Herpetofaunal survey of the Ongeluksnek (Malekgalonyane

Introduction

The herpetofaunal richness of South Africa is considered to be amongst the highest in the world (Branch, 1998; Bates et al., 2014; Du Preez and Carruthers, 2017; Tolley et al., 2019). Even though South Africa’s herpetofauna is considered fairly well studied, new species are still being described (for example, frogs: Channing, 2012; Channing et al., 2013; Channing and Baptista, 2013; Conradie, 2014; Wilson and Channing, 2019; reptiles: Jacobson et al., 2014; Travers et al., 2014; Whiting et al., 2015; Heinicke et al., 2017; Conradie et al., 2018, Bauer et al., 2019; Bates and Stanley, 2020). Some areas of the country remain poorly surveyed, especially parts of the Northern Cape, Eastern Cape and North West provinces (Bates et al., 2014). Within the Eastern Cape Province, much work

has been done in the southern and western regions (e.g. Branch and Braack, 1987), while the northern and central areas associated with the former homelands of the Ciskei and Transkei remained poorly surveyed. In recent years a series of rapid biodiversity studies has increased our knowledge of herpetofauna occurring along the Wild Coast (in the former Transkei homeland) (Venter and Conradie, 2015) and the arid interior of the Eastern Cape Province (Conradie et al., 2016).

The Ongeluksnek (~Malekgalonyane) Nature Reserve (NR) is located in the north-eastern part of the Eastern Cape Province, on the border between South Africa and Lesotho. It is approximately 13 250 ha in extent. The surrounding land tenure is primarily communal and land use is predominantly large and small stock farming. The area around Ongeluksnek NR was farmed commercially until the early 1980s when land was expropriated during the formation of the erstwhile Transkei homeland. The lower lying portions of the area were allocated to communal or subsistence farmers while the catchment areas were set aside as a conservation area, the Ongeluksnek NR, which was declared in 1990. The reserve has a history of consumptive utilisation. Local farmers were previously allowed to graze the lowland areas of the reserve under a permit system. An inspection of vegetation condition in 1994 revealed that the area was heavily overgrazed and vulnerable to accelerated

Herpetology Notes, volume 13: 717-730 (2020) (published online on 25 August 2020)

Herpetofaunal survey of the Ongeluksnek (Malekgalonyane) Nature Reserve in the foothills of the

Drakensberg, Eastern Cape Province, South Africa

Werner Conradie1,2,* Brian Reeves3, Sandile Mdoko3, Lwandiso Pamla3, and Oyama Gxabhu3

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

2 School of Natural Resource Management, George Campus, Nelson Mandela University, George, South Africa.

3 Eastern Cape Parks & Tourism Agency, 17-25 Oxford Street, East London, South Africa.

* Corresponding author. E-mail: [email protected]

Abstract. The results of a herpetofaunal survey of Ongeluksnek Nature Reserve, Eastern Cape Province, South Africa are presented here. Combination of visual encounter survey methods and standard Y-shape trap arrays were used to conduct the survey. A total of 26 species (eight amphibians and 18 reptiles) were recorded, representing 29 quarter-degree grid cell records, of which 62% represented the first records for these units. Furthermore, we document the presence of three species of snakes (Crotaphopeltis hotamboeia, Hemachatus haemachatus and Homoroselaps lacteus) for the first time for the whole degree square of 3028 (approx. 100 km2). This study highlights the need to survey poorly known regions to enable us to understand and document the full distributional extent of species. We also discuss the impact of uncontrolled fires on the absence of grassland specialised species during our survey.

Keywords. Amphibia, Reptilia, karroid, conservation, biodiversity, fire

Werner Conradie et al.718

soil erosion. Recommendations were made to rest the vegetation and manage by fires, but these were not implemented. At this point the reserve was severely invaded by alien plants and subjected to frequent illegal grazing, uncontrolled fire and poaching of wildlife (ECPTA, 2012). In 1997 attempts were made by the post-apartheid government to improve the management of the reserve, in partnership with traditional leaders and surrounding communities. The Eastern Cape Parks and Tourism Agency (ECPTA) assumed responsibility for the reserve in 2004 but, due to limited resources, this organisation has experienced challenges in increasing management effectiveness on the reserve (ECPTA, 2012).

