First Herpetological Appraisal of the Parque Nacional De Banhine, Gaza Province, Southern Mozambique

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First herpetological appraisal of the Parque Nacional de Banhine, Gaza Province, southern Mozambique

Darren W. Pietersen1,2*, Errol W. Pietersen3 and Wulf D. Haacke4 1Department of Zoology and Entomology, University of Pretoria, Private Bag X20, Hatfield, Pretoria, 0028 South Africa 2Research Associate, Department of Vertebrates, Herpetology Section, Ditsong National Museum of Natural History, P.O. Box 413, Pretoria, 0001 South Africa 3Ecological Consultant, P.O. Box 293, Upington, 8800 South Africa 4Honorary Curator of Herpetology, Ditsong National Museum of Natural History, P.O. Box 413, Pretoria, 0001 South Africa PIETERSEN, D. W., PIETERSEN, E. W. and HAACKE, W. D., 2013. First herpetological appraisal of the Parque Nacional de Banhine, Gaza Province, southern Mozambique. Annals of the Ditsong National Museum of Natural History 3: 153–163. The herpetofauna of Mozambique remains poorly documented despite recent surveys. We present here the first report on the herpetofauna occurring in Parque Nacional de Banhine in Gaza Province.Four chelonian, 13 snake, one amphisbaenid, 16 lizard and 15 frog species were recorded in the park, while an additional six snake, five lizard and two frog taxa have been recorded in close proximity and probably also occur in the park. Numerous range extensions are recorded, and these are discussed in the light of our present knowledge of the distribution of herpetofauna in Mozambique. Keywords: Banhine National Park, Gaza Province, Mozambique, Reptiles, Amphibians.

INTRODUCTION Gaza Province (Fig. 1). PNB was proclaimed in The herpetofauna of Mozambique remains poorly 1972 after representation by Ken Tinley to protect sampled and our knowledge of the distribution of the vast numbers of game that roamed these plains. most species is scant. Most of our knowledge on The park lies between latitudes 22°30’–23°20’S and the Mozambique herpetofauna stems from ad hoc longitudes 32°15’–33°25’E, and consists of near- and often opportunistic sampling (Downs and pristine habitat. PNB is located in Gazaland, probably Wirminghaus, 1997) and as such the distribution of the driest province in Mozambique (Tinley, 1972), reptiles along the coast, especially in areas attract- with mean annual rainfall ranging from 399 mm in ing large numbers of tourists, is better known than the west to 427 mm in the east (Stalmans, 2003). the distribution of taxa inland. Broadley (1966, 1983) The topography is predominantly flat, with a slight and Poynton and Broadley (1985a,b, 1987, 1988) rise in elevation from 69 m above sea level (a.s.l.) in provided initial distributions for the herpetofauna of the southeast to 181 m a.s.l. in the northwest Mozambique, while more recently Broadley (1990, (Stalmans and Wishart, 2005; McNamara and 1992), Channing (2001) and Jacobsen et al. (2010) Larsen, 2006). Geology consists of deep sandy added to the slowly growing volume of literature. soils overlying calcrete at a depth of c. 7m Northern and central Mozambique have received (Stalmans and Wishart, 2005). Occasionally this more attention than southern Mozambique in recent calcrete reaches the surface and extends as years, with a number of new taxa recently being intermittent calcrete bands for a few hundred described from the former regions (e.g., Branch metres (Stalmans and Wishart, 2005; DWP pers. et al., 2005; Branch and Bayliss, 2009; Branch and obs.). Clayey sediments are present in the Tolley, 2010). northeast of the park (Stalmans, 2003) and are A herpetological survey of Parque Nacional de largely associated with closed woodlands along the Banhine was undertaken while one of the authors seasonal drainage lines and some of the mopane (EWP) was stationed there, with occasional visits by woodland communities (Stalmans and Wishart, DWP.Additional unpublished records from an earlier 2005). The grasslands and main wetland also have collecting trip by WDH are also included. a moderate to high clay content (Stalmans, 2003). Using the Koppen classification system, the study Study site area can be described as having a warm, dry Parque Nacional de Banhine (PNB) is a 700 000 ha climate with dry winters (van Rooyen et al., 1981). national park situated in southern Mozambique’s Eleven plant communities have been identified in *Author for correspondence. PNB, which can be further assigned to five major E-mail: [email protected] landscape types. These are (with their respective 154 ANNALS OF THE DITSONG NATIONAL MUSEUM OF NATURAL HISTORY, VOLUME 3, 2013

Fig. 1 Location of Parque Nacional de Banhine in Mozambique, and its geographic proximity to other major protected areas. Place names used in the text are indicated on the main map.

