rupicolous species living in crevices between rocks or un- der flakes of exfoliating rocks in boulders at elevations of 920—2,055 m a.s.l. Specimens were collected in cracks of relatively small- to medium-sized boulders of carbonatitic origin surrounded by montane grassland. Some individu- als were also found under flakes in large granite boulders and often in steep vertical rock faces and overhangs. Distribution and conservation. This species is current- ly known from southern Cuanza-Sul, central Huambo and northern Benguela Provinces, Angola (Fig. 1). It ap- pears to have a relatively large, but patchy, distribution on the Angolan highlands and may extend its range into neighbouring provinces. Although sometimes locally common, it appears to be absent from vast areas in-be- tween, where the species would be expected to occur. Populations in isolated granite outcrops may be threat- ened by removal of rock flakes for construction of homes and other buildings. Afroedura praedicta sp. nov. http://zoobank.org/FAECE4F2-AC75-4E57-8B78-D76FFA8C282D Serra da Neve Flat Gecko Osga-achatada da Serra da Neve Figures 5C, 8; Tables 3, 6 Holotype. NB 854, adult male, collected from Serra da Neve (-13.77354, 13.24825, 1944 m a.s.l.), Namibe Prov- ince, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo Antonio on 30 November 2017. Paratypes. ZMB 91607 (NB 853), NB 855, adult males, collected from Serra da Neve (-13.77354, 13.24825, 1944 m_a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo Antonio on 30 November 2017. ZMB 91608 (NB 1053), NB 1054, adult females, collected from Serra da Neve (-13.77354, 13.24825, 1944 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo Antonio on 30 November 2017. NB 1055, juvenile, collected from Serra da Neve (-13.77354, 13.24825, 1944 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo Antonio on 30 November 2017. Etymology. The specific epithet reflects the earlier pre- diction by WRB of the potential existence of an isolated population of Afroedura at Serra da Neve. We use the specific epithet “praedicta’, the Latin participle mean- ing predicted or anticipated, formed in the feminine gen- itive to match the gender of Afroedura. zse.pensoft.net 70 William R. Branch et al.: New species of Afroedura for Angola Figure 8. Holotype of Afroedura praedicta sp. nov. (NB 854) from Serra da Neve, Namibe Province, Angola. Photos: Telmo Antonio. Diagnosis. A member of the greater ‘transvaalica’ group as it possesses two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail (Jacobsen et al. 2014). Part of the A. bogerti-group which differs from other members of the ‘transvaalica’ group by having less than 78 mid-body scale rows (vs. 97-102 in A. gorongo- sa, 113—120 in A. loveridgei, 102—119 in A. transvaalica), by the rostral bordering the nostril (nostril excluded from rostral in A. /overidgei); by the anterior nasals always be- ing in contact (separated by 1—3 granules in A. gorongosa; always in broad contact in A. /overidgei; usually in broad contact in A. transvaalica ~ 3—18%); and in having 12—15 scales between the anterior borders of the eyes (19-22 in A. gorongosa, 15-19 in A. loveridgei, 15—20 in A. trans- vaalica) (comparative data fide Branch et al. 2017a). Afroedura praedicta sp. nov. differs from other mem- bers of the 4. bogerti-group by a combination of the fol- lowing characters (see Tables 3 and 4): 73—78 (mean 74.4 mid-body scale rows (69-77 [mean 73.5] in A. bogerti, 76-88 [mean 79.3] in A. wulfhaackei sp. nov., 64-78 [mean 72.8] in A. donveae sp. nov., 73—86 [mean 80.3] in A. vazpintorum sp. nov.); by the anterior nasals always being in contact (similar to A. donveae sp. nov. and A. vazpintorum sp. nov.; ~ 33% of the time in contact in A. bogerti, ~ 68% of the time in contact in A. wulfhaackei Sp. nov.); each verticil having 4 ventral and 5 dorsal rows of scales (similar to the 4 and 5 in A. bogerti, 4—S [mean zse.pensoft.net 4.0] and 5-6 [mean 5.1] in A. wulfhaackei sp. nov.; but lower than the 5—6 (mean 5.5) ventral