the herpetofauna of the langjan nature reserve (limpopo

HERPETOZOA 15 (3/4): 121 - 135 121Wien, 30. Dezember 2002

The herpetofauna of the Langjan Nature Reserve(Limpopo Province, Republic of South Africa)

(Amphibia, Reptilia)

Die Herpetofauna des Langjan-Naturreservates(Provinz Limpopo, Republik Südafrika)

(Amphibia, Reptilia)

ALMUTH D. SCHMIDT

KURZFASSUNG

Das Langjan Naturreservat ist ein 4774 ha großes Schutzgebiet in der Limpopo Provinz Südafrikas, 130 kmnördlich der Provinzhauptstadt Pietersburg gelegen. Während einer Feldstudie von Januar bis April 1998 und dreikürzeren Aufenthalten zwischen 1999 und 2001 konnten innerhalb des Schutzgebietes insgesamt 43 Reptilien- (3Schildkröten, 23 Eidechsen, 17 Schlangen) und 7 Amphibienarten nachgewiesen werden. Die Anzahl der aus demGebiet bekannten Formen erhöht sich damit auf 47 bei den Reptilien und 10 bei den Amphibien. Die von derAutorin im Untersuchungsgebiet nachgewiesenen Arten werden hinsichtlich ihrer relativen Häufigkeit, ihrer allge-meinen Lebensraumansprüche und Verbreitung im Reservat charakterisiert. Neun weitere, bisher nur außerhalb derReservatsgrenzen nachgewiesene Reptilienarten kommen wahrscheinlich auch im Reservat selbst vor. Die Fund-ortbeschreibungen ausgewählter Formen werden durch ökologische, zoogeographische oder taxonomische Hin-weise ergänzt. Vier Arten {Psammobates oculiferus, Typhlosaurus lineatus subtaeniatus, Scelotes limpopoensisalbiventris, Colopus wahlbergii wahlbergü) verdienen aufgrund ihrer eingeschränkten Gesamtverbreitungsgebietebzw. Seltenheit besondere faunistische Beachtung.

ABSTRACT

The Langjan Nature Reserve, 4774 ha in size, is situated in South Africa's Limpopo Province, 130 km northof the Province's capital Pietersburg. During a field study from January to April 1998 and three shorter stays bet-ween 1999 and 2001, 43 reptilian (3 chelonians, 23 lizards, 17 snakes) and 7 amphibian species were recorded wit-hin the reserve. Thus, the known herpetofauna of the reserve now consists of 47 reptilian and 10 amphibian taxa.From the reptile and amphibian species recorded by the author, estimates of the abundance, the general habitatrequirements and distributions in the reserve are presented. Nine more reptile species are specified which are like-ly to occur in the reserve but were observed only outside its boundaries so far. The description of the records ofselected species in the reserve is complemented by considerations on ecological, taxonomic and distributionalaspects. Because of their endemic status, their generally limited distribution or rarity, four species {Psammobatesoculiferus, Typhlosaurus lineatus subtaeniatus, Scelotes limpopoensis albiventris, Colopus wahlbergii wahlbergii)are of specific faunistic interest.

KEY WORDS

Reptilia, Amphibia, Colopus wahlbergii wahlbergii, Psammobates oculiferus, Scelotes limpopoensis albi-ventris, Typhlosaurus lineatus subtaeniatus; check list, ecology, herpetofauna of Langjan Nature Reserve, LimpopoProvince, South Africa

INTRODUCTION

In the Limpopo Province of South Af- local climate account for the typical faunarica there is a number of small nature reser- and flora in the Reserve which are highlyves of special zoological interest. One of influenced by Kalahari elements. Unlike thethem is Langjan Nature Reserve, which was situation in mammals and birds, a completeestablished in the year 1954 to protect the checklist of the herpetofauna is lacking up tolast autochthonous population of the South now.African Gemsbok (Oryx gazelld) in former Within the short period of a few daysTransvaal. The geographical location of the JACOBSEN (1989) investigated the areareserve, its specific soil conditions and the during his reptile and amphibian survey of

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122 A. D. SCHMIDT

the Transvaal and some occasional collec-ting was carried out over the years. Twofield studies on species of the generaScelotes and Typhlosaurus occurring in thereserve were published (JACOBSEN 1987a,1987b). The rare subspecies Scelotes limpo-poensis albiventris JACOBSEN, 1987 was des-cribed during these studies based on speci-mens from Langjan Nature Reserve (= locustypicus).