percentage area): wetland (1.1%), grassland PNB forms part of the Gazaland Plain (Smithers (13.6%), mopane woodland (33.9%), sandveld and Tello, 1976). This plain most probably has its (46.7%) and Androstachys forest (4.7%). A detailed origins in fluvial deposits of the Zambezi, Save, and vegetation map can be found in Stalmans and Limpopo Rivers, originating from headwater erosion Wishart (2005). The main wetland is located in the together with marine depositions prior to the northeast of the park and represents a seasonally Plio-Pleistocene (Moore and Larkin, 2001). During flooded deltaic system with limited overflow. The the Plio-Pleistocene, major flexing occurred along water level is governed by an alluvial plug (fan) at the Ciskei-Swaziland flexure axis (du Toit,1933; Par- the confluence of the main wetland and the tridge, 1998), resulting in the down-warping of the Changane River, formed by sediment deposition by margins of the subcontinent (King, 1967; Moore the Changane River during flooding events and Larkin, 2001) and ultimately the raised marine (McNamara and Larsen, 2006). A number of terrace that today is the Gazaland plain. smaller, cut-off pools occur along the margins of the main wetland, and permanent to semi-permanent MATERIAL AND METHODS pools usually persist in portions of the wetland Observations were made on an ad hoc basis during dry spells. A number of temporary pans also between November 2006 and May 2008. From form in the mopane woodlands after heavy rains. November 2006 to December 2007, the main wet- The grasslands form a large plain surrounding the land was dry owing to a prolonged drought, and wetland, and may also be seasonally inundated water remained only in a few scattered natural pans forming moist grassland. Sandveld is the dominant and man-made wells. In December 2007, the wet- landscape and occurs in the western, south-central land flooded for the first time in five years, and and eastern portions of the park, with some isolated remained inundated until at least May 2008. pockets in the north as well. Mopane woodland is Reptiles were actively searched for from a vehicle most prominent in the northern and central portions traversing roads by day and night, and by exploring of the park, with isolated pockets of varying sizes various habitats on foot. All likely microhabitats scattered throughout the remainder of the park. were actively sampled in search of reptiles and Androstachys forests are present in the south and amphibians. The old subterranean water tank at the extreme northeast border areas of the park. Pio Cabral park headquarters acted as a de facto PIETERSEN ET AL.: HERPETOLOGICAL SURVEY OF PARQUE NACIONAL DE BANHINE, MOZAMBIQUE 155 pit-fall trap and was checked regularly. The clearing some instances. Whenever lengths or scale counts of vegetation for the new airstrip also provided were recorded, these have been included in the opportunities to search for additional fossorial relevant species accounts. species. Frogs were actively searched for on a number of summer nights following rains. A search SYSTEMATIC ACCOUNT was also made of the herpetological records of the Ditsong National Museum of Natural History REPTILIA (formerly the Transvaal Museum), which houses TESTUDINES material collected by WDH on a much earlier field CRYPTODIRA trip to the Banhine swamps in 1964. Where taxo- TESTUDINIDAE nomic changes have been effected subsequent to this early collection trip specimens were Stigmochelys pardalis* (Bell, 1828) re-examined to confirm their identity. Records were One individual was observed near Cantine on the also gleaned from published sources. eastern boundary of the park and another was The distributions of species recorded from PNB noted on the edge of the main wetland to the west of were compared to the distributions as currently the old Pio Cabral homestead. A carapace of this depicted in major reference works (Broadley, 1983; species is on display in the warden’s caravan at the Branch, 1998; du Preez and Carruthers, 2009). park headquarters. WDH collected a specimen at Nomenclature follows the most recent list available Chigubo (TM 29330). from Johan Marais (available online at www. Kinixys zombensis* (Gray, 1831) reptileventures.com, downloaded on 5 June 2012), Asingleindividualwasobservedandphoto- with the exception of the families Lamprophiidae graphed in sandveld after crossing the road and Acontiinae. For the former family, we follow the between Xlekhane and Mungazi. nomenclature of Vidal et al. (2008). Current Acontiinae nomenclature follows Lamb et al. (2010). PLEURODIRA However, the findings of Lamb et al. (loc. cit.) have PELOMEDUSIDAE been called into question by S. Daniels (pers. comm.) and D.W. Pietersen et al. (in prep.) and we Pelomedusa subrufa* (Lacépède, 1788) therefore follow the former accepted nomenclature An individual was observed and photographed for this subfamily. on the edge of the wetland east of Fish-Eagle Most specimens collected were deposited in Research Camp. the Ditsong National Museum of Natural History Pelusios subniger subniger (Lacépède, 1788) (abbreviated TM) in Pretoria, South Africa. Some A specimen was collected in the Banhine Swamp specimens were also deposited in the Eduardo area by WDH in 1964 (TM 29316) and the anterior Mondlane National Museum of Natural History portion of a plastron was collected in front of (MHN) in Maputo, Mozambique, and in the National Fish-Eagle Research Camp by DWP (TM 85585). Museum Bloemfontein (NMB), South Africa. Acces- These records are 200 km east of the populations sion numbers have been incorporated