Extending along the Namibian coastline into southwestern Angola, the Namib Desert system includes one of the oldest and driest deserts in the world and is recognised as a hotspot for the evolution of geckos (Bauer, 1999; Herrmann and Branch, 2013). A long period of evolution on a highly heterogenous landscape promoted isolation, niche specialisation and explosive radiation in this region, leading to high species diversity and local endemicity rates (Bauer, 1999). Some gecko species have become highly specialised (Heinicke et al., 2017), with unusual morphological features and behavioural adaptations, like the loss of subdigital pads (Lamb and Bauer, 2006), vocalisation (Hibbitts et al., 2005) or even biofluorescence (Prötzel et al., 2021).

One of the most unique and iconic African geckos is the Namib endemic Kolekanos plumicaudus (Haacke, 2008), characterised by very slender limbs, dorsoventrally flattened body, and a bizarre feather-like tail (Haacke, 2008). It should be noted that the Reptile Database (Uetz et al., 2021) states that the correct species name should be plumicauda according to ICZN Article 34.2.1, which states that a noun in apposition does not need to match the gender of the genus. Haacke (2008) did not make any specific statement regarding the composition of the name but from the context it

appears that he intended it as an adjective “feather-tailed” instead of a noun “feather-tail”, in which case no change to the species name is needed (Bauer, pers. comm. August 2021). Based on its single pairs of enlarged digital pads, this species was thought to be related to other leaf-toed geckos and initially described as Afrogecko plumicaudus (Haacke, 2008). Subsequent molecular data revealed that southern African leaf-toed geckos (which all have single paired scansors, excluding the distantly related Goggia) represented a paraphyletic group with deeply separated lineages which were split into four genera (Afrogecko, Cryptactites, Kolekanos and Ramigekko). The latter three form monotypic lineages, while Afrogecko remains represented by two species, A. porphyreus and A. ansorgii (Heinicke et al., 2014). However, the recent rediscovery of the Angolan endemic A. ansorgii (Vaz Pinto et al., 2019) allowed for additional phylogenetic analyses which suggest that Afrogecko needs to be further split into two separate monotypic genera (Conradie et al., 2019; Lobón-Rovira et al., in prep.). This will consequently lead to the recognition of five monotypic genera within southern African leaf-toed geckos. While the knowledge of the evolutionary relatedness of this group is improving, little is known about the natural history, habitat, and distribution for most of these unique species (e.g., Ceríaco et al., 2020).

Kolekanos plumicaudus has been associated with the desert environments of the Angolan Kaokoveld. It is mainly rupicolous, sheltering under thin granite flakes in rock outcrops during the day while foraging in nearby bushes at night (Agarwal et al., 2017). It has only been recorded from five main sites in southwestern Angola, inside or in close proximity to Iona National Park (NP) and only separated by approximately 80 km at the most extreme locations (Marques et al., 2018). Haacke (2008) collected material just outside Iona NP at Tambor (type locality), a second record about 20 km south at the Curoca River crossing of Salondjamba and a third locality inside the park near the road to Oncócua, north of Iona village. The same author explored accessible

Herpetology Notes, volume 14: 1207-1212 (2021) (published online on 13 September 2021)

Updated distribution of Kolekanos plumicaudus (Sauria: Gekkonidae), with some comments on its natural history

Pedro Vaz Pinto1,2,*, Werner Conradie3,4, François S. Becker5, and Javier Lobón-Rovira1

1 Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Rua Padre Armando Quintas, Campus de Vairão, Vairão 4485-661, Portugal.

2 Fundação Kissama, Rua 60 Casa 560, Lar do Patriota, Luanda, Angola.

3 Port Elizabeth Museum (Bayworld), P.O. Box 13147, Humewood 6013, South Africa.

4 Department of Nature Conservation Management, Natural Resource Science and Management Cluster, Faculty of Science, George Campus, Nelson Mandela University, George, South Africa.