No formal herpetological surveys have been undertaken for this reserve, although in the late 1980s the late Bill Branch (Port Elizabeth Museum) conducted a brief one-day survey at Ongeluksnek NR, while Marius Burger (former Eastern Cape Nature Conservation) visited the reserve briefly in 1996. Both ‘surveys’ were

time-constrained and, between them, documented only four species of reptiles and amphibians.

This study forms part of a series of rapid surveys to improve biodiversity inventory data of protected areas managed by ECPTA within the Eastern Cape Province. The purpose of these surveys are to: a) develop and update the biodiversity inventories of protected areas, b) assess the biodiversity features in terms of conservation value, c) identify habitats and sites of conservation concern, and d) to create benchmark datasets for environmental change and anthropogenic impact research.

Material and Methods

Study Site.—Ongeluksnek NR falls within the Eastern Cape Province part of the Drakensberg Mountains (Fig. 1). The elevation is highest in the west, along the Lesotho border, at about 2,700 m above sea level (a.s.l.) and lowest in the east, where it drops to about 1,500 m a.s.l. A high-lying (at approximately 2,500 m a.s.l.) plateau occurs on the western boundary. The geology

Figure 1. Topographic map indicating the position of Ongeluksnek Nature Reserve (green polygon) and the quarter-degree grid cells it covers (blue lines: major rivers, grey lines: provincial or country boundaries).

of the reserve is predominantly of the Drakensberg Group, which is basaltic lava with minor sandstone, tuff and agglomerate. The Clarens Formation occurs on the lower-lying eastern portion of the reserve and comprises fine-grained sandstone and siltstone. More recent Quaternary alluvial deposits occur in the river systems (Johnson and Wolmarans, 2008).

The reserve is situated predominately within the Grassland Biome. The higher lying areas in the west are vegetated by Lesotho Highland Basalt Grassland. There are extensive wetlands on the plateaus in these areas and these are classified as Lesotho Mires. At intermediate elevations in the central part of the reserve, the vegetation is Southern Drakensberg Highland Grassland, while the lower lying areas in the east is East Griqualand Grassland. A small section of Mabela Sandy Grassland also occurs here (Mucina and Rutherford, 2006).

Ongeluksnek NR is mainly influenced by subtropical anticyclones and receives predominantly summer rain, principally as orographic rain from squall-line thunderstorms (ECPTA, 2012). Annual precipitation

is about 1,000 mm in the foothills and around 1,800 mm on the escarpment. Snow occurs with an average frequency of approximately eight days per year, mainly in winter, although snow falls have been recorded for all months of the year. Berg winds are prevalent between August and September while local topographically-induced winds occur as a result of solar heating of the ground during the day and cooling during the night (ECPTA, 2012).

Survey.—A herpetofauna survey was conducted in the reserve from 18 to 28 February 2019. The survey involved active searches and passive trapping. Four Y-shape trap arrays (each array consisting of three 10–15 m long, 50 cm high, drift fences positioned in a Y-shape with a single pitfall trap in the middle, two one-way funnels per fence, and a one-way funnel at the end of each arm, see Fig. 2 and Table 1) were set up in different habitats. An additional trap array (Trap 3) comprised a single straight drift fence of about 30 m that utilised six funnel traps with a pitfall trap in the middle.

Vouchers (two to five individuals) were collected for each species, while all other specimens were released

Herpetofaunal survey of the Ongeluksnek Nature Reserve, South Africa 719

Figure 2. Arial images demonstrating the arrangement of the four trap arrays employed at Ongeluksnek Nature Reserve. All images are orientated in a northern direction.

at capture site. Voucher specimens were humanely euthanised by injecting reptiles with, and submerging frogs in, tricaine methanesulfonate (MS222) solution (Conroy et al., 2009). Specimens were then fixed in 10% formalin before being preserved in ethanol. Genetic material was collected from each voucher specimen and stored in 96% ethanol. All specimens are deposited at Port Elizabeth Museum, South Africa.

Species Identification.—Species identifications were based on the use of relevant field guides or published identification keys (reptiles: Broadley, 1983; Branch, 1998; frogs: Channing, 2001; Du Preez and Carruthers, 2017; Channing and Rödel, 2019; tadpoles: Channing et al., 2012) and comparison to other material housed in the Port Elizabeth Museum. Nomenclature was based on established online databases (amphibian: Frost, 2020; reptiles: Uetz et al., 2020), updated where appropriate.