Due to the conservation status of thestudy area, the main goal of this field studywas the investigation of aspects of reptileecology and population biology. The presentpaper documents the reptiles and amphibi-ans recorded during a 4 months researchstay from January to the end of April 1998and 3 additional stays in February 1999 (14days), January 2000 (7 days) and January2001 (8 days).

STUDY AREA AND METHODS

The Langjan Nature Reserve is situat-ed 130 km north of Pietersburg (now Polok-wane) between 22°47' - 22°52' S and 29° 11'- 29° 17' E. The topography of the reserve isplane at an altitude of approximately 800 mabove sea level (STUART & STUART 1993).The present size of the reserve is 4774 ha.The Brak River flows through the south-eastern part of the reserve (fig. 1). In 1994 adam was built at the foot of the Blouberg, 40km south of the Langjan Nature Reserve, toprovide water for citrus plantations. Sincethat time, the formerly seasonal Brak hasbecome temporary downstream of the damin the Langjan Reserve. The river conveyswater only a few days a year when thewaterlevel exceeds the crown of the damafter heavy rains. Throughout the rest of theyear only pools can be found along the riverbed.

Conglomerates, limestone and sand-stone form the lithic substratum of theground. The soil on the south-eastern side ofthe Brak River as well as in the centre of thereserve consists of deep red Kalahari sand(course sand - see fig. 2). Large areas of thereserve are covered by loam soil of differentcomposition.

The vegetation is of the 'Arid SweetBushveld' type (Veld Type 14 - ACCOCKS1988). The vegetation can be classified asshort open woodland with patches of tallshrubland. On the harder soils (Hardveld,fig. 3) in the north-western area of thereserve, the dominant plants include aGrewia flava veld with dense growth ofGrewia flava and Acacia erubescens, Acaciamellifera and Dichrostachys cinerea in vary-ing proportions, some Commiphora pyra-

canthoides and scattered taller trees ofBoscia albitrunca and Acacia tortilis. Insections where the soil is shallower, the veldbecomes more scrubby and shrubs likeCommiphora spp., Terminalia prunoides,Catoph-actes alexandri, Boscia foetida reh-manniana and Sesamothammnus lugardiiare common. Patches of Acacia thicketswith A. mellifera, A. karroo, A. nebrowniiand A. tortilis are characteristic. South-eastof the Brak River in the Sandveld (fig. 4),open woodland communities are dominant,represented by Sclerocarya caffra, Acacianigrescens, Terminalia sericea, Grewiabicolor, Combretum apiculatum, Commi-pora pyracanthoides, Grewia flavescenstogether with the common finger grassDigitaria eriantha and Enneapogon sco-par ias.

The average annual temperature isminimum in July (11 °C) and maximum inJanuary (35 °C). The average annual precip-itation is 300 - 400 mm (STUART & STUART1993), rain falling mainly in the summerbetween November and March.

During an intensive field study whichwas predominantly focussed on the lacertidfauna of the Langjan Nature Reserve, dataand records of other reptiles and amphibianswere collected as well. Within three repre-sentative study areas, different in vegetationand soil types, the amphibians and reptileswere trapped by daily controlled live trapsover a period of four months During thisperiod the traps were active altogether for 78days. The size of these areas varied between5 and 15 ha. They were situated south-eastof the Brak River and in the northern andwestern parts of the reserve. Twenty one live


Herpetofauna of the Langjan Nature Reserve 123

LANGJAN N. R.

/ \To Vivo

Camping huts / Campinghütten AWaterhole / Wasserloch ORoads / Straßen _

2000 m

Fig. 1: Map of the Langjan Nature Reserve (Limpopo Province, Republic of South Africa). Modified from SouthAfrica topographical maps 1:50.000 (sheets no. 2229CD and 2229CC), CTP Book Printers, Cape 1980.

Abb. 1: Karte des Langjan Naturreservates (Provinz Limpopo, Republik Südafrika). Verändert nach Topographi-sche Karten Südafrikas 1:50.000 (Blätter Nr.. 2229CD und 2229CC), CTP Book Printers, Cape 1980.


124 A. D. SCHMIDT

2000 m.\\v.v.v

Course Sand Course Sandy Loam Sandy Clay Loam

lockerer Tiefsand verfestigter Sand Toniger Lehm(grobkörnig) (mittelfein gekörnt)

Sandy LoamSandiger

Lehm

Clay

Ton

Fig. 2: Distribution of the soil types in the Langjan Nature Reserve (Limpopo Province, Republic of SouthAfrica) based on a guidemap of the Langjan N. R. authorities and own investigations.