into the known to occur in temporary pans in northeastern systematic account below. Photographs taken of Kruger National Park (KNP) (Jacobsen, 1989) and reptiles and amphibians in PNB (species marked 230 km southwest of the population on the San with a ‘*’ in the species accounts) have been lodged Sebastian Peninsula (Jacobsen et al., 2010). online and can be accessed at http://www. facebook.com/darrenpietersen.herpetology/pho- SQUAMATA tos SERPENTES Snout–vent length (SVL) was measured from the TYPHLOPIDAE tip of the snout to the posterior border of the anal plate with a Vernier calliper to an accuracy of Rhinotyphlops lalandei* (Schlegel, 1844) 0.02 mm, or with a material rule to an accuracy of A specimen was collected in mopane woodland 1 mm. Totallength (TL) was measured with a Vernier during the construction of the new airstrip calliper or material rule from the tip of the snout to (TM 85595), while two other individuals that were the tip of the tail. Ventral scales were counted from captured at the same location were examined and the first scale posterior to the chin shields to the last released. TM 85595 has SVL: 226.0 mm, TL: scale bordering, but excluding, the anal plate. 229.1 mm and 26 scale rows at midbody. All individ- Subcaudal scales were counted from the first scale uals were unearthed among the roots of trees that posterior to the cloacal opening to the tail tip. The were being cleared for the airstrip. These records number of upper labials (UL), lower labials (LL) and are 200 km east of the records in northeastern KNP number of scale rows at midbody were recorded in (Jacobsen, 1989). 156 ANNALS OF THE DITSONG NATIONAL MUSEUM OF NATURAL HISTORY, VOLUME 3, 2013

LEPTOTYPHLOPIDAE (Jacobsen, 1989), and 230 km southwest of the population on the San Sebastian Peninsula Myriopholis longicauda (Peters, 1854) (Jacobsen et al., 2010). WDH collected a specimen at Chigubo in 1964 (TM 29288). This record is c. 120 km east of the Xenocalamus bicolor lineatus* Roux, 1907 distribution for this species shown in Branch (1998), Three individuals (two adults and a juvenile) were although within the distribution presented by observed in tall mopane woodland at the new air- Broadley (1983). strip during bush-clearing activities. One adult was collected after being injured (TM 85594). This speci- BOIDAE men has SVL: 467 mm, TL: 498 mm, 17 scale rows at midbody, 242 ventrals, 26 subcaudals, 6 UL and 6 LL. Python natalensis A. Smith, 1840 Two large individuals were seen in the wetland Lamprophiinae during the aerial census in October 2004 (Stalmans, 2004). Three individuals were killed during 2007 by Boaedon capensis* (Duméril & Bibron, 1854) members of the local community who descended Encountered fairly regularly in the park, especially on the wetland to catch fish in the drying ponds (A. around Fish-Eagle Research Camp and on the new Guenya, pers. comm.). Owing to its large size and airstrip. One of the individuals observed displayed perceived threat to humans and livestock, together the xanthic colour form. with it being a potential source of protein, this species may be less common in the park at present Psammophiinae than it was historically. Hemirhagerrhis nototaenia (Günther, 1864) Broadley (1983) recorded this species in the vicin- LAMPROPHIIDAE ity of the Banhine Swamps. It is likely to occur in the Atractaspidinae extensive belts of mopane woodland Colopho- Atractaspis bibronii A. Smith, 1849 spermum mopane, a habitat that was not exten- A single individual believed to be referable to this sively covered during either of the survey periods. species was observed crossing the road between Rhamphiophis rostratus* Peters, 1854 Pio Cabral headquarters and Xlekhane. Broadley An adult was observed in sandveld on the road (1983) also recorded this species from the vicinity of between Xlekhane and Mungazi and was also the Banhine Swamps. recorded in this area by Broadley (1983). Amblyodipsas microphthalma microphthalma* Psammophis subtaeniatus* (Peters, 1854) (Bianconi, 1850) Commonly encountered in sandveld around Pio Two individuals were unearthed while clearing tall Cabral Headquarters and Fish-Eagle Research mopane woodland on the new airstrip. One of Camp. One individual examined had 8 UL and these, an injured individual, was collected (TM 120 subcaudals. Although the UL count may be 85596). This specimen (SVL: 363.5 mm, TL: indicative of P.orientalis, the high subcaudal count 400 mm) has 15 scale rows at midbody, 144 ventrals, and coloration confirm that it is P. subtaeniatus. 31 paired subcaudals, anal scale divided, 5 UL and Although these records are c. 100 km southeast of 6LL.ThereleasedindividualmeasuredSVL291mm, the distribution range indicated in Branch (1998), TL 319.5 mm and was unearthed amongst the roots Broadley (1983) recorded this species in the vicinity of a Grewia bicolor, about 40 cm below the soil of the Banhine swamps. surface. The tail : total length ratio (10.96) and number of Psammophis mossambicus Peters, 1882 ventral scales indicate that TM 85596 is a female. Individuals were seen foraging in the reeds at the The subcaudal count of TM 85596 is higher than edge of the wetland after it flooded. previously recorded for this species: Broadley (1983) records subcaudal counts ranging from COLUBRIDAE 18–26. Branch (1998) records the maximum SVL of Colubrinae this species to be 313 mm for a female and 320 mm Dasypeltis scabra (Linnaeus, 1758) for a male, while Broadley (1971b, 1983) gives maxi- A specimen was collected by WDH at Chigubo mum total lengths of 320 mm and 330 mm for males (TM 29286). and females, respectively. These measurements are surpassed by TM 85596, with a total length of Telescopus semiannulatus semiannulatus 400 mm (363.5 mm + 36.5 mm) and represents a A. Smith, 1849 new size record for this species. These records are This species was recorded from the vicinity of the 200 km east of the population in northeastern KNP Banhine Swamps by Broadley (1983). PIETERSEN ET AL.: HERPETOLOGICAL SURVEY OF PARQUE NACIONAL DE BANHINE, MOZAMBIQUE 157

