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Language editing: Will Simonson (Cambridge), and Proofreading Pal Translation of abstracts to Portuguese: Ana Filipa Guerra Silva Gomes da Piedade Page desing & layout: Marit Arnold, Klaus A. Hess, Ria Henning-Lohmann Cover photographs: front: Thunderstorm approaching a village on the Angolan Central Plateau (Rasmus Revermann) back: Fire in the miombo woodlands, Zambia (David Parduhn) Cover Design: Ria Henning-Lohmann

ISSN 1613-9801

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Suggestion for citations: Volume: Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N. (eds.) (2018) Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions. Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek.

Articles (example): Archer, E., Engelbrecht, F., Hänsler, A., Landman, W., Tadross, M. & Helmschrot, J. (2018) Seasonal prediction and regional climate projections for southern Africa. In: Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions (ed. by Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N.), pp. 14–21, Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek.

Corrections brought to our attention will be published at the following location: http://www.biodiversity-plants.de/biodivers_ecol/biodivers_ecol.php Biodiversity & Ecology

Journal of the Division Biodiversity, Evolution and Ecology of Plants, Institute for Plant Science and Microbiology, University of Hamburg

Volume 6:

Edited by

Rasmus Revermann1, Kristin M. Krewenka1, Ute Schmiedel1, Jane M. Olwoch2, Jörg Helmschrot2,3, Norbert Jürgens1

1 Institute for Plant Science and Microbiology, University of Hamburg 2 Southern African Science Service Centre for Climate Change and Adaptive Land Management 3 Department of Soil Science, Faculty of AgriSciences, Stellenbosch University

Hamburg 2018

RPlease cite the article as follows:

Baptista, N., António, T. & Branch, W.R. (2018) Amphibians and reptiles of the Tundavala region of the Angolan Escarpment. In: Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions (ed. by Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N.), pp. 397-403, Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek. doi:10.7809/b-e.00351 mhibians and retiles of the undavala region of the ngolan Escarment

inda Batista, elmo ntnio, illiam Branch, Instituto Suerior de incias da Educao da Hula, ua Sarmento odrigues, ubango, ngola Port Eliabeth Museum Bayorld, P Bo , Humeood , South frica esearch ssociate, Deartment of oology, P Bo , elson Mandela Metroolitan University, Port Eliabeth , South frica orresonding author nindabatistagmailcom Biodiversity The most poorly known section of the African Great Escarpment is located in Angola. It has been highlighted as a potential center of endemism for several biological groups, including herpetofauna. The region, which is critical for the conservation of Angolan biodiversity, reuires urgent research. In the scope of the SASSCAL project, a herpetofauna moni- toring plan is being implemented in Tundavala, in the southern Angolan escarpment. In total, 13 species of amphibians, 12 species of liards and 9 species of snakes have been registered so far, and more are expected to occur. Among them are some important rediscoveries of Angolan endemics uncollected for decades, such as Anchietas treefrog (Leptopelis anchietae) and Ansorges whip snake (Psammophis ansorgii).

A seco menos conhecida da Grande Escarpa Africana localia-se em Angola. Esta j foi destacada como um potencial centro de endemismo para vrios grupos, incluindo a herpetofauna, sendo por isso uma regio crtica para a con- servao da biodiversidade angolana ue carece urgentemente de investigao. No mbito do projecto SASSCAL, est a ser implementado um plano de monitoriao de herpetofauna na Tundavala, no sul da escarpa de Angola. At ao momento foram registadas 13 espcies de anfbios, 12 espcies de lagartos e nove espcies de cobras, e presume-se ue existam mais. Entre estas, esto redescobertas importantes de endemismos angolanos no registados h vrias dcadas, como a r-arbor- cola-de-Anchieta (Leptopelis anchietae), e a cobra-de-Ansorge (Psammophis ansorgii).