Data collection.—The Port Elizabeth Museum database and other major museums’ databases housing South African herpetological material were consulted in order to obtain data of voucher specimens previously collected within the boundaries of Ongeluksnek NR. Additionally, we consulted virtual museum platforms (e.g. ReptileMap ~ http://vmus.adu.org.za and iNaturalist ~ www.inaturalist.org) in order to obtain additional (non-vouchered) records.

Results

A total of 26 species, representing eight amphibian (four families) and 18 reptile (11 families: 10 lizards and eight snakes) species were recorded during the survey. Previous ad hoc surveys by Bill Branch in 1980 and Marius Burger in 1996 recorded four common species (Amietia delalandi, Semnodactylus wealii, Pseudocordylus subviridis and Trachylepis punctatissima), all of which were also recorded during

the current survey. No additional historic reptile records exist for the reserve, although one other reptile species (Psammophylax rhombeatus) has been recorded from the same Quarter Degree Grid Cell (QDGC). Although Branch and Burger also recorded additional species (Breviceps verrucosus, Strongylopus grayii and Xenopus laevis), these were not substantiated by voucher specimens and only mentioned in their personal unpublished field notes or internal reports. The South Africa Frog Atlas Project (Minter et al., 2014) also recorded additional amphibian records (Sclerophrys capensis and Tomopterna natalensis), mostly originating from the records of staff at the reserve. The additional records bring the total number of recorded amphibian species on the reserve to 11.

The survey documents a total of 29 QDGC species records, of which 83% of reptile and 27% of amphibian records are new records for those QDGCs (Table 2). Furthermore, three records, namely, Crotaphopeltis hotamboeia, Hemachatus haemachatus and Homoroselaps lacteus, represent the first full degree grid cell record for unit 3028 (approx. 100 km2). Four additional records from our survey represent only the second species record for the entire degree grid cell (Pachydactylus maculatus, Dasypeltis scabra, Gerrhosaurus flavigularis, and Trachylepis capensis). Our survey recorded one undescribed species in the Leptoptyphlops sylvicolus group (see species account).

Species Accounts.—The species accounts are grouped into three sections, Amphibia (frogs), Sauria (lizards) and Serpentes (snakes). Within these groups we ordered species accounts by Family and then alphabetically within these families. We provide information on voucher numbers and sampling localities (latitude and longitude in decimal degrees, and elevation in meters). Brief notes on identification, taxonomy, and/or natural history are also given when appropriate under

Table 1. Trap array sites for the survey, with brief habitat descriptions and numbers of days installed.Table 1. Trap array sites for the survey, with brief habitat descriptions and numbers of days installed.

Trap Site Latitude (S) Longitude (E) Altitude (m) Description Trap days

1 -30.33425˚ 28.35612˚ 1596 Rocky outcrop with young grass 8


3 -30.33087˚ 28.35196˚ 1540 Bank of artificial dam 8

4 -30.34124˚ 28.32787˚ 1770 Grassy Scrubland 7

5 -30.34236˚ 28.31880˚ 1767 Dry Wetland with young grass 7 Table 2. Summary of the number of species and subspecies recorded at Ongeluksnek Nature Reserve, and the number of new quarter-degree grid cell (QDGC) records for South Africa. Ongeluksnek Nature Reserve

Number of reptile species 18

Number of QDGC records 18

Number of new QDGC records 15

Percentage new QDGC records (%) 83

Number of amphibian species 11





‘Comments’. Abbreviations: TM – Distsong Museum of Natural History (formerly Transvaal Museum), DM – Durban Natural Science Museum.

Amphibia

Brevicipitidae

Breviceps verrucosus Rapp, 1842Plaintive Rain Frog

Comments: No new material was collected during the current survey. Branch (unpublished field notes) reported that he had heard numerous Breviceps (possibly B. verrucossus) calling during the evenings, after mist had rolled in and during the night’s rainstorm (approx. -30.33880˚ 28.27371˚, 2,199 m a.s.l., 22 November 1980). This population of rain frogs has historically been assigned to B. maculatus FitzSimons, 1947, but its distinctiveness from B. verrucossus was disputed by Passmore and Carruthers (1979) and Channing (1989) based on similarity of their advertisement calls.

Bufonidae

Sclerophrys capensis Tschudi, 1838Raucous Toad (Fig. 3A)

Material (2 specimens): PEM A14160, -30.29804˚ 28.36038˚, 1565 m a.s.l., 20 February 2019; PEM A14152, -30.33724˚ 28.34777˚, 1,682 m a.s.l., 21 February 2019.