Dispholidus typus typus (A. Smith, 1829) AMPHISBAENIA An adult female was captured and examined AMPHISBAENIDAE 10 km south of Xlekhane in sandveld. This female Monopeltis sphenorhynchus* Peters, 1879 displayed the typical brown colour pattern, and was Seven individuals were unearthed in mopane readily identified by the large eyes and presence of woodland during construction of the new airstrip, rear fangs. two of which were collected (TM 85590, TM 85591). TM 85590 has 28 dorsals and 19 ventrals per body ELAPIDAE annulus, 279 body annuli and 9 subcaudals; SVL: Elapinae 248.73 mm, TL: 258.34 mm. TM 85591 has 28 dorsals and 16 ventrals per body annulus, Aspidelaps scutatus fulafula* (Bianconi, 1849) 224 body annuli and 8 subcaudals; SVL: 83.28 mm, This species occurs widely throughout the park. TL: 88.10 mm. This species is likely to occur Individuals were observed in sandveld around throughout most of the park, but has been largely Fish-Eagle Research Camp (TM 85620), in mopane overlooked due to its fossorial habits. PNB records woodland near the northwestern boundary of the are 200 km east of the population in northeastern park and in the grassland plains around Xlekhane. KNP (Jacobsen, 1989) and 230 km southwest of the TM 85620 has 121 ventrals, 41 subcaudals and population on the San Sebastian Peninsula and 23 scale rows at midbody. Bazaruto Archipelago (Broadley, 1992; Jacobsen Elapsoidea sundevallii longicauda* Broadley, et al., 2010). 1971 An adult was captured and examined at SAURIA Fish-Eagle Research Camp after being unearthed SCINCIDAE from a Yellow Golden Mole (Calcochloris Acontiinae obtusirostris) tunnel and a second individual was Typhlosaurus sp. nov. Pietersen et al., in prep. captured 16 km north of Mungazi after it crossed the TM 85604–12, NMB R8840–43, 1251/MHN/09 road early in the morning. Morphological data for (five specimens). This new taxon is presently known the research camp individual are: 180 ventrals, only from PNB. Specimens were regularly encoun- 30 subcaudals, 13 scale rows at midbody, tered on the southern edge of the wetland in SVL 655 mm, TL 714 mm. The Mungazi individual sandveld habitat, wetland-fringing vegetation, in had 175 ventrals, 30 subcaudals, 13 scale rows at bush clumps and in mopane woodland during the midbody, 7 UL, 6 LL, SVL 816 mm, TL 899 mm. Both construction of the new airstrip, but may be more individuals had white bellies, a buff stripe bordering widespread in the park. the ventrals dorsally and a glossy black back. WDH also collected a specimen at Chigubo (TM 29287). Lygosomatiinae Note: Branch (1998: 106) confused the descrip- tions of E. s. longicauda and E. s. decosteri, errone- Mochlus sundevallii sundevallii (A. Smith, 1849) ously indicating that E. s. longicauda has a lower Two specimens were collected near Chigubo by number of ventrals compared to E. s. decosteri (see WDH (TM 29255, TM 29256). Broadley, 1971a). Trachylepis depressa (Peters, 1854) Naja annulifera* Peters, 1854 WDH collected a specimen 20 km southeast of A large male was captured in the research camp Machailla (TM 29248) and another at Chigubo ablution section while additional individuals were (TM 29280). This species was not observed during observed in the buffer zones around the park. 2007 and 2008. Trachylepis punctulata* (Bocage, 1872) Dendroaspis polylepis (Günther, 1864) Afairlycommonspecies,encounteredmost A single individual was observed about 20 km frequently in mopane woodland in the northeast of south of Chigubo. Although this record is the park, but likely to be more widespread. WDH extralimital, this species is likely to occur within the collected seven specimens at the Banhine Swamps park. (TM 29306–12) and four at Chigubo (TM 29251, TM 29281, TM 29282, TM 31254). A single speci- VIPERIDAE men was collected in mopane woodland to the north of Fish-Eagle Research Camp (TM 85582). Bitis arietans arietans* (Merrem, 1820) A common species observed in most vegetation Trachylepis striata* (Peters, 1844) types throughout the park. Individuals from PNB are This species was commonly observed throughout pale in colour. the park and was regularly encountered on build- 158 ANNALS OF THE DITSONG NATIONAL MUSEUM OF NATURAL HISTORY, VOLUME 3, 2013

ings and large baobab trees (Adansonia digitata). Chigubo (TM 29326), but the species was not WDH collected a specimen at the Banhine Swamps observed during the recent survey. TM 29326 has (TM 29313). four supraciliaries, a robust appearance, and closely resembles other G. nigrolineatus material Trachylepis varia* (Peters, 1867) housed in the Ditsong Museum collection. An adult was observed foraging at the edge of the wetland in front of the research camp at 22:05. A CORDYLIDAE juvenile and adult were subsequently observed in the research camp. Cordylus jonesii* (Boulenger, 1891) Afroablepharus walbergii* (A. Smith, 1849) A specimen was collected at Chigubo by WDH An uncommon species, but likely to occur (TM 29287), and a second individual was observed throughout the park. WDH collected two specimens and photographed in sandveld on the road between at the Banhine Swamps (TM 29304, TM 29305). Pio Cabral and Chigubo. One individual