and 6—7 (mean 6.6) in A. donveae sp. nov. and 5—6 (mean 5.0) and 6—7 (mean 6.1) in A. vazpintorum sp. nov.); ventral surfaces grey with black specks on scales (similar to A. bogerti and A. wulfhaackei sp. nov.; immaculate in A. donveae sp. nov. and A. vazpintorum sp. nov.). Afroedura praedicta sp. nov. also differs from its sister highland species A. bogerti sp. nov. and A. wulfhaackei sp. nov. in that the nasals are always in direct contact (versus mostly separated). Holotype description. Adult male; SVL 51.6 mm; tail 37.0 mm (regenerated tail, except for the first verticil), with a small mid-ventral horizontal incision for the re- moval of liver sample. Measurements and meristic char- acters of holotype presented in Table 6. Head and body dorsoventrally compressed; HL 12.9 mm, HW 7.9 mm, broadest at posterior level of eye and 1.6 times longer than wide. Eye large (2.6 mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, elongated at upper anterior margin, the most upper poste- rior scale with very small upward pointing spines. Snout rounded, 5.0 mm long, slightly larger than the distance between eye and ear openings (4.2 mm). Scales on top of snout slightly granular and elevated, rounded, mostly equal in size, with no intervening minute granules. Scales on snout slightly subequal in size to those on back of Zoosyst. Evol. 97 (1) 2021, 55-82 71 Table 6. Measurements (in mm) and scale counts for the type series of Afroedura praedicta sp. nov. Catalogue number NB 854 ZMB 91607 Type status Holotype Paratype Sex Male Male Snout vent length (max) 5G 51.4 Head length 1:29 12.4 Tail length 37.0 29:2 Tail condition Regenerated Regenerated Head width re) 723 Snout length 5.0 4.8 Eye distance 2.6 27 Eye-ear distance 4.2 3.9 Precloacal pores (males) 8 8 Dorsal rows per tail verticil 4 4 Ventral rows per tail verticil 5 5 Scales below 4'" toe 9 11 Mid-body scale rows 74 75 Scales between eyes 12 13 Scales: nostril to eye 9 11 Scales: ear to eye 16 15 Anterior nasals in contact Yes Yes Supralabials 8 10 Infralabials 1 i] head or nape, which in turn is irregular in size and mostly smooth. Scales on eyelids larger than those on the crown, six scales deep from circumorbital scales to crown. Cir- cumorbital scales are separated by a row of smaller scales from the larger scales on eyelid. Nostril pierced between rostral, three nasal scales; 1** supralabial in contact with nostril; the supranasal being much larger than the sub- equal postnasals and are separated from each other by two smaller scales. Nostrils slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to ex- tensions into the nostril. Eight supralabials on each side, the labial margin flexing upwards at the rictus (approx. mid-orbital position), with 2-3 minute scales proximal to the flexure. Nine infralabials on either side, with a small scale proximal to the flexure. At the lip, mental slightly narrower than adjacent infralabial, only three quarters the width of rostral and in contact with three distinctly elon- gate postmental scales. Scales on throat much smaller than those on belly; scales touching infralabials larger. Twelve scales across the crown at level of front of eye; 9 scales between nostril and front of eye; 16 scales from ear to eye; 74 scales around mid-body. Ear opening deep, oblique and roughly round, backward pointing, nearly equal as long as wide (0.7 x 0.6 mm). Scales on dorsum smooth, non-over- lapping, largest at mid-body, smaller on nape and tail base. Sales on ventre flattened, not overlapping, more-or-less ovate at mid-ventrum and twice the size of lateral gran- ules and 1.5 times those along backbone. Tail regenerated, except for the first verticil, with obvious lateral constric- tion before regenerated tail start; first verticil comprising 5 imbricate rows of scales dorsally and 4 imbricate scale rows ventrally and with ventral scales approximately twice the size of those on the dorsal surface. Limbs well developed, hindlimbs slightly longer than forelimbs, both without obvious mite