Comments: Specimens were caught in traps and collected under stones.

Vandijkophrynus gariepensis gariepensis (Smith, 1848)Karoo Toad (Fig. 3B)

Material (4 specimens): PEM A14161, -30.33815˚ 28.25941˚, 2,359 m a.s.l., 26 February 2019; PEM A14166–7, PEM A14169, -30.33828˚ 28.25230˚, 2,519 m a.s.l., 26 February 2019.

Comments: The new material conforms to the typical form, V. g. gariepensis. Two subspecies are recognised, a largely karroid species, V. g. gariepensis and a montane grassland species, V. g. nubicolis. Despite concern over the validity of the latter subspecies (e.g. Branch & Braack 1989; Branch 1990; Bates & Haacke 2003), its status remains equivocal and a modern phylogenetic study is required to resolve its status.

Hyperoliidae

Kassina senegalensis (Duméril and Bibron, 1841)Bubbling Kassina

Material: PEM A14154 (tadpoles), -30.33001˚ 28.36814˚, 1,572 m a.s.l., 27 February 2019.

Comments: Calls were heard, but only tadpoles were collected, from a small dam at the main gate of the reserve.

Semnodactylus wealii (Boulenger, 1882)Rattling Frog

Material (2 specimens): PEM A6906, -30.33028˚ 28.35250˚, 1,580 m a.s.l., 24 October 1996: PEM A14155 (tadpoles), -30.33001˚ 28.36814˚, 1,572 m a.s.l., 22 February 2019.

Table 2. Summary of the number of species and subspecies recorded at Ongeluksnek Nature Reserve, and the number of new quarter-degree grid cell (QDGC) records for South Africa.

Table 1. Trap array sites for the survey, with brief habitat descriptions and numbers of days installed.

Trap Site Latitude (S) Longitude (E) Altitude (m) Description Trap days



3 -30.33087˚ 28.35196˚ 1540 Bank of artificial dam 8

4 -30.34124˚ 28.32787˚ 1770 Grassy Scrubland 7

5 -30.34236˚ 28.31880˚ 1767 Dry Wetland with young grass 7 Table 2. Summary of the number of species and subspecies recorded at Ongeluksnek Nature Reserve, and the number of new quarter-degree grid cell (QDGC) records for South Africa. Ongeluksnek Nature Reserve

Number of reptile species 18




Number of amphibian species 11





Comments: Calls were heard, but only tadpoles were collected, from a small dam at the main gate of the reserve.

Pipidae

Xenopus laevis (Daudin, 1802)Common Platanna (Fig. 3C)

Material (5 specimens): PEM A14156–9, -30.33001˚ 28.36814˚, 1,572 m a.s.l., 22 February 2019.

Comments: Both tadpoles and adults were collected in the small dam at the main gate of the reserve.

Pyxicephalidae

Amietia delalandii (Duméril and Bibron, 1841)Common River Frog (Fig. 3D)

Material (7 specimens): PEM A384, approx. -30.33815˚ 28.25941˚, 2,396 m a.s.l., 22 November 1980; PEM A7047, -30.33222˚ 28.35528˚, 1,574 m a.s.l., 24 October 1996, PEM A14162–4, -30.33611˚ 28.35800˚, 1,662 m a.s.l., 20 February 2019; PEM A14150–1, -30.33920˚ 28.33389˚, 1,685 m a.s.l., 25 February 2019.

Comments: This species was found to be very common in the reserve, associated with streams or seepages.

Cacosternum boettgeri (Boulenger, 1882)Boettger’s Caco (Fig. 3E)

Material (2 specimens): PEM A14165, -30.33611˚ 28.35800˚, 1,662 m a.s.l., 20 February 2019; PEM A14168, -30.33087˚ 28.35196˚, 1,540 m a.s.l., 23 February 2019.

Comments: Specimens were collected under stones far from water.

Cacosternum nanum Boulenger, 1887Bronze Caco

Material (tadpoles): PEM A14153, -30.33001˚ 28.36814˚, 1,572 m a.s.l., 22 February 2019.

Comments: Specimens were heard calling, but no adults were captured. Tadpoles were caught in the small dam at the main gate of the reserve.