trodctio it one of the most important tourist des- the region with their livestock (cows and tinations in the country. The creation of goats) and plant crops. Other threats in- The Angolan section is the most poorly a nature reserve in Tundavala was pro- clude man-made res and the dumping known of the African Great Escarpment, posed by Huntley & Matos (1994), and of rubble and domestic, commercial, and and yet it supports the highest number of the reserve is mentioned in the Angolan even medical waste. vertebrate endemics after those in South National Biodiversity Strategy and Ac- Southwestern Angola is one of the Africa (Clark et al., 2011). It is rich in tion Plan (NBSAP, 2006) as a protected better-surveyed regions of Angola for all endemic plants and birds and is a po- area to be implemented. It is also classi- vertebrate groups (Crawford-Cabral & tential hotspot for Angolan biodiversity ed as an Important Bird Area (BirdLife Mesuitela, 1989), and the herpetofauna (Hall, 1960). However, more research International, 2017), and the Tundavala is no exception, with numerous studies is reuired to identify and meaningfully Crevice was classi ed as a Cultural addressing it (Bocage, 1895 Schmidt, implement conservation objectives, es- Landscape in 2012 (Executive decree no. 1933, 1936 Parker, 1936 Monard, pecially detailed biodiversity surveys 261/12). Despite its social and biological 1937a,b Mertens, 1938 Bogert, 1940 and systematic studies. These will allow importance, the region lacks o cial na- Loveridge, 1944 Hellmich, 1957 FitSi- the e ects of predicted climate change, tional protected status and is threatened mons, 1959 Laurent, 1964 Poynton & highlighted by Clark et al. (2011), to be by progressively increasing human activ- Haacke, 1993 Haacke, 1997). Despite identi ed, and refugia and migration hy- ities, especially logging and burning for this, new species continue to be described potheses to be formulated and tested. charcoal production and the harvesting from the region (Haacke, 2008 Conradie Tundavala is located on the Angolan of natural resources such as medicinal et al., 2012 Stanley et al., 2016). Little escarpment in southwestern Angola (Fig. plants and rocks for building purposes. work has focused speci cally on the es- 1), and its outstanding landscape makes Increasing numbers of villagers inhabit carpment region, and its herpetofauna is

B E 6 2018 397 Biodiversity

igure eft ocation of the study site in the frican reat Escarment ight ocalities near the study area mentioned here additional secies eected in undavala ere recorded

far from being completely understood. The Humpata Plateau, including Tun- davala, has been identi ed as a potential center of endemism for montane herpeto- fauna and is therefore a priority area for herpetofaunal surveys (Laurent, 1964). The main goal of the SASSCAL Projects observatories (task 210) is to monitor biodiversity to understand the e ects of climate change and land use on plant communities in the long term. Several observatories have been im- plemented throughout Angola, includ- ing one in Tundavala. The herpetofauna igure Montane grassland grassy habitat at rst monitoring site is freuently cited as a useful indicator for environmental monitoring thanks to its importance in ecological function- ing and its sensitivity to environmental change (Smith & Rissler, 2010). In this context, a monitoring program of the Tundavala herpetofauna was initiated in 2016. The compilation of baseline infor- mation, such as presence and absence of species and their relative abundances in the mid- and long term, is essential for the e ective management of this critical region for the conservation of Angolan biodiversity. This is the rst herpetofauna monitoring plan implemented in Angola. The project is ongoing, and preliminary and more relevant ndings are presented in this publication. igure eathered sandstone outcros rocy habitat and associated oody and grassy vegetation at second monitoring site