5 National Museum of Namibia, Windhoek, Namibia.* Corresponding author. E-mail: pedrovazpinto@gmail.com

© 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0.

Pedro Vaz Pinto et al.1208

areas in northern Namibia along the Angolan border on multiple occasions in search of this species, without any success (Haacke, 2008). During the 2009 SANBI/ISCED/UAN Angolan Biodiversity Assessment and Capacity Building Project expedition to southwestern Angola, new material was collected north of Iona village, near the third original collecting site (Huntley, 2009). Recently, a fifth site at Omauha farm was added to the species’ distribution (Agarwal et al., 2017). Various herpetological surveys conducted in Iona NP and across the Angolan province of Namibe failed to record the species elsewhere (Ceríaco et al., 2016; Marques et al., 2018; Branch et al., 2019), strongly suggesting a highly restricted distribution range. In all these cases the species was found associated with granite outcrops overlaying a sandy substrate at mid-elevations below 800 m elevation. A recent IUCN assessment awarded the species a Least Concern status, suggesting it to be common in suitable habitat yet seldom encountered, with an estimated extent of occurrence (EOO) of 4,000 km2 and present between 200 and 600 m elevation, while recommending further studies on its biology (Ceríaco et al., 2020).

Recognising that the poorly understood natural history of K. plumicaudus may undermine future conservation efforts for the species, we performed a series of herpetological surveys across Namibe Province, in southwestern Angola, between September 2019 and May 2021. During these surveys we visited the known K. plumicaudus localities, nearby sites within the known range, and expanded our search into new regions, also covering different habitats and a wider elevation gradient. The surveys consisted of active searches by looking inside cracks and under rock flakes during the day and inspecting rocks and vegetation near rocky outcrops at night. Specimens found far from the known distribution were collected as vouchers. Specimens collected as vouchers were humanely euthanised with oral application of benzocaine gel or injection of tricaine methanesulfonate (MS222) (Conroy et al., 2009). After euthanasia, specimens were fixed in 96% ethanol, after which they were transferred to 70% ethanol for long-term storage in the natural history collection of Fundação Kissama (FKH), Luanda, Angola, and the National Museum of Namibia (NMNW), Windhoek, Namibia. For future molecular analyses, liver or muscle

Figure 1. Distribution of Kolekanos plumicaudus within Angola. Red circles depict the previously known records of the species, and the red shading the corresponding IUCN polygon (Ceríaco et al., 2020). Green circles depict new records reported in this work, and the green shading the updated EOO obtained using GeoCAT. The red star depicts the type locality of the species.

samples were collected and stored in 96% ethanol. For each observation or sample collected the location was recorded with precise coordinates, using the WGS84 coordinate system. Due to the morphological uniqueness of the species (slender body and feather-like tail), photographic records were considered as valid records for individuals not vouchered.

Resulting from these extensive surveys, we have confirmed the occurrence of K. plumicaudus at 12 sites (Table 1), including two previously known localities, Omauha farm, and at Conguingulo, which is along the road from Iona to Ocócua and very close to the third site reported by Haacke (2008) (Fig. 1). The remaining sites expanded the species’ distribution ~70 km southwest to the margins of the Cunene River and ~30 km southeast, in both cases to near the Namibian border, and to the north of the town of Moçamedes, a ~120 km northern range extension. Moreover, the species was recorded from across a large elevational range, spanning from less than 100 m to above 2,000 m elevation (Table 1).

Specimens were found in different habitats and geological settings, not only near isolated large granite

outcrops as previously reported. These included small or scattered igneous rock outcrops in steep valleys bordered by large sand dunes (e.g., Muende-Curoca, Fig. 2H), in rocky mountain summits (e.g., Serra Cafema, Fig. 2D), and in quartzite sandstone cliffs in mountain ridges (e.g., Tchamalinde, Fig. 2E). Consistent with previous findings (Haacke, 2008; Agarwal et al., 2017), K. plumicaudus were found sheltering under very narrow rock flakes during the day or foraging among nearby small bushes at night. As previously reported, they were often found stretched flat on thin branches (Agarwal et al., 2017), and proved to be extremely agile when disturbed, running rapidly along and between twigs and jumping readily. Such jumps were often impressive, allowing them to disappear quickly, either by obtaining cover in rock cracks or by reaching relatively distant branches and bushes. On one occasion we observed a female jumping more than 40 cm between branches.