Strongylopus grayii (Smith, 1849)Clicking Stream Frog

Comments: No material was collected. Branch (unpublished field notes) reported the presence of a juvenile Strongylopus grayii at the old farm Killarney, but it was not collected. This is a widely distributed

winter-breeding species that is expected in the reserve and is thus included in the final species list, although specimens need to be collected to validate its presence.

Tomopterna natalensis (Smith, 1849)Natal Sand Frog

Comments: No new material was collected during this survey. This record is based on a call recorded during the Frog Atlas Project and needs verification. Until such time, we include this record as it is within the species’ predicted range (Minter et al., 2014).

Reptilia

Squamata

Sauria

Agamidae

Agama atra Daudin, 1802Southern Rock Agama (Fig. 4A)

Material (4 specimens): PEM R24835, -30.34561˚ 28.31956˚, 1,841 m a.s.l., 22 February 2019; PEM R24845, -30.29247˚ 28.34484˚, 1,620 m a.s.l., 19 February 2019; PEM R24847, -30.29116˚ 28.34615˚, 1,588 m a.s.l., 19 February 2019; PEM R24854, -30.33611˚ 28.35800˚, 1,662 m a.s.l., 19 February 2019.

Comments: This species was commonly observed throughout most of the reserve basking on elevated positions on rocks or termite mounds.

Cordylidae

Cordylus cordylus (Linnaeus, 1758)Cape Girdled Lizard (Fig. 4B)

Material (2 specimens): PEM R24830, -30.33441˚ 28.33194˚, 1,689 m a.s.l., 21 February 2019; PEM R24836, -30.34433˚ 28.32208˚, 1,541 m a.s.l., 22 February 2019.

Comments: Specimens were collected from only a small part of the reserve associated with sandstone and basalt outcrops.

Pseudocordylus subviridis (Smith, 1838)Drakensberg Crag Lizard (Fig. 4C)

Material (19 specimens): PEM R2516–2523, 2665–69, approx. -30.33815˚ 28.25942˚, 2,399 m a.s.l., 22 November 1980; PEM R13365, -30.33889˚ 28.25556˚, 2,484 m a.s.l., 25 October 1996; PEM R24849–50,


-30.33853˚ 28.25525˚, 2,449 m a.s.l., 26 February 2019; PEM R24851–2, -30.33815˚ 28.25942˚, 2359 m a.s.l., 26 February 2019; PEM R24861, -30.33828˚ 28.25230˚, 2,519 m a.s.l., 26 February 2019.

Comments: Very common at higher altitudes on the mountain pass to Lesotho.

Gekkonidae

Pachydactylus maculatus Gray, 1845Spotted Gecko (Fig. 4D)

Material (1 specimens): PEM R24829, -30.33648˚ 28.35274˚, 1,670 m a.s.l., 25 February 2019.

Comments: One juvenile was collected under a stone. This is only the second record for the entire degree cell. The other record is from near Umhlangeni (PEM R13359‒60, -30.31056˚ 28.63833˚, 3028BC).

Figure 3. Plate of selected amphibians collected at Ongeluksnek Nature Reserve. A – Sclerophrys capensis (adult female), B – Sclerophrys capensis (juvenile), C – Vandijkophrynus gariepensis gariepensis, D– Xenopus laevis, E – Amietia delalandi, F – Cacosternum boettgeri.


Figure 4. Plate of selected lizards collected at Ongeluksnek Nature Reserve. A – Agama atra, B – Cordylus cordylus, C – Pseudocordylus subviridis (male), D – Pachydactylus maculatus (PEM R24829), E – Gerrhosaurus flavigularis, F – Nucras lalandii, G – Pedioplanis burchelli, H – Trachylepis capensis, I – Trachylepis punctatissima, J – Trachylepis varia (PEM R24841).


Gerrhosauridae

Gerrhosaurus flavigularis Wiegmann, 1828Yellow-throated Plated Lizard (Fig. 4E)

Material (5 specimens): PEM R24853, -30.34430˚ 28.29603˚, 1,926 m a.s.l., 26 February 2019; PEM R24868, -30.33087˚ 28.35196˚, 1,540 m a.s.l., 19 February 2019; PEM R24869, -30.33087˚ 28.35196˚, 1,540 m a.s.l., 21 February 2019; PEM R24870, -30.33087˚ 28.35196˚, 1,540 m a.s.l., 24 February 2019; PEM R24875, -30.34124˚ 28.32787˚, 1,770 m a.s.l., 24 February 2019.