398 C A ŽŵŵŽŶŶĂŵĞ ^ƉĞĐŝĞƐ &ĂŵŝůLJ able mhibian secies recorded from the undavala ŶĐŚŝĞƚĂ͛ƐdƌĞĞĨƌŽŐ ;Ϳ>ĞƉƚŽƉĞůŝƐĂŶĐŚŝĞƚĂĞ;ŽĐĂŐĞ͕ϭϴϳϯͿ ƌƚŚƌŽůĞƉƚŝĚĂĞ observatory and close ŽĐĂŐĞ͛ƐdƌĞĞĨƌŽŐ >ĞƉƚŽƉĞůŝƐĐĨ͘ďŽĐĂŐŝŝ;'ƺŶƚŚĞƌ͕ϭϴϲϱͿ ƌƚŚƌŽůĞƉƚŝĚĂĞ surroundings E indicates 'ƵƚƚƵƌĂůdŽĂĚ ^ĐůĞƌŽƉŚLJƌƐĐĨ͘ŐƵƚƚƵƌĂůŝƐ;WŽǁĞƌ͕ϭϵϮϳͿ ƵĨŽŶŝĚĂĞ secies endemic to the &ůĂƚͲĂĐŬĞĚdŽĂĚ ^ĐůĞƌŽƉŚƌLJƐĐĨ͘ƉƵƐŝůůĂ;DĞƌƚĞŶƐ͕ϭϵϯϳͿ ƵĨŽŶŝĚĂĞ ngolan highlands DŽŶĂƌĚ͛ƐZĞĞĚ&ƌŽŐ ;Ϳ,LJƉĞƌŽůŝƵƐĐŝŶĞƌĞƵƐDŽŶĂƌĚ͕ϭϵϯϳ ,LJƉĞƌŽůŝŝĚĂĞ ŶŐŽůĂZĞĞĚ&ƌŽŐ ,LJƉĞƌŽůŝƵƐĐĨ͘ƉĂƌĂůůĞůƵƐ'ƺŶƚŚĞƌ͕ϭϴϱϴ ,LJƉĞƌŽůŝŝĚĂĞ ĞŶŐƵĞůĂ>ŽŶŐZĞĞĚ&ƌŽŐ ,LJƉĞƌŽůŝƵƐĐĨ͘ďĞŶŐƵĞůůĞŶƐŝƐ;ŽĐĂŐĞ͕ϭϴϵϯͿ ,LJƉĞƌŽůŝŝĚĂĞ ƵďďůŝŶŐ<ĂƐƐŝŶĂ <ĂƐƐŝŶĂƐĞŶĞŐĂůĞŶƐŝƐ;ƵŵĠƌŝůĂŶĚŝďƌŽŶ͕ϭϴϰϭͿ ,LJƉĞƌŽůŝŝĚĂĞ DĂďĂďĞWƵĚĚůĞ&ƌŽŐ WŚƌLJŶŽďĂƚƌĂĐŚƵƐĐĨ͘ŵĂďĂďŝĞŶƐŝƐ&ŝƚnj^ŝŵŽŶƐ͕ϭϵϯϮ WŚƌLJŶŽďĂƚƌĂĐŚŝĚĂĞ WĞƚĞƌƐ͛ůĂǁĞĚ&ƌŽŐ yĞŶŽƉƵƐĐĨ͘ƉĞƚĞƌƐŝŝŽĐĂŐĞ͕ϭϴϵϱ WŝƉŝĚĂĞ ŶĐŚŝĞƚĂ͛ƐZŝĚŐĞĚ&ƌŽŐ WƚLJĐŚĂĚĞŶĂĐĨ͘ĂŶĐŚŝĞƚĂĞ;ŽĐĂŐĞ͕ϭϴϲϴͿ WƚLJĐŚĂĚĞŶŝĚĂĞ ŶŐŽůĂZŝǀĞƌ&ƌŽŐ ŵŝĞƚŝĂĂŶŐŽůĞŶƐŝƐ;ŽĐĂŐĞ͕ϭϴϲϲͿ WLJdžŝĐĞƉŚĂůŝĚĂĞ ZŽƵŐŚ^ĂŶĚ&ƌŽŐ dŽŵŽƉƚĞƌŶĂĐĨ͘ƚƵďĞƌĐƵůŽƐĂ;ŽƵůĞŶŐĞƌ͕ϭϴϴϮͿ WLJdžŝĐĞƉŚĂůŝĚĂĞ Biodiversity