This species was frequently found in sympatry with several other rupicolous lizard species, such as Rhoptropus spp., Pachydactylus cf. oreophilus, Chondrodactylus pulitzerae, and Trachylepis sulcata. At

Updated distribution of Kolekanos plumicaudus 1209

Table 1. Detailed collection and observational records of Kolekanos plumicaudus in southwestern Angola. Catalogue numbers refer to vouchers housed in collections in the California Academy of Science (CAS), Florida Museum of Natural History (UF), Kissama Foundation (FKH, and field numbers P1 and JLRZ), National Museum of Namibia, Windhoek (NMNW), Ditsong National Museum of Natural History (formerly the Transvaal Museum; TM) and Port Elizabeth Museum (PEM). Where material was not collected, vouchers are stated as Not Available (NA).

Catalogue Locality Date Latitude Longitude Elevation (m) Source

TM 40521–31 Tambor 01/04/1971 -16.1356 12.4297 388 Haacke (2008)

TM 40553–55 Curoca River Crossing 02/04/1971 -16.3027 12.4165 192 Haacke (2008)

TM 40755–61 11 km NE from Iona 08/04/1971 -16.8606 12.6107 815 Haacke (2008)

PEM R18047; PEM R18010–5; CAS 248782

7 km NE from Iona 12/12/2009 -16.8583 12.6128 808 Port Elizabeth Museum

UF 187219–22; CAS 262389–91

Omauha 25/11/2016 -16.1996 12.3988 341 Agarwal et al. (2017)

FKH 0235–6 Camp Baptista Cunene 30/09/2019 -17.1604 12.0182 68 This work

FKH 0343–6; JLRZC0109

Omauha 03/10/2019; 03/12/2020; 10/03/2021

-16.1988 12.4013 338 This work

NA Mutuovano 08/03/2021 -15.9153 12.3848 319 This work

NA Muende-Curoca 10/03/2021 -16.2892 12.3180 248 This work

NA Tchitchaki 13/03/2021 -16.2878 12.2754 223 This work

NA Humbi 17/03/2021 -16.9859 12.5415 875 This work

NA Congundo 17/03/2021 -17.0396 12.6013 916 This work

NA Conguiungulo 18/03/2021 -16.8437 12.6141 788 This work

P1 021–22 Maongo-Giraul 01/04/2021 -15.0327 12.4147 365 This work

NA Chamaleva 22/04/2021 -15.6863 12.6124 427 This work

NMNW R11011–2 Tchamalindi 27/04/2021 -16.9753 12.8834 1447 This work

P1 127 Serra Cafema 28/04/2021 -17.1290 12.5139 2035 This work

Omauha, the species was frequently observed sharing a similar niche with another flat-bodied gecko species, Afroedura donveae, sheltering under very thin granite flakes. The latter species was mostly found foraging among thicker branches of large bushes and small trees sprouting directly from the steep faces of rock boulders, while K. plumicaudus preferred thinner twigs of smaller bushes, and were often found near the ground at the base of rock outcrops. At Tchamalindi and Serra Cafema, we retrieved specimens of Hemirhagerrhis viperina in similar circumstances and in near proximity to where

K. plumicaudus were collected, strongly suggesting that this snake species may be predating on these geckos.

As an added observation, we report blue bone-based fluorescence of the skull, vertebrae and hindlimb joints, through the translucent skin, of K. plumicaudus after exposing one individual to ultraviolet light (~395 nm) (Fig. 3). This observation adds to the growing knowledge of gekkonid species with bone-based fluorescence, including Chondrodactylus (Sloggett, 2018), Cyrtodactylus (Jeng, 2019; Top et al., 2020) and Pachydactylus (Prötzel et al., 2021). No dermal