Comments: This was one of the most common lizards caught in our traps. The record represents only the second one for the whole degree cell. Other records are near Zitapile (PEM R20388-9, -30.55472˚ 28.85417˚).

Lacertidae

Nucras lalandii (Milne-Edwards, 1829)Delalande’s Sandveld Lizard (Fig. 4F)

Material (5 specimens): PEM R24831, -30.33396˚ 28.33226˚, 1,674 m a.s.l., 21 February 2019; PEM R24832–3, -30.33724˚ 28.34777˚, 1,682 m a.s.l., 21 February 2019; PEM R24865, -30.33424˚ 28.35612˚, 1,596 m a.s.l., 19 February 2019; PEM R24867, -30.33473˚ 28.35295˚, 1,607 m a.s.l., 19 February 2019.

Comments: Numerous specimens were collected by hand under rocks, and others captured in traps.

Pedioplanis burchelli (Duméril and Bibron, 1839) Burchell’s Sand Lizard (Fig. 4G)

Material (6 specimens): PEM R24834, -30.33868˚ 28.34764˚, 1,691 m a.s.l., 21 February 2019; PEM R24839–40, -30.34561˚ 28.31956˚, 1,841 m a.s.l., 22 February 2019; PEM R24857–8, -30.33611˚ 28.35800˚, 1662 m a.s.l., 19 February 2019; PEM R24858, -30.33611˚ 28.35800˚, 1,662 m a.s.l., 19 February 2019.

Comments: Common in the reserve and associated with rocky areas.

Scincidae

Trachylepis capensis (Gray, 1831)Cape Skink (Fig. 4H)

Material (3 specimens): PEM R24828, -30.33814˚ 28.33870˚, 1,687 m a.s.l., 23 February 2019; PEM R24878, -30.34237˚ 28.31880˚, 1,767 m a.s.l., 22 February 2019; PEM R24879, -30.34237˚ 28.31880˚, 1,767 m a.s.l., 26 February 2019.

Comments: Only the second record for the entire degree grid cell. The other record is from west of Matatiele (-30.29833˚ 28.64528˚).

Trachylepis punctatissima (Smith, 1849)Speckled Rock Skink (Fig. 4I)

Material (11 specimens): PEM R2524–8, PEM R2698, approx. -30.33880˚ 28.27371˚, 2,199 m a.s.l., 22 November 1980; PEM R13366, -30.33889˚ 28.25556˚, 2,477 m a.s.l., 25 October 1996; PEM R24842–3, -30.33953˚ 28.31861˚, 1,788 m a.s.l., 19 February 2019; PEM R24859–60, -30.33611˚ 28.35800˚, 1,662 m a.s.l., 19 February 2019.

Comments: This is one of the most abundant reptile species observed in the reserve, in both natural habitat and among man-made structures (e.g. buildings, vulture hide).

Trachylepis varia (Peters, 1844)Variable Skink (Fig. 4J)

Material (3 specimens): PEM R24841, -30.34561˚ 28.31956˚, 1,841 m a.s.l., 22 February 2019; PEM R24844, -30.28843˚ 28.34070˚, 1,564 m a.s.l., 19 February 2019; PEM R24846, -30.29247˚ 28.34484˚, 1,620 m a.s.l., 19 February 2019.

Comments: This is another very abundant species observed throughout the reserve.

Serpentes

Colubridae

Crotaphopeltis hotamboeia (Laurenti, 1768)Red-lipped Snake (Fig. 5A)

Material (1 specimens): PEM R24876, -30.34237˚ 28.31880˚, 1,767 m a.s.l., 27 February 2019.

Comments: This represents the first record for the degree cell. The nearest other record is for Farm Lyndale (TM 49363, 3027DB).

Dasypeltis scabra (Linnaeus, 1758)Rhombic Egg-eater (Fig. 5B)

Material (2 specimens): PEM R24855–6, -30.33611˚ 28.35800˚, 1,662 m a.s.l., 19 February 2019.