ŽŵŵŽŶŶĂŵĞ ^ƉĞĐŝĞƐ &ĂŵŝůLJ able etile secies >ŝnjĂƌĚƐ;^ĂƵƌŝĂͿ recorded from the undavala observatory and close sur 'ƌŽƵŶĚŐĂŵĂ ŐĂŵĂĐĨ͘ĂĐƵůĞĂƚĂ;DĞƌƌĞŵ͕ϭϴϮϬͿ ŐĂŵŝĚĂĞ roundings E indicates se EĂŵŝďZŽĐŬŐĂŵĂ ŐĂŵĂĐĨ͘ƉůĂŶŝĐĞƉƐWĞƚĞƌƐ͕ϭϴϲϮ ŐĂŵŝĚĂĞ cies endemic to the ngolan ŽƵďůĞͲƐĐĂůĞĚŚĂŵĞůĞŽŶ ŚĂŵĂĞůĞŽĂŶĐŚŝĞƚĂĞŽĐĂŐĞ͕ϭϴϳϮ ŚĂŵĂĞůĞŽŶŝĚĂĞ highlands indicates DĂĐŚĂĚŽ͛Ɛ'ŝƌĚůĞĚ>ŝnjĂƌĚ ;ͿŽƌĚLJůƵƐŵĂĐŚĂĚŽŝ>ĂƵƌĞŶƚ͕ϭϵϲϰ ŽƌĚLJůŝĚĂĞ cases here taonomy fol ^ƉĞĐŬůĞĚdŚŝĐŬͲƚŽĞĚ'ĞĐŬŽ WĂĐŚLJĚĂĐƚLJůƵƐĐĨ͘ƉƵŶĐƚĂƚƵƐWĞƚĞƌƐ͕ϭϴϱϰ 'ĞŬŬŽŶŝĚĂĞ los aurent DŽŶƚĂŶĞĂLJ'ĞĐŬŽ ;ͿΎZŚŽƉƚƌŽƉƵƐďŽƵůƚŽŶŝŵŽŶƚĂŶƵƐ>ĂƵƌĞŶƚ͕ϭϵϲϰ 'ĞŬŬŽŶŝĚĂĞ tĞƐƚĞƌŶ^ĞƌƉĞŶƚŝĨŽƌŵ^ŬŝŶŬ ƵŵĞĐŝĂĂŶĐŚŝĞƚĂĞŽĐĂŐĞ͕ϭϴϳϬ ^ĐŝŶĐŝĚĂĞ ŶŐŽůĂďƵƌƌŽǁŝŶŐ^ŬŝŶŬ ^ĞƉƐŝŶĂĐĨ͘ĂŶŐŽůĞŶƐŝƐŽĐĂŐĞ͕ϭϴϲϲ ^ĐŝŶĐŝĚĂĞ ,ŽĞƐĐŚ͛ƐƐŬŝŶŬ dƌĂĐŚLJůĞƉŝƐŚŽĞƐĐŚŝ;DĞƌƚĞŶƐ͕ϭϵϱϰͿ ^ĐŝŶĐŝĚĂĞ tĞƐƚĞƌŶZŽĐŬ^ŬŝŶŬ ;ͿΎdƌĂĐŚLJůĞƉŝƐƐƵůĐĂƚĂĂŶƐŽƌŐĞŝ;ŽƵůĞŶŐĞƌ͕ϭϵϬϳͿ ^ĐŝŶĐŝĚĂĞ sĂƌŝĂďůĞ^ŬŝŶŬ dƌĂĐŚLJůĞƉŝƐĐĨ͘ǀĂƌŝĂ;WĞƚĞƌƐ͕ϭϴϲϳͿ ^ĐŝŶĐŝĚĂĞ tĂŚůďĞƌŐ͛Ɛ^ŬŝŶŬ dƌĂĐŚLJůĞƉŝƐǁĂŚůďĞƌŐŝ;WĞƚĞƌƐ͕ϭϴϲϵͿ ^ĐŝŶĐŝĚĂĞ ^ŶĂŬĞƐ;^ĞƌƉĞŶƚĞƐͿ ŶŐŽůĂŶ'ƌĞĞŶ^ŶĂŬĞ WŚŝůŽƚŚĂŵŶƵƐĂŶŐŽůĞŶƐŝƐ;ŽĐĂŐĞ͕ϭϴϴϮͿ ŽůƵďƌŝĚĂĞ ŽŶĨƵƐŝŶŐŐŐͲĂƚĞƌ ĂƐLJƉĞůŝƐĐĨ͘ĐŽŶĨƵƐĂ;dƌĂƉĞΘDĂŶĠ͕ϮϬϬϲͿ ŽůƵďƌŝĚĂĞ ŶĐŚŝĞƚĂ͛ƐĐŽďƌĂ EĂũĂĂŶĐŚŝĞƚĂĞŽĐĂŐĞ͕ϭϴϳϵ ůĂƉŝĚĂĞ sŝƉĞƌŝŶĞZŽĐŬ^ŶĂŬĞ ,ĞŵŝƌŚĂŐĞƌƌŚŝƐǀŝƉĞƌŝŶĂ;ŽĐĂŐĞ͕ϭϴϳϯͿ >ĂŵƉƌŽƉŚŝŝĚĂĞ ^ƉŽƚƚĞĚtŽůĨ^ŶĂŬĞ >LJĐŽƉŚŝĚŝŽŶĐĨ͘ŵƵůƚŝŵĂĐƵůĂƚƵŵŽĞƚƚŐĞƌ͕ϭϴϴϴ >ĂŵƉƌŽƉŚŝŝĚĂĞ ŶƐŽƌŐĞ͛ƐtŚŝƉ^ŶĂŬĞ ;ͿWƐĂŵŵŽƉŚŝƐĂŶƐŽƌŐŝŝ;ŽƵůĞŶŐĞƌ͕ϭϵϬϱͿ >ĂŵƉƌŽƉŚŝŝĚĂĞ DŽnjĂŵďŝƋƵĞ'ƌĂƐƐ^ŶĂŬĞ WƐĂŵŵŽƉŚŝƐĐĨ͘ŵŽƐƐĂŵďŝĐƵƐ;WĞƚĞƌƐ͕ϭϴϴϮͿ >ĂŵƉƌŽƉŚŝŝĚĂĞ KĐĞůůĂƚĞĚ^ŬĂĂƉƐƚĞŬĞƌ ;ͿWƐĂŵŵŽƉŚLJůĂdžƌŚŽŵďĞĂƚƵƐŽĐĞůůĂƚƵƐ;ŽĐĂŐĞ͕ϭϴϳϯͿ >ĂŵƉƌŽƉŚŝŝĚĂĞ ůŝŶĚ^ŶĂŬĞ ĨƌŽƚLJƉŚůŽƉƐĐĨ͘ĂŶŽŵĂůƵƐ;ŽĐĂŐĞ͕ϭϴϳϯͿ dLJƉŚůŽƉŝĚĂĞ  etods oitorig April). Monitoring involves the installa- Herpetofauna monitoring in the tion of two 15-meter-long traplines, each rvey re SASSCAL Tundavala observatory