Figure 2. Habitat photographs of Kolekanos plumicaudus: (A) Omauha, (B) Salondjamba, (C) Maongo-Giraul, (D) Serra Cafema, (E) Tchamalindi, (F) Congundo, (G) Camp Baptista-Cunene, (H) Muende-Curoca, (I) Tchichaki, and (J) Chamaleva. Photographs by Pedro Vaz Pinto (D, E, G, J), Michel Morais (C) and Javier Lobón-Rovira (A, B, F, H, I).

Pedro Vaz Pinto et al.1210

fluorescence was observed, similar to what has been reported for Pachydactylus rangei (Prötzel et al., 2021).

This work provides an updated distribution for K. plumicaudus within Namibe Province, the known range now spanning ~240 km from north of the city of Moçamedes southwards to the margins of the Cunene River on the Namibian border. Furthermore, using the GeoCAT IUCN tool (following Bachman et al., 2011) we update the EOO to 11,333 km2, which almost triples the previous estimates (Ceríaco et al., 2020; Fig. 1). These results support the hypothesis of K. plumicaudus being a species characteristic of the Angolan Kaokoveld within the Namib Desert system, but also suggest that they may be marginally present in north-western Namibia near the Cunene River at the edge of the sand sea, a possibility that should be investigated further in future works. Our new findings highlight some key habitat requirements of this species: sparse bushes with

thin branches for nocturnal foraging, in close proximity to rock formations with deep, narrow crevices (often formed by flaked weathering patterns) for diurnal shelter. While it was previously assumed to occur only on large granite boulders, smaller outcrops of quartzite or other formations with similar weathering patterns are also suitable for this species. We further demonstrate a wide elevational tolerance, ranging from below 100 m to above 2,000 m elevation. Finally, we provide the first evidence of fluorescence in a leaf-toed gecko, adding to at least three other distantly related species of geckos in the Namib Desert who are known to fluoresce, suggesting that this feature may have evolved independently in different lineages in this environment.

Acknowledgements. This work greatly benefitted from an existing institutional collaboration with the Ministry of Culture, Tourism and Environment of the Republic of Angola (MCTA), and in particular the support received by Dr Albertina Nzuzi, Director of Instituto Nacional de Biodiversidade e Conservação (INBC).

Figure 3. Kolekanos plumicaudus in dorsal view under (A) natural light, and (B) 395 nm ultraviolet (UV) light, showing the bone-based blue fluorescence of the skull, vertebrae and hindlimb joints. Photos by Javier Lobón-Rovira.

Updated distribution of Kolekanos plumicaudus 1211

Material was collected under permit ISCED/001/21. A very special thanks is due to the owners of Omauha farm and lodge, Álvaro and Varito Baptista, for their generous logistical support, friendship and sharing of knowledge. We also thank African Parks, and in particular the warden of Iona NP, Bruce Bennet, and its operations manager, Bastiaan Boon, who accompanied and greatly contributed to the success of various surveys. We acknowledge the important role played by Rolf Becker, Ansie Bosnam and Vera de Cauwer in organising the mountain surveys, and Fernanda Lage and Vladimir Russo for coordinating institutional relationships and facilitating work authorisations. Surveys for the Project for Creation of Marine Protected Areas in Angola was financially supported by GEF6, and the SCIONA - Biodiversity Survey of Mountain Tops in the Kaokoveld Centre of Endemism, done in partnership with MCTA and the Ministry of Environment, Forestry and Tourism of Namibia, was funded by the European Union under grant agreement FED/2017/394-802. JLR is currently supported by Fundação para a Ciência e Tecnologia (FCT) contract PD/BD/140808/2018. Finally, we thank Franco Andreone for the pre-review of the manuscript and Aaron Bauer for advice on the correct species name composition.

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Accepted by Darren Pietersen

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