Comments: This is only the second record for the whole degree grid cell. The nearest other record is at Cedarville, Mvenyane Mission (DM R18 and 22, -30.4950˚ 28.9375˚). A third specimen was collected from the Jordan River side of the reserve (-30.29247˚


Figure 5. Plate of selected snakes collected at Ongeluksnek Nature Reserve. A – Crotaphopeltis hotamboeia (PEM R24876), B – Dasypeltis scabra, C – Hemachatus haemachatus (PEM R24871), D – Homoroselaps lacteus (PEM R24864), E – Lycodonomorphus rufulus, F – Psammophylax rhombeatus, G – Psammophis crucifer, H – Leptotyphlops sp.


28.34484˚, 1,620 m a.s.l.), but escaped before being photographed or processed.

Elapidae

Hemachatus haemachatus Bonnaterre, 1790Rinkhals (Fig. 5C)

Material (1 specimen): PEM R24871, -30.33087˚ 28.35196˚, 1,540 m a.s.l., 24 February 2019.

Comments: This represents the first record for the full degree cell. The closest other record is at Farm Clifton (TM 49612, 3027DB).

Lamprophiidae

Homoroselaps lacteus (Linnaeus, 1758)Spotted Harlequin Snake (Fig. 5D)

Material (1 specimen): PEM R24864, -30.33424˚ 28.35612˚, 1,596 m a.s.l., 26 February 2019.

Comments: Broadley (1983) recognised three colour varieties: blotched, striped and barred. The striped form is mostly associated with inland montane grassland and our specimen conforms to it.

Lycodonomorphus rufulus (Lichtenstein, 1823)Brown Water Snake (Fig. 5E)

Material (2 specimens): PEM R24872, -30.33087˚ 28.35196˚, 1,540 m a.s.l., 19 February 2019; PEM R24873, -30.33087˚ 28.35196˚, 1,540 m a.s.l., 24 February 2019.

Comments: Both specimens were collected in a trap near a water body. The closest other record is 50 km east, from near Matatiele (PEM R21007, -30.46972˚ 28.93889˚).

Psammophylax rhombeatus (Linnaeus, 1758)Spotted Grass Snake / Skaapsteker (Fig. 5F)

Material (4 specimens): PEM R24862, -30.33152˚ 28.34776˚, 1,576 m a.s.l., 23 February 2019; PEM R24863–4, -30.33152˚ 28.34776˚, 1,576 m a.s.l., 25 February 2019; PEM R24877, -30.34237˚ 28.31880˚, 1,767 m a.s.l., 21 February 2019.

Comments: This is one of the most common snakes encountered during the survey.

Psammophiinae

Psammophis crucifer (Daudin, 1803)Cross-marked Grass Snake (Fig. 5G)

Material (2 specimens): PEM R24848, -30.29509˚ 28.37040˚, 1,553 m a.s.l., 19 February 2019; PEM

R24866, -30.33424˚ 28.35612˚, 1,596 m a.s.l., 23 February 2019.

Comments: This represents the first record for the QDGC. The closest record is from the Maclear region (Broadley, 1983).

Leptotyphlopidae

Leptotyphlops sp.Undescribed thread snake (Fig. 5H)

Material (3 specimens): PEM R24827, -30.33814˚ 28.33870˚, 1,687 m a.s.l., 23 February 2019; PEM R24837–8, -30.34561˚ 28.31956˚, 1,841 m a.s.l., 22 February 2019.

Comments: During a recent large-scale phylogenetic study, Leptoptyphlops specimens were collected from near Matatiele (PEM R18154–5, Fever Village, 79 km SW Cedarville, -30.53556˚ 28.82722˚), 60 km south-east of Ongeluksnek NR. It is part of the L. sylvicolus group (Adalsteinsson et al., 2009). True L. sylvicolus is known from forest habitat, and the above material is from grassland and is genetically different from the forest samples. Busschau (2019) expanded on this study and included the Ongeluksnek NR samples, documented additional genetic structure, and assigned the grassland clade of the interior of the Eastern Cape Province to an undescribed species. Further studies are underway to formally describe this grassland form.

Discussion

To the best of our knowledge our survey is the first formal reptile and amphibian inventory of Ongeluksnek NR. Although we found no threatened amphibian or reptile species, and most of the species encountered are widespread and common, our survey resulted in a considerable number of new QDGC records, and even a few new records for the entire 3028 degree grid cell. This study has thus contributed to the understanding of the distribution and occurrence of South Africa’s herpetofauna by filling a substantial spatial gap in survey effort and focusing on an area that has been poorly studied for herpetofauna.