examined and photographed at Fish-Eagle Research Camp had five white spots VARANIDAE between the ear opening and the armpit insertion, Varanus albigularis albigularis (Daudin, 1802) with scattered white spots continuing along the Not observed during the recent observation period, flank to the groin, as does TM 29304. These individ- although WDH collected a specimen at Chigubo uals did, however, also have a vermiculated belly (TM 29289). and a white stripe bordering the dark lateral band ventrally, as is typical for A. walbergii. AGAMIDAE LACERTIDAE Agama armata* Peters, 1854 Heliobolus lugubris (A. Smith, 1838) Acommonterrestrialagamaoccurringwidely The most common lacertid in the park. Adults and throughout the park. WDH collected two specimens juveniles were regularly observed in sandveld and at Chigubo (TM 29284, TM 29285) while a gravid mopane woodland throughout the north of the park. female was collected in mopane woodland in the These records are approximately 100 km east of this northwest of the park in January (TM 85583). species’ distribution as plotted by Branch (1998). TM 85583 (SVL 80.52 mm, TL: 182.86 mm) has 89 scale rows at midbody, 10 UL and 11 LL. The Ichnotropis capensis (A. Smith, 1838) scales on the back are strongly keeled and directed A specimen was collected at Chigubo by WDH obliquely inwards towards the backbone. There are (TM 29272). This record is 230 km southeast of the six rows of enlarged, keeled spines along the body. nearest population in southeastern Zimbabwe The dorsal crest is small and the fifth toe does not (Branch, 1998) and 235 km southwest of the popu- extend as far as the first. This specimen corre- lation on the San Sebastian Peninsula (Jacobsen sponds well to other A. armata specimens in the et al., 2010). Ditsong Museum collection, and is clearly distinct Ichnotropis squamulosa* Peters, 1854 from the A. mossambica specimens housed in this WDH collected a series of six specimens from the collection. Banhine Swamps (TM 29319–TM 29324) and five from Chigubo (TM 29274–TM 29278). This species CHAMAELEONIDAE was encountered in summer and autumn virtually Chamaeleo dilepis* Leach, 1819 throughout the park. These records extend this An adult was observed in tall wetland-fringing species’ range 150 km east (see Branch, 1998). sandveld vegetation in the research camp during Nucras caesicaudata Broadley, 1972 light rain, while WDH collected specimens at the Specimens were collected by WDH at the Banhine Banhine Swamps (TM 29314, TM 29315) and at swamps (TM 29317, TM 29318) and at Chigubo Chigubo (TM 29258, TM 29325). This species prob- (TM 29279). Observations of this species appear to ably occurs widely throughout the park, but may be be closely correlated with termite emergences more common in sandveld habitat and in the vicinity following thunderstorms (Pienaar, 1978; Branch, of the wetland. 1998), which might explain the lack of subsequent observations. GEKKONIDAE Chondrodactylus turneri* (Gray, 1864) GERRHOSAURIDAE This species was commonly observed in Gerrhosaurus nigrolineatus Hallowell, 1857 Fish-Eagle Research Camp at night, often entering AsinglespecimenwascollectedbyWDHat buildings. PIETERSEN ET AL.: HERPETOLOGICAL SURVEY OF PARQUE NACIONAL DE BANHINE, MOZAMBIQUE 159

Hemidactylus mabouia Morreau de Jonnes, HEMISOTIDAE 1818 Hemisus marmoratus (Peters, 1854) Common on buildings and tents in the research WDH collected two specimens at Chigubo in camp and park headquarters. Also observed forag- February 1964 (TM 29547, TM 29548). This species ing along cement pathways at night. WDH collected is probably present in the park and may have been specimens at the Banhine Swamps (TM 29301) and overlooked during the recent survey. Chigubo (TM 29264, TM 29266–TM 29268). Hemidactylus platycephalus Peters, 1854 HYPEROLIIDAE WDH collected two specimens at both Chigubo Hyperolius argus Peters, 1854 (TM 29269, TM 29270) and the Banhine Swamps Individuals were observed on emergent vegetation (TM 29302, TM 29303). A single individual was in a small seasonal pan situated northeast of the observed on the stem of a large nyala berry research camp. This species probably occurs Xanthocercis zambesiaca tree near the edge of the throughout the main wetland, as well as in seasonal wetland in the vicinity of the old Pio Cabral home- pans with suitable emergent and waterside vegeta- stead. This species is likely to occur on large Adan- tion. sonia and Xanthocercis trees throughout the northern portions of the park. These records are 235 km Hyperolius marmoratus marmoratus Rapp, southwest of the nearest known population on the 1842 San Sebastian Peninsula (Jacobsen et al., 2010). WDH collected two specimens at Chigubo (TM 29547, TM 29548) where they occurred Lygodactylus capensis capensis* (A. Smith, sympatrically with H. m. taeniatus (TM 29545). 