region associated with six funnel and two pit fall The Tundavala region of the plateau is involved standardied sampling in the two traps. These are placed for 14 consecutive composed of a mosaic of vegetation types habitat types occurring in the observatory days and are checked daily. In addition, and rugged topography. Patches of rel- area: sandy soils with montane grassland 20 time-constrained transects with visual ict Afromontane forest with Podocarpus (grassy habitat, Figure 2), and corridors encounter surveys (VES) are done on m i l a n j i a n u s occur in deep humid ravines of undi erentiated woody montane com- sunny mornings (10) and afternoons (10). and at altitudes above 1,800 m (Huntley munities between large weathered sand- Both techniues are undertaken eually & Matos, 1994). They co-occur on the es- stone outcrops (rocky habitat, Figure 3). in grassy and rocky habitats. Additional carpment plateau with patches of open Pro- Preliminary herpetological surveys began VESs are undertaken opportunistically tea savanna, Pteridium bracken, miombo in November 2015. Standardied moni- in unmonitored habitats during day and woodlands on sands, montane grasslands, toring was initiated in April 2016, with night, as are auditory surveys for adult thickets along seasonal streams, and poorly three monitoring sessions each year: fol- frogs and tadpole sampling. Small mam- drained grassy patches in valleys (BirdLife lowing early rains (October–November), mals are also included in the Tundavala International, 2017), and provide diverse at the peak of the rainy season (February), monitoring plan, but results are not ad- habitats for the herpetofauna. and at the end of the rainy season (late dressed in the present work.