As a result of the recent fires, the reserve had low habitat heterogeneity, being composed almost entirely of young grassland (at lower elevations) and young shrubland (at higher elevations). Habitat heterogeneity is generally positively correlated with species richness for herpetofauna and other taxonomic groups (see Báldi, 2008; Tews et al., 2003). The reserve may be at an early stage of faunal succession with respect to


herpetofauna, with several climax-stage species (those requiring mature vegetation, e.g. Chamaesaura aenea and C. anguina) being absent (see Letnic et al., 2004).

The relatively small home ranges of amphibians and reptiles make them vulnerable to local extinction after disturbances (Rios-López and Aide, 2007) such as fire. Fire is a natural occurrence in the Grassland Biome (Coombs, 2015) and the species occurring there have evolved mechanisms to deal with it (Russell et al., 1999). However, humans have directly and indirectly altered the pattern of fire in the landscape, changing aspects such as frequency, intensity and seasonality, and possibly pushing species beyond their evolved capacity to mitigate this disturbance (see Branch, 2014). The outcomes of our survey were probably influenced by the fact that almost the entire reserve had burned in the previous year. Responses of herpetofaunal assemblages to fire have been poorly studied (Russell et al., 1999; Ruthven et al., 2008). Data from Texas (United States of America) suggest that, in the long-term, fire could increase diversity and abundance of grassland herpetofaunal species, and that fire can be used to manage rangelands without negative effects on herpetofauna (Ruthven et al., 2008). Similarly, Masterson et al. (2008) found that the long-term fire history of grassland sites within the Suikerbosrand Nature Reserve in Gauteng, South Africa, had no significantly negative effects on herpetofaunal species richness or composition. However, the findings of these studies must be considered with the caveat that the sites examined contained unburned patches of intermediate-aged and older vegetation that allowed for recolonisation after fire. The near complete burn of Ongeluksnek NR, and large parts of the surrounding landscape, left virtually no such patches. As a possible consequence, we did not encounter the two specialised Chamaesaura species (C. aenea and C. anguina), which we suspect to be present in the reserve. These species do not seek refuge in burrows or rock crevices as some other reptiles do, but rather flee from fires (Coombs, 2015). Large fires will kill most individuals, leaving small populations surviving in unburned patches to later repopulate burned areas (Du Toit et al., 2003; Bates, 2014a, b). The absence of unburned patches in Ongeluksnek NR may have left no surviving populations to recolonise burned areas, and this may have resulted in local extinctions of these species. It is also possible that these species are present on the reserve, but were simply not encountered. Across their distributions both species are suspected to have undergone a substantial population reduction in the past two decades and may soon be regarded as threatened

(Bates, 2014a, b), with frequent anthropogenic fire being suspected as a major contributor. Tetradactylus spp. (another taxon that we expected to encounter during the survey, but did not find) are similarly vulnerable to fire (Bates, 2014c, d).

Over and above the direct effects of fire through mortality, fire also influences herpetofaunal diversity through its effect on vegetation cover. Vegetation cover is generally positively correlated with post-fire age, although post-fire vegetation accumulation can vary with land use (e.g. grazing intensity), the timing of fires, rainfall and soil conditions. Masterson et al. (2008) found vegetation cover to be the driver of herpetofaunal species richness at all sites surveyed in the Suikerbosrand Nature Reserve. If our survey effort is repeated at a later date, once the vegetation matures and cover increases, additional species may be recorded.

It is important that reserve management strives to allow for habitat heterogeneity on the reserve by not allowing large fires to burn down the entire reserve all at once. Firebreaks should be implemented, and fires should be responded to where feasible, according to an adaptive fire management plan. Formal protected areas are generally regarded as secure refugia for the conservation of herpetofauna (Tolley et al., 2019). However, if they are to serve as such, it is important that the habitat requirements of reptile and frog species are taken into account, and that appropriate reserve management is practised.

Acknowledgments. The authors would like to thank Mbuyiselo Gxashi (Reserve Manager: Ongeluksnek Nature Reserve) and his staff, including the field rangers, environmental monitors and the Working on Fire team, for their support of the survey. The authors would like to extend a special word of thanks to Sandile Mdoko for managing the logistics for the survey. Permits were issued by Eastern Cape Parks and Tourism Agency and ethical clearance was granted by Bayworld Ethics Committee (# 2017_02).

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Accepted by Anamarija Zagar


Documents

Herpetofaunal survey of the Ongeluksnek (Malekgalonyane