pockets at posterior margin of limb insertions, mite pockets present at anterior margin of hind- limbs. All digits with a large pair of distal scansors, sep- NB 855 ZMB 91608 NB 1054 NB 1055 Paratype Paratype Paratype Paratype Male female female Juvenile 49.6 5 al 46.0 28.0 12.8 10.1 10.0 G23 — 43.4 - 2251 Truncated Regenerated Truncated Original 6.8 6.7 6.3 4.7 4.8 4.2 4.3 2.5 eal 2.4 2.3 1.8 4.8 3.8 Say 2.4 4 - 4 - 5 - 5 10 9 - 2) 74 75 3 78 12 14 1-5 15 10 9 #1. It 15 Li 15 13 Yes Yes Yes Yes 8 10 9 10 i] 8 8 8 arated by a large, curved claw and followed after a large gap (twice the length of terminal scansor) by a smaller pair of scansors; infero-median row of digital scales enlarged transversely, particularly towards the scansors, where the terminal scale adjoining the first pair of scansors may be medially constricted, swollen and scansor-like; 9 enlarged scale rows under 4" toe. Precloacal pores 8. Paratypes variation (see Table 6 for more measure- ments and scale counts of type series). SVL varied from 46.0-51.6 mm; head length 1.09-1.46 times head width; snout 1.71 times diameter of eye. The supranasals always in contact; the first supralabial enters the nostril (except for ZMB 91607 and NB 855) and rostral always enters the nostril and the width of the rostral at the lip margin is always wider than that of the mental; 2—3 post- mental scales; supralabials 8—10, infralabials 8—9; scales between anterior edge of eye 13-15; scales between nos- tril and anterior edge of orbit 9-11; scales between ear and eye 13-16; scales around mid-body 73-78; subdigital lamellae on 4" toe 9-11; dorsal scales per tail verticil 5; ventral scales per tail verticil 4; precloacal pores 8. Colouration. Jn life (holotype NB 854, Fig. 5C). Dark grey above with random yellow-olive scattered scales, with six irregularly-spaced, darker grey to black W-shaped crossbars, first one broken up into three blotch- es, each crossbar anteriorly bordered by a row of light white to yellow scales; head mostly grey with scattered darker grey and yellow scales, dark black bar from nos- tril to posterior of eye, continuing anteriorly of the eye to ear opening; no thin pale white canthal stripe; upper and lower labials dark grey with diffuse lighter grey edg- es; lateral sides of the body with a mix of dark grey and yellow blotches; limbs darker grey above with scattered yellow scales; tail (regenerated) with dark black blotches and irregular white to light grey mottling; iris dark brown zse.pensoft.net 72 to golden with a narrow black elliptic pupil with crenu- lated edge and black reticulation; ventre uniform greyish with scattered black specks; ventral surface of limbs with scattered black specks, more prominent than on the ven- trum. /n preservative (holotype NB 854, Fig. 8): dorsum with five distinct, irregularly-spaced, dark grey W-shaped crossbars anteriorly, with beige intervening blotches, the posteriorly crossbar on the nape are broken up dark grey blotches; dorsally, the arms and legs are beige with irregular darker grey blotches; tail (regenerated part) with light grey to white mottling on darker grey to black background; dorsally, the head has mottled dark brown scales, dark grey bar running from the nasals through eye to anterior of the ear opening; supralabials dark brown- edged ventrally; infralabials scattered with dark brown markings dorsally; ventrum uniform greyish with scat- tered black specks; ventrally, limbs with scattered black specks, more prominent than on the ventrum. Variation: Similar colouration and patterning as the holotype (pre- served) and paratype (life). Dorsal dark W-shaped cross- bars number 5—6. Regenerated tails with fine dark brown to black mottling. Juveniles with more sharply-defined pattern and darker colouration. Natural history and habitat (Fig. 4C). A rupiculous species found inside deep crevices and fractured rocks in large boulders of extrusive origin. All specimens were found on one site between 1,900—2,000 m a.s.1. The sur- rounding vegetation