1849) These records are much farther north than most A common dwarf gecko recorded on buildings reference books depict their distribution (Channing, and trees throughout the park. WDH collected spec- 2001; Minter et al., 2004; du Preez and Carruthers, imens at the Banhine Swamps (TM 29292–TM 29299) 2009). However, Jacobsen et al. (2010) recorded and at Chigubo (TM 29271, TM 29273). both subspecies on the San Sebastian Peninsula, Pachydactylus punctatus* Peters, 1854 Vilanculos, together with what appears to be a An uncommon species found under leaves and hybrid individual and suggested that the peninsula logs in sandveld vegetation at Fish-Eagle Research may lie on a contact zone between the two subspe- Camp and the new airstrip. WDH collected speci- cies. These records suggest that H. m. marmoratus mens at the Banhine Swamps (TM 29300) and at is more widely distributed in southern Mozambique Chigubo (TM 29283, TM 29327, TM 29328). These than presently appreciated, that H. m. marmoratus records extend the known distribution of this species and H. m. taeniatus may occur sympatrically at c. 100 km east (see Branch, 1998). some localities, and that these two taxa may therefore represent distinct species. Further surveys are AMPHIBIA required to assess the distribution and taxonomic ANURA status of these taxa. BREVICEPTIDAE Hyperolius marmoratus taeniatus Peters, 1854 Breviceps adspersus adspersus Peters, 1882 Numerous individuals were seen and heard in the Calling individuals in the vicinity of the research pan to the northeast of Fish-Eagle Research Camp, camp were considered referable to this species, but as well as in the main wetland between the research this requires confirmation. camp and the alluvial plug. WDH collected specimens at the Banhine Swamps (TM 29554– Breviceps mossambicus Peters, 1854 TM 29557) and at Chigubo (TM 29545). This sub- Individuals presumably belonging to this species species is likely to be common in suitable habitat were heard calling in the vicinity of the research throughout the park. camp. This species requires confirmation. Kassina maculata Duméril, 1853 BUFONIDAE This species was heard calling in the small wetland to the northeast of Fish-Eagle Research Camp. Amietophrynus garmani * Meek, 1897 Numerous individuals were seen between Kassina senegalensis Duméril and Bibron, 1841 Fish-Eagle Research Camp and the salt pan to the WDH collected a specimen at Chigubo east of the alluvial plug. Individuals were also (TM 29553). A single individual was observed in the observed in a sunken cement well in the north of the research camp and others were heard calling in the park, and WDH collected a specimen at Chigubo vicinity. This species may occur in wet locations (TM 29549). throughout the park. 160 ANNALS OF THE DITSONG NATIONAL MUSEUM OF NATURAL HISTORY, VOLUME 3, 2013

MICROHYLIDAE subarticular tubercle beneath the first finger, as well as call. Specimens were discerned from Tomopterna Phrynomantis bifasciatus Smith, 1847 sp. Beira by the presence of one (not two) double A number of individuals were heard calling from subarticular tubercles beneath the first finger. the direction of the small wetland to the northeast of the research camp. Tomopterna sp. Banhine Additional Tomopterna individuals were observed PHRYNOBATRACHIDAE and photographed at Fish-Eagle Research Camp, but were not collected. These frogs had differing Phrynobatrachus sp. dorsal colour patterns and could not be positively Puddle frogs were heard calling from the small assigned to any presently recognized taxa. They pan to the northeast of the research camp, but we may represent undescribed species, or merely were unable to confirm which species were present. aberrant colour variations of described taxa. A description of the different forms follows: PTYCHADENIDAE Form A*: Two individuals seen in sandveld at Fish-Eagle Research Camp in January 2007. Metallic Ptychadena mossambica Peters, 1854 green dorsally, with glandular elevations coloured Numerous froglets were observed in vegetation gold (anteriorly) or yellow (medially) and enclosed fringing the pan to the northeast of Fish-Eagle by a broad black circle. A dark interorbital bar is Research Camp and adults were also heard calling present, broadest medially and directed back- from fringe vegetation at night. Adults were also wards. Flanks mottled with black. Concealed heard and observed between the research camp surfaces pale blue. The first individual changed and the alluvial plug after the wetland flooded. from metallic green to pale blue, especially laterally, while being photographed which may have been PYXICEPHALIDAE stress-induced. There is a glandular skin ridge Pyxicephalus edulis* Peters, 1854 present on the jaw-line, which continues to behind Numerous individuals were observed in sandveld the tympanum. The tympanum is obscured. A dead in the research camp, along the border of the wet- female containing eggs was observed in Fish-Eagle land between the research camp and the alluvial Research Camp in January, but was too degraded plug and in mopane woodland on the new airstrip to collect. Although it is possible that the observed following the onset of rains in November. It is likely to colour change (metallic green to light blue) was be common throughout most of the park, but its induced by envenomation, it is unlikely that two indi- presence may go unnoticed for many years owing viduals would be envenomated and not consumed to this species’ habit of remaining dormant until suit- at the same site and within four days of each other. able conditions prevail. The first individual also did not show any signs of envenomation (other than the colour change) while Tomopterna sp. Beira* Dawood et al., in prep. being photographed and hopped away upon A novel Tomopterna that was previously known release. The presence of two double proximal only from the vicinity of Beira, but confirmed to be tubercles beneath the first finger suggests that this present in PNB through genetic characterization form may be referable to Tomopterna sp. Beira. (Dawood, pers. comm.). Seven individuals were Form B*:SeeninsandveldatFish-EagleResearch collected in sandveld in Fish-Eagle Research Camp Camp. This form is pale pinkish dorsally and white (TM 85562–TM 85568). This species can be distin- ventrally. The dorsal surface is largely smooth, but guished from all other southern