B E 6 2018 399 Reptiles and amphibians were collect- in Tundavala given their habitat associ- ecies list tooy d co ed and deposited in the herpetological ations, but have not been recorded so far. servtio relevce collection currently deposited at the In- Among these, some have been found in Many of the species listed in Tables 1 and stituto Superior de Cincias da Educao Humpata: snout burrower, Hemisus sp. 2 have been only provisionally identi ed. da Hula (ISCED-Hula), and tissue for (Branch et al., unpub. Data) long-head- This re ects the current poor state of genetic analysis has been preserved in ed tropical house gecko, Hemidactylus knowledge of the Angolan herpetofauna. 95 ethanol for use in ongoing and fur- longicephalus Bocage, 1873 Angolan Many of the provisional identi cations ther research. Preliminary identi cation rough-scaled liard, Ichnotropis bivitta- involve wide-ranging species or species of species is based on Channing (2001, ta pallida Laurent, 1964 Bayons skink, complexes in which cryptic diversity has 2012) and Branch (1998), supplemented Trachylepis bayonii huilensis (Laurent, recently been identi ed but for which no, with taxa-speci c literature when need- 1964) Angolan garter snake, Elap- or little, Angolan material was included. ed. Taxonomy follows Frost (2017) for soidea semiannulata Bocage, 1882) (all Other species are poorly known Ango- amphibians and Uet & Hoek (2017) for records from Laurent, 1964). Others lan species, some known from very few reptiles, except where noted. All pictures have been recorded in Lubango: Afri- specimens and for which the types were of landscape and animals are from Tun- can house snake, Boaedon cf. fuligino- lost in the re that destroyed the Lisbon Biodiversity davala. sus (Boie, 1827) leopard grass snake, Museum in 1978. These problematic Psammophis leopardinus Bocage, 1887 species, of which some may be new to three-lined grass snake, Psammophylax science, given the geographic setting of eslts tritaeniatus (Günther, 1868) (Va Pinto, Tundavala, are either currently under in- pers. obs.), and in Lubango, Leba Pass, vestigation or will reuire further inves- lig sccess and the Estao Zootcnica: ap-necked tigation when additional, often topotypic Trapping success of herpetofauna at Tun- chamaeleon, Chamaeleo dilepis (Leach, material becomes available. Taxonomic davala has been low, regardless of the 1819) (Brach et al., unpub. data Baptis- issues associated with some of these spe- sampling season. In the four sampling ta, pers. obs.) at gecko Afroedura sp., cies are addressed below. seasons performed, fewer than 15 indi- snake-eyed skink Panaspis sp. (Va Pin- viduals were caught in the traplines, and to & Baptista, unpub. data), and the en- in one sampling season in particular, only demic habitat specialist Chela reed frog, one individual was captured during the Hyperolius chelaensis (Conradie et al., 14 days of sampling. As a result of low 2012). catch rates at Tundavala, the data do not allow in-depth or temporal analysis, and thus we will focus on the observed herpe- iscssio tofaunal diversity and its biogeographic importance. oitorig sccess The herpetological monitoring program ecies list in the Tundavala observatory region has A total of 13 species of frogs (Tab. 1), 12 been of only limited success and reuires igure nchietas treefrog Leptopelis anchietae, breeding male. species of liards, and 9 species of snakes adjustment. New approaches considered (Tab. 2) were recorded during the moni- include surveys at di erent times during toring and opportunistic surveys in the the day, checking traplines twice daily, Leptopelis anchietae(Bocage, 1873) observatory and close surroundings. and modifying the trapline arrangement This species is known from Misso da Most species were caught too few to use a -shaped (3 arms of 15 m) array Hula, Caconda, and uindumbo (Bo- times to establish habitat associations. in grassy areas. Opportunistic acoustic cage, 1895) between Benguela and Some reptile species were regularly as- surveys and tadpole collection proved Bi (Boulenger, 1905) Caluuembe, sociated with rocky habitats (e.g., Agama productive and will be standardied and Ebanga, Misso do Cubango (Monard, cf. planiceps, Rhoptropus b. montanus, included in the monitoring plan on a sys- 1937a) Chitau (Schmidt, 1936) and Alto and Hemirhagerrhis viperina), whilst tematic basis. Chicapa (Laurent, 1964). It is endemic to others were found in both habitats (e.g., The Tundavala plateau has varied the Angolan plateau (Monard, 1937a). Psammophylax r. ocellatus). As expect- soils, geology, topography, and vegeta- The specimens from Tundavala (Fig. 4) ed, most amphibians except Sclerophrys tion types that provide di erent habitats correspond in morphology and coloration spp., Tomopterna tuberculosa, and Lep- and microhabitats for the herpetofauna. with the type description and plate in Bo- topelis bocagii were found only near wa- Currently, systematic monitoring covers cage (1895). Details of this rediscovery ter bodies. only two habitat types and therefore does and those of other Leptopelis from north- Additional species recorded from not fully survey the entire herpetofaunal ern areas of the Angolan escarpment will Lubango and adjacent regions of the diversity of the plateau and the escarp- be presented elsewhere (Baptista et al., in escarpment may be expected to occur ment. prep.).