included altitude dwarf miombo and montane elements. Found in syntopy with Cordylus pho- nolithos at Serra da Neve. Distribution and conservation. This species is known only from the alkaline mountain complex of Serra da Neve (Fig. 1). Afroedura donveae sp. nov. http://zoobank.org/B2939E 10-5F5D-45D0-9CBF-352D 1 D8A6B9A Iona Flat Gecko Osga-achatada do Iona Figures 5D, 9; Tables 3, 7 Synonym. Afroedura bogerti — Haacke 2008:6, Huntley 2009:84, Rosler 2000:57, Barts and Haacke 2010:39, Ja- cobsen et al. 2014:456 & 468 (part), Branch et al. 2017b:157 (part), Marques et al. 2018: 177 (part), Branch et al. 2019a: 287 (part); Afroedura cf. bogerti— Agarwal et al. 2017:649; Afroedura bogerti (clade 1) — Branch et al. 2017a:146. Holotype. PEM R17937, adult female, collected from Omauha Lodge, 15 km south of Tambor (-16.20061, 12.40183, 341 m a.s.l.), Namibe Province, Angola, by William R. Branch, Werner Conradie, Krystal Tolley and John Measey on 18 January 2009. Paratype. PEM R17936, collected from Omauha Lodge, 15 km South Tambor (-16.20061, 12.40183, 341 m a.s.l), Na- zse.pensoft.net William R. Branch et al.: New species of Afroedura for Angola mibe Province, Angola, by William R. Branch, Werner Con- radie, Krystal Tolley and John Measey on 18 January 2009. Additional material examined. Males: TM 40508, TM 40512, TM 40516, TM 40518, collected from Tambor (-16.06667, 12.43333, 355 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 1 April 1971. Females: PEM R18041-2, collected 0.5 km south of Tambor (-16.07414, 12.43328, 352 m as.l.), Namibe Province, Angola, by William R. Branch, Krystal Tolley and John Measey on 23 January 2009; TM 40509-11, TM 40513-—5, TM 40517, TM 40536-8, collected from Tambor (-16.06667, 12.43333, 355 m as.l.), Namibe Province, Angola, by Wulf Haacke on | April 1971. Additional referred material (not examined). TM 40519-20, collected from Tambor (-16.06667, 12.43333, 355maz.s.l.), Namibe Province, Angola, by Wulf Haacke on 1 April 1971; FKH 0341-2, collected from Omauha Lodge (-16.20061, 12.40183, 338 ma.s.l.), Namibe Province, An- gola, by Pedro Vaz Pinto on 3 October 2019; CAS:HERP 263012-3, collected from Omauha (-16.19858, 12.40073, 338 m as.l.), Namibe Province, Angola, by Luis M.P. Ceriaco, Suzana Bandeira and Isham Agarwal on 25 and 27 November 2017; CAS:HERP 248780-1, collected 0.5 km south of Tambor (-16.07414, 12.43328, 352 m a.s.l.), Namibe Province, Angola, by William R. Branch, Krystal Tolley and John Measey on 23 January 2009. Etymology. This gecko is named after Donvé Branch, WRB’s wife, with the following personal quote: “This, the most beautiful of all the Angolan flat geckos, is named for my wife, Donvé Branch (‘Dove’) who bore the long periods I was away on fieldwork, and to whose nest I re- turned, and surrounded me with love until the end”. The name is constructed in the feminine singular genitive. Diagnosis. A member of the greater ‘transvaalica’ group as it possesses two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail (Jacobsen et al. 2014). Part of the A. bogerti-group which differs from other members of the ‘transvaalica’ group by having less than 78 mid-body scale rows (vs. 97-102 in A. gorongo- sa, 113-120 in A. loveridgei, 102—119 1n_A. transvaalica); by the rostral bordering the nostril (nostril excluded from rostral in A. /overidgei); by the anterior nasals always in contact (separated by 1-3 granules in A. gorongosa; al- ways in broad contact in A. /overidgei; usually in broad contact in A. transvaalica ~ 3—-18%); and in having 11-14 scales between the anterior borders of the eyes (19-22 in A. gorongosa, 15-19 in A. loveridgei; 15—20 in A. trans- vaalica) (comparative data fide Branch et al. 2017a). Afroedura donveae sp. nov. differs from other members of the A. bogerti-group by a combination of the following characters (see Tables 3, 4): 64-78 (mean 72.8) mid-body scale rows (69-77 [mean 73.5] in A. bogerti, 76-88 [mean 79.3] in A. wulfhaackei sp. nov., 73—86 [mean 80.3] in A. vazpintorum sp. nov., 