African species by with scattered glandular elevations. The lateral having two double proximal tubercles beneath the glandular elevations are orange; the median eleva- first finger and it is currently being described tions are elongated, reddish medially and surrounded (Dawood, pers. comm.). by a narrow black circle. A dark interorbital bar is Tomopterna cryptotis Boulenger, 1907 present, being broadest medially and pointing Acommonspecies,collectedatChigubo backwards, followed by a pale scapular patch. A (TM 29551, TM 29552) and at Fish-Eagle Research glandular skin ridge is present on the jaw-line but is Camp (TM 85589). not pronounced, and extends approximately to the posterior border of the concealed tympanum. Tomopterna krugerensis* Passmore and Small, conical orange elevations are present above Carruthers, 1975 the eyes. Specimens were collected in sandveld at Fish- Form C*: Seen in sandveld at Fish-Eagle Research Eagle Research Camp (TM 85588) and Chigubo Camp. This individual is pale grey dorsally and later- (TM 29546, TM 29550, TM 29599, TM 29600). Spec- ally, and white ventrally. The dorsal surface is imens were identified by the presence of a double smooth to slightly granular, with a mosaic of slightly PIETERSEN ET AL.: HERPETOLOGICAL SURVEY OF PARQUE NACIONAL DE BANHINE, MOZAMBIQUE 161 darker patterns on the pale ground colour, including then re-appear in northeastern South Africa or a darker inter-orbital bar that is broadest medially southeastern Zimbabwe. This apparent disjunct and points backwards, followed by a paler scapular distribution may be an attribute of inadequate patch. A pale glandular skin ridge is present but sampling rather than a true gap in distribution, an indistinct on the upper jaw, becoming distinct from hypothesis that is lent support by the discovery of behind the angle of the jaw and extending to the arm A. m. microphthalma, M. sphenorhynchus, T. auran- insertion. There is a darker patch behind the eye, tiacus species complex and I. capensis in PNB. extending across the concealed tympanum to the Conversely, some taxa with a mainly western distri- top of the arm insertion. bution extend eastwards along the sandy alluvium of northern South Africa, although few of these RHACOPHORIDAE species have been recorded on the Gazaland plain as yet. The occurrence of Monopeltis spheno- Chiromantis xerampelina* Peters, 1854 rhynchus and Trachylepis punctulata in PNB is Nests of this species were seen above temporary therefore significant and may portend that other pans throughout the park, including at Lipasse. species such as Chirindia l. langi, Monopeltis WDH collected a specimen 30 km southeast of leonhardi, M. rhodesiana, M. decosteri, Zygaspis Machailla (TM 29544). quadrifrons and Nucras intertexta could occur here as well. The former species are all fossorial and their DISCUSSION presence may only be noticed after many years Atotaloffourchelonian,18snake,one of continuous fieldwork. The apparent lack of amphisbaenid, 21 lizard and 17 frog species were Scelotes, a genus that is well-represented in north- recorded from PNB, or in the areas immediately eastern KNP, southeastern Zimbabwe and on the adjacent to the park. This herpetological diversity San Sebastian Peninsula and Bazaruto Archipel- (61 species) is lower than the diversity in southeast- ago, is surprising as this genus is usually located ern Zimbabwe (105), northeastern KNP (116) and quite easily when searched for. Further fieldwork in the San Sebastian Peninsula and Bazaruto Archi- all habitat types is required to ascertain whether this pelago (75), all of which are also on sandy sub- genus is truly absent, or whether it has merely been strates (Jacobsen, 1989; Broadley, 1990, 1992; overlooked. Branch, 1998; Jacobsen et al.,2010).Theapparently Three species of Tomopterna have already been lower diversity at the Mozambican sites might be recorded from PNB, although phenotypic traits due to less sampling having been done at these suggest that there may be as many as three addi- sites to date, but may also suggest that the habitat is tional undescribed taxa. One of the species recorded less complex at these sites, which in turn translates in PNB was only recently discovered near Beira for to a lower herpetological diversity. Although the the first time, and is still to be described. Ascer- current list almost certainly does not represent the taining the total diversity of Tomopterna in PNB is total herpetofaunal diversity of PNB, it does provide likely to prove an interesting avenue of research. a baseline that future studies can build on. The sympatric occurrence of Hyperolius m. Considering its geographic proximity, it is perhaps marmoratus and H. m. taeniatus is interesting, as not surprising that PNB shows the closest the former was considered to only occur further herpetological affinities to the northeastern KNP south and east (Channing, 2001; Minter et al., 2004; and southeastern Zimbabwe. Of the 61 species du Preez and Carruthers, 2009; but see Jacobsen recorded in PNB, 54 have also been recorded in et al.,2010).Althoughthesesubspeciesare northeastern KNP and/or southeastern Zimbabwe, supposed to occur parapatrically only in the vicinity while only 35 have been recorded on the San of St Lucia, KwaZulu-Natal, where intergrades have Sebastian Peninsula and/or Bazaruto Archipelago. been recorded (Lambiris, 1989; Minter et al., 2004), There are also a number of common species that Jacobsen et al. (2010) found them together on the have been recorded at the other three locations, but San Sebastian Peninsula, where an apparent hybrid not as yet in