400 C A Leptopelis bocagii(Günther, 1865) which has recently been revised (Chan- only subseuent material comprises six This species, originally described from ning et al., 2013). Both species are wide- heads and one body from Bella-Vista Calandula (formerly Duue de Bra- spread in Angola and have several names (Hellmich, 1957). The Tundavala speci- gana), is considered widespread in Af- in their synonymy, with variations in mens (Fig. 7) are the rst with detailed rica, including the north, west, and south shape, sie, and coloration. Comprehen- locality and habitat. They agree mor- of Angola (Bocage, 1895 Ferreira, 1904, sive studies, including wide geographical phologically with the description and the 1906 Schmidt, 1936 Monard, 1937a surveys combined with genetics and ad- plate provided in the original description Laurent, 1954, 1964 Hellmich, 1957 vertisement calls, are needed to resolve (Boulenger, 1905), and its phylogenetic Laurent, 1954, 1964). It is thought to be a the taxonomy of both groups. An Ango- a nities within Psammophis are cur- complex of cryptic species (Frost, 2017), lan reed frog from Tundavala is shown in rently being addressed (Branch et al., in and the availability of Angolan material Figure 5. prep.). on Hylambates angolensis Bocage, 1893, currently in the synonymy of L. bocagii, Tomopterna tuberculosa (Boulenger, needs further study (Perret, 1967). 1882) One of the most common species of frog Sclerophrys spp. in the monitoring region, with a variety of Biodiversity The status of the Angolan toad taxonomy color morphs. Originally described from needs to be readdressed (Ruas, 1996). S. Pungo Andongo on the Angolan central gutturalis is widespread in the country, plateau, this species is widespread in the and the recently renamed Sclerophrys country (Ruas, 1996), extends east to pusilla (Poynton et al., 2016) is known central coastal Moambiue, and may igure nsorges hi snae mostly from western Angola (Ruas, comprise a complex of cryptic species. Psammophis ansorgii 1996). However, both species have wide Other available names for Angolan mate- distributions in Africa, and their taxon- rial currently subsumed under T. tubercu- Eumecia anchietae Bocage, 1870 omy and the availability of names for losa include Rana cacondana and Rana This serpentiform skink was described Angolan material still reuires further signata. Advertisement calls and genetics from Huilla Plateau (Bocage, 1870), investigation.. are crucial to resolving these issues. and the Tundavala material can therefore be considered topotypic material and Hyperolius cinereus Monard, 1937 the rst collected since the type descrip- An endemic from the Angolan plateau, tion. The species has a curiously disjunct historically known from only a few lo- distribution, with isolated populations calities, this species distribution has known from the Upemba Plateau, DRC, been found to be much more widespread NW Zambia, northern Serengeti, etc. (Conradie et al., 2016). Various subspecies have been proposed (Laurent, 1964) but have never been reas- sessed within a modern, integrated taxo- nomic framework. Whether the species is a complex of cryptic, vicariant species remains unresolved and is the subject of igure cellated saasteer Psammophylax rhombeatus ocellatus. ongoing research (Branch et al., in prep.).