73-78 [mean 74.8] in A. praedicta Zoosyst. Evol. 97 (1) 2021, 55-82 73 Figure 9. Holotype of Afroedura donveae sp. nov. (PEM R17937) from Omauha Lodge, 15 km south of Tambor, Namibe Province, Angola. Photos: Werner Conradie. sp. nov.); by the anterior nasals always in contact (similar to A. vazpintorum and A. praedicta sp. nov.; in contact in ~ 33% of A. bogerti; in contact in ~ 68% of A. wulfhaackei sp. nov.); in each verticil having 5—6 (mean 5.5) ventral and 6—7 (mean 6.6) dorsal rows of scales (5—6 [mean 5.0] and 6—7 [mean 6.1] to A. vazpintorum sp. nov.; 4 and 5 in A. bogerti and A. praedicta sp. nov., 4-5 [mean 4.0] and 5-6 [mean 5.1] in A. wulfhaackei sp. nov.); ventral surfaces immaculate (similar to A. vazpintorum sp. nov.; greyish with black spots in A. bogerti, A. wulfhaackei sp. nov. and A. praedicta sp. nov.); larger average adult size 57.6 mm SVL (versus 50.0 mm in A. bogerti, 51.7 mm in A. wulfhaackei sp. nov., 51.3 mm in A. vazpinto- rum sp. nov.; 49.9 mm A. praedicta sp. nov.). Afroedura donveae sp. nov. differs more specifically from its sister lowland species A. vazpintorum sp. nov. in being larger (57.6 mm versus 51.3 mm average SVL) and having lower mid-body scale counts (64—78 [mean 72.8] versus 73—86 [mean 80.3]), higher numbers of precloacal pores (11—12 [mean 11.5] versus 9-11 [mean 10.2]), bolder colouration and distinct tail banding (versus duller colouration and less distinct tail banding). Holotype description. Adult female: SVL 61.0 mm; tail 59.0 mm (full original tail), with a small mid-ventral incision for the removal of liver sample. Measurements and meristic characters of holotype presented in Table 7. Head and body dorsoventrally compressed; HL 13.6 mm, HW 11.4 mm, broadest at posterior level of eye and 1.19 times longer than wide. Eyes large (3.4 mm wide), pu- pil vertical with indented margins; circumorbital scales small and smooth, elongated at upper anterior margin, upper three posterior scales with small upward pointing spines. Snout rounded, 5.6 mm long, longer than distance between eye and ear openings (4.5 mm). Scales on top of snout smooth, rounded, equal in size, with no inter- vening minute granules. Scales on snout slightly larger than those on back of head or nape. Scales on eyelids larger than those on the crown, 5 scales deep from circu- morbital scale to crown. Nostril pierced between rostral, three nasal scales; 1“ supralabial narrowly excluded from nostril; the supranasals are much larger than the subequal smaller postnasals, in broad contact with each other. Nos- trils slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to extensions to the nostril. Nine supralabials on each side, the labial margin flexing upwards at the rictus (approx. mid-orbital posi- tion), with 1-2 minute scales proximal to the flexure. Ten infralabials on either side, with a small scale proximal to the flexure. At the lip, mental slightly narrower than adyja- cent infralabial, 63% of rostral and in contact with three distinctly elongated postmental scales. Scales on throat much smaller than those on belly, scales touching infral- abials larger. Thirteen scales across the crown at level of zse.pensoft.net 74 Table 7. Measurements (in mm) and scale counts for the type series of Afroedura donveae sp. nov. Catalogue number PEM R17937 PEM R17936 Type Status Holotype Paratype Sex Female Female Snout vent length 61.0 64.0 Tail length 59.0 38.0 Tail condition Original Regenerated Head length 16.6 se Head width 11.4 ti Sey Snout length a6 brs Eye distance 3.4 si Eye-Ear distance 4.4 4.4 Dorsal rows per tail verticil 6 - Ventral rows per tail verticil 7 - Scales below 4‘ toe 8 7 Mid-body scale rows 74 68 Scales between eyes 3 12 Scales: nostril to eye 11 10 Scales: ear to eye ake: 14 Anterior nasals in contact Yes Yes Supralabials 8 8 Infralabials 10 10 front of eyes; 11 scales between nostril and front of eye; 13 scales from ear to eye; 74 scales around mid-body. Ear opening deep, oblique and roughly oval, only half as high as wide (0.4