PNB, and which are expected to occur was also collected. These PNB records are the there as well. second known instance of sympatry in southern Also of interest are the distribution patterns of Mozambique, indicating that sympatry may be various taxa. A number of taxa such as Afrotyphlops more widespread than is currently believed. This fornasinii, Amblyodipsas m. microphthalma, suggests that H. m. marmoratus and H. m. taeniatus Zygaspis vandami, Monopeltis sphenorhynchus, may represent distinct species rather than merely Typhlosaurus aurantiacus species complex and geographic variants although further research, Ichnotropis capensis exhibit apparently isolated including molecular characterization, is required to populations in northern KwaZulu-Natal and southern investigate this possibility. It is also believed that Mozambique, but skip the Gazaland plain and H. m. marmoratus is more widely distributed in 162 ANNALS OF THE DITSONG NATIONAL MUSEUM OF NATURAL HISTORY, VOLUME 3, 2013

southern Mozambique than is currently appreciated. BROADLEY, D. G., 1990. The herpetofaunas of the islands The current survey of PNB, together with docu- off the coast of south Moçambique. Arnoldia Zimbabwe 9(35): 469–493. mentation of the long-overlooked specimens BROADLEY, D. G., 1992. Reptiles and amphibians from the collected by WDH, has started to fill some gaps in Bazaruto Archipelago, Mozambique.Arnoldia Zimbabwe the known distribution of a number of reptile and 9(38): 539–548. amphibian taxa in southern Mozambique. However, CHANNING, A., 2001. Amphibians of central and southern a great deal more sampling is required before we Africa. Cornell University Press, New York. can deduce phylogeographic patterns. The species DOWNS, C. T.and WIRMINGHAUS, J. O., 1997. The terrestrial checklist for PNB would benefit from further vertebrates of the Bazaruto Archipelago, Mozambique:A biogeographical perspective. Journal of Biogeography surveys, especially surveys employing traps to 24: 591–602. increase sampling ability. DU PREEZ, L. and CARRUTHERS, V., 2009. A complete guide to the frogs of southern Africa. Struik Nature ACKNOWLEDGEMENTS Publishers, Cape Town. DU TOIT, A. L., 1933. Crustal movements as a factor in the The authors thank the African Wildlife Foundation geographical evolution of South Africa. South African for providing accommodation during the latest Geography Journal 16: 3–20.As cited by Moore (1999). survey. Armando Guenya (PNB) is thanked for his JACOBSEN, N. H. G., 1989. A herpetological survey of the comments on the occurrence of pythons in PNB. Transvaal. Ph.D. thesis, University of Pretoria, South Abeda Dawood (National Zoological Gardens of Africa. South Africa) is thanked for commenting on the JACOBSEN, N. H. G., PIETERSEN, E. W. and PIETERSEN, D. W., 2010. A preliminary herpetological survey of the identity of Tomopterna specimens from PNB. Vilanculos Coastal Wildlife Sanctuary on the San Lauretta Mahlangu and Lemmy Mashinini are Sebastian Peninsula, Vilankulo, Mozambique.Herpetology thanked for providing access to specimens housed Notes 3: 181–193. in the Ditsong National Museum of Natural History KING, L. C., 1967. South African scenery. Oliver and Boyd, and for access to this museum’s electronic data- Edinburgh. As cited in Jacobsen (1989). base. Lauretta Mahlangu is also thanked for exam- LAMB, T., BISWAS, S. and BAUER, A. M., 2010. A phylogen- etic reassessment of African fossorial skinks in the ining the Gerrhosaurus nigrolineatus specimen. subfamily Acontiinae (Squamata: Scincidae): evidence Savel Daniels (Department of Botany and Zoology, for parallelism and polyphyly. Zootaxa 2657: 33–46. University of Stellenbosch) is thanked for his com- LAMBIRIS, A. J. L., 1989. A review of the amphibians of ments on the taxonomy of the Acontiinae. Mike Natal. Lammergeyer 39: 1–210. Bates (National Museum, Bloemfontein) and Niels McNAMARA, M.and LARSEN, E.J., 2006.Banhine National Jacobsen are thanked for greatly improving an Park, Gaza Province, Mozambique – Watershed assess- ment. Report 10/2006, 38 pp. earlier draft of this manuscript. MINTER, L. R., BURGER, M., HARRISON, J. A., BRAACK, H.H., BISHOP,P.J.and KLOEPFER, D., eds, 2004. Atlas REFERENCES and red data book of the frogs of South Africa, Lesotho BRANCH, W. R., 1998. Field guide to the snakes and other and Swaziland. SI/MAB Series #9. Smithsonian Institu- reptiles of southern Africa.ThirdEdition.Struik tion, Washington, DC. Publishers, Cape Town. MOORE, A. E. and LARKIN, P.A., 2001. Drainage evolution BRANCH, W. R. and BAYLISS, J., 2009. A new species of in south-central Africa since the breakup of Gondwana. Atheris (Serpentes: Viperidae) from northern Mozam- South African Journal of Geology 104: 47–68. bique. Zootaxa 2113: 41–54. PARTRIDGE, T. C., 1998. Of diamonds, dinosaurs and BRANCH, W. 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Appendix Gazetteer of localities in Parque Nacional de Banhine mentioned in the text.

Locality Latitude Longitude Altitude

Alluvial Plug 22°39’35”S 33°18’53”E 70 m a.s.l. Cantine 22°45’24”S 33°24’28”E 86 m a.s.l. Chigubo 22°49’54”S 33°31’11”E 103 m a.s.l. Fish-Eagle Research Camp 22°37’58”S 33°16’02”E 76 m a.s.l. Lipasse 22°28’46”S 33°03’10”E 80 m a.s.l. Machailla 22°15’14”S 32°54’56”E 113 m a.s.l. Mungazi 23°10’50”S 32°53’51”E 100 m a.s.l. New Airstrip 22°40’28”S 33°15’07”E 79 m a.s.l. Old Pio Cabral Homestead 22°37’41”S 33°14’38”E 75 m a.s.l. Pio Cabral Headquarters 22°38’04”S 33°15’46”E 86 m a.s.l. Salt Pan 22°39’33”S 33°21’16”E 68 m a.s.l. Xlekhane 22°44’46”S 32°57’11”E 81 m a.s.l.

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