Psammophylax rhombeatus ocellatus Chamaeleo anchietae Bocage, 1872 (Bocage, 1873) Double-scaled chameleons from Tundava- This taxon has morphological di erences la (Fig. 8) correspond morphologically to (Fig. 6) and is separated from the typical

igure ngolan reed frog Hyperolius cf race in South Africa, with a few scattered parallelus, breeding male. records in northern Namibia (Kaman- jab Broadley, 1977 Kaross Ho man, Hyperolius parallelus (Günther, 1989). Its status as a full species is cur- 1858) and Hyperolius benguellensis rently being investigated (Branch et al., (Bocage, 1893) in prep.). Both these species consist of super com- plexes in Africa. H. parallelus is closely Psammophis ansorgii (Boulenger, 1905) related to the problematic H. marmoratus, This snake is very poorly known. The and H. cf. benguellensis belongs to the type locality of the single type is vague igure Doublescaled chameleon eually unresolved H. nasutus complex, (Benguela to Bihe, Angola), and the Chamaeleo anchietae

B E 6 2018 401 the original description (Bocage, 1872). and lack of oologists. The specimens of tunity to acknowledge (in alphabetical As with Eumecia anchietae, this taxon Psammophylax rhombeatus ocellatus, order) Abdelaia Moyo, Afonso Va was described from Huilla ( Bocage, Chamaeleo anchietae, and Eumecia an- Pinto, Fernanda Lages, Julian Glos, Kai 1872) and also has similarly disjunct dis- chietae, recorded recently (this study and Schütte, Kerllen Costa, Pedro Va Pinto, tribution (i.e., Marunga Plateau, Katanga, Branch et al., unpub. data, and Va Pinto, Rasmus Revermann, Segunda dos San- Kivu, DRC Udungwa Mountains, Tan- pers. comm.), are the rst for many de- tos, and Valter Chissingui. ania). Various subspecies have been pro- cades in Angola. The type specimens for posed (Laurent, 1952), but their status re- all of the species described by Bocage, mains unresolved (possibly comprising a were lost in the 1978 re that burnt the complex of cryptic, vicariant species) and Lisbon Museum, and thus the rediscovery eereces is the subject of ongoing research (Branch of these species is critical for taxonomic et al., in prep.). studies and may facilitate the nomination BirdLife International (2017) Important Bird Areas factsheet: Tundavala. Downloaded of neotypes and the resolution of cryptic from http://www.birdlife.org on 28 June 2017. Trachylepis hoeschi (Mertens, 1954) diversity within disjunct populations. Bocage, J.V.B. (1870) Description dun Sau- This fat terrestrial skink was found in rien nouveau de lAfriue occidentale. Jor- nal de Sciencias Mathematicas, Physicas e Biodiversity montane grasslands, often near termite oservtio Naturaes, , 66–68. mounds in which it may shelter. It is The presence of a high number of en- Bocage, J.V.B. (1872) Diagnoses de uelues espces nouvelles de Reptiles dAfriue oc- endemic to Angola and Namibia, and demic amphibians and reptiles in the Tun- cidentale. Jornal de Sciencias Mathematicas, previously known in Angola only from davala region, though the taxonomic sta- Physicas e Naturaes, , 72–82. lowlands in Namibe province (Laurent, tus of many species remains unresolved, Bocage, J.V.B. (1895) Herpétologie d’Angola et du Congo. Ministrio da Marinha e das Col- 1964 Haacke unpub. data Branch un- highlights the relevance and urgency of nias, Lisbon. pub. data Ceraco et al., 2016). This e ective protection of the region. Further Bogert, C.M. (1940) Herpetological results of record constitutes an important range studies on the recorded species may re- the Vernay Angola Expedition with notes on African reptiles in other collections. Part 1, extension of the species onto the plateau, veal further cryptic diversity and thus am- Snakes, including an arrangement of African above 2200 m a.s.l.. Further studies will plify further the urgency for its protection. Colubridae. Bulletin of the American Museum help reveal if the species is tolerant to al- More comprehensive surveys in adjacent of Natural History, , 1–107. Boulenger, G.A. (1905) A list of the batrachians titudinal range, or if the upland popula- habitats will also likely increase the num- and reptiles collected by Dr. W.J. Ansorge in tion re ects cryptic speciation. ber of species recorded in the region. Angola, with descriptions of new species. An- In many respects, Tundavala rep- nals and Magazine of Natural History, ser. 7, 6, 8–115. resents only the tip of the iceberg when it Branch, W.R. (1998) Field guide to snakes and comes to the immense and poorly known other reptiles of southern Africa. Struik. Cape herpetological diversity of the Angolan Town. Broadley, D.G. (1977) A revision of the African escarpment. As Clark et al. (2011) have snake genus Psammophylax Fit Ingersoll highlighted, further research and conser- (Colubridae). Arnoldia (Rhodesia), , 1–44. Channing, A. (2001) Amphibians of central and vation are urgently needed. The study of southern Africa. Comstock Pub. Associates. the biogeography and herpetofauna of Cornell University Press. New ork. the rest of the Angolan escarpment is of Channing, A., Rödel, M.O. & Channing, J. (2012) Tadpoles of Africa: The biology and high priority. Herpetological surveys in igure Hoeschs sin Trachylepis identifi cation of all known tadpoles in sub- hoeschi northern regions of the escarpment, such Saharan Africa. Edition Chimaira, Frankfurt. as Kwana Sul and Kwana Norte, have Channing, A., Hillers, A., Lötters, S., Rödel, M.O., Schick, S., Conradie, W., Rödder, D., taken place and will be continued (Bap- Mercurio, V., Wagner, P., Dehling, J.M., Du Although the Angolan escarpment has tista et al., in prep.). Pree, L.H., Kielgast, J. & Burger, M. (2013) a nities with the adjoining biomes, it Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa acts as a barrier between the drier coastal (Anura: Hyperoliidae), with descriptions of plains and the inland plateaus, allowing six new species. Zootaxa, 3620, 301–350. speciation to develop (Huntley, 1974), coledgeets Clark, V.R., Barker, N.P. & Mucina, L. (2011) The Great Escarpment of southern Africa: a explaining the high level of endemism new frontier for biodiversity exploration. Bio- observed in the region (see Tab. 1 and The research was carried out in the diversity and Conservation, , 2543–2561. Conradie, W., Bills, R. & Branch, W.R. (2016) Tab. 2). At least ve of the recorded spe- framework of SASSCAL and was spon- The herpetofauna of the Cubango, Cuito, and cies consist of important rediscoveries of sored by the German Federal Ministry of lower Cuando river catchments of south-east- Angolan plateau endemics. Psammophis Education and Research (BMBF) under ern Angola. Amphibian & Reptile Conserva- tion, , 6–36. ansorgii and Leptopelis anchietae have promotion number 01LG1201M. This Conradie, W., Branch, W.R., Measey, G.J. & not been recorded for 60 and 53 years, work was improved with advice, records Tolley, K.A. (2012) A new species of Hyper- respectively, which is not a total surprise, of species, eldwork assistance, and ad- olius Rapp, 1842 (Anura: Hyperoliidae) from the Serra da Chela mountains, south-western given the lack of studies in Angola for ministrative, logistical, and technical Angola. Zootaxa, 6, 1–17. decades as a conseuence of the civil war support, for which we take this oppor-

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