Abb.2: Verteilung der Bodentypen im Langjan Naturreservat (Provinz Limpopo, Republik Südafrika). Karteerstellt nach Unterlagen der Reservatsleitung des Langjan Naturreservates und eigenen Untersuchungen.



trap systems were distributed over the studyareas. Each trap system was made of four 10or 20 liter buckets, which were dug in theground and connected through 10 meterlong, strong plastic fences, similar to classicamphibian fences. Additionally, these studyareas were monitored about twice a day inthe morning (between 07:00 and 12:00) andafternoon (between 15:00 and 19:00) bythree to four people. Altogether, nineteenspotlight surveys were made during nights,especially after rainfall, in order to recordnocturnal species. Additional spots of her-petological interest, like waterholes, riverbeds, and roadsides were observed fromtime to time during all stays.

Specimens were captured alive, stud-ied, determined and released after takingphotographs. Details of locality, habitat,date, sex and reproductive state were record-ed. The study was permitted by the NatureConservation Department of the Limpopo

Province. Permits for collecting of voucherspecimens were not generally granted, butissued under special circumstances only(e.g., for the new record of Colopus wahl-bergii wahlbergii PETERS, 1869). All vouch-er specimens are stored at the TransvaalMuseum (TM), Pretoria.

Scientific nomenclature and determi-nation of species follows for the reptiles(except Chelonians) FITZSIMONS (1943,1962) as modified by BALLETO (1968),BRANCH & BROADLEY (1985), BROADLEY(1968, 1972, 1977a,b,c, 1990), BROADLEY &WATSON (1976), HAACKE (1975, 1976),JACOBSEN (1987a) and LOVERJDGE (1957);for the Chelonians LOVERJDGE (1941) andLOVERIDGE & WILLIAMS (1957) as modifiedby BROADLEY (1981) and BOUR (1983) andfor the amphibians POYNTON (1964), asmodified by POYNTON & BROADLEY (1985a,b, 1987, 1988).

SPECIES ACCOUNT

In total, 50 reptile and amphibianspecies were recorded within the Reserve (3chelonians, 23 lizards, 17 snakes, 7 amphib-ians).

The species lists (table 1 - Chelonia,Serpentes, Sauria; table 2 - Amphibia), werecomplemented by ecological aspects anddata on the distribution and the frequency ofobservation in the reserve.

Seven reptile species were subject toadditional observations or are of herpetofau-nal interest and will be treated in more detailbelow.

Psammobates oculiferus (KUHL, 1820).All five tortoise records are localised in thesandveld area south-east of the Brak River.The specimens were observed betweenJanuary and early March 1998, mostly afterrainfall. The observation of five specimensof this rare and endemic species in one rainseason suggests that the population inLangjan N. R. seems not to be seriouslythreatened when compared to other popula-tions in the Limpopo Province or Botswana(W. HAACKE pers. com. 1999; P. SNYMANpers. com. 2000; own data). The status of thespecies in the Limpopo Province is vulnera-

ble owing to low densities and limited avail-able habitat (JACOBSEN 1989).

Pseudaspis cana (LINNAEUS, 1754).The Mole Snake appears to be abundant inthe sandy parts of the reserve. It was exclu-sively found during the day. Juveniles (10specimens) and subadults (3 specimens) wererecorded between mid Februar and midApril; adults were never found. Juveniles ofthis viviparous snake are normally born dur-ing March and April, with an average of 30 to50 young at a time (AUERBACH 1981 ; BROAD-LEY 1990). We observed the first juvenilesearlier (on the 1 lth and 26th of February) ina sandy area south of the Brak River.

Nucras intertexta A. SMITH, 1838. Intotal, four specimens of the endemic SpottedSandveld Lizard were recorded. During therain season of 1998 they were observed intwo localities characterized by sandy soiland more or less dense vegetation. Threelizards were adults, which emerged at theend of January and mid of March, one was ahatch ling recorded on April 7. This speciesseems to be the rarest lacertid lizard in thereserve. However, its apparent „rarity"could also be a result of the species' short


126 A. D. SCHMIDT

I

Fig. 3: A Grewia flava Veld (Hardveld) in the western part of the Langjan Nature Reserve(Limpopo Province, Republic of South Africa); in the background the Soutpansberg mountain range.

Abb. 3: Grewia flava Steppe (Hardveld) im westlichen Bereich des Langjan Naturreservats (Provinz Limpopo,Republik Südafrika); im Hintergrund ist ein Teil des Bergmassivs der Soutpansberge zu sehen.

Fig. 4: Sandveld open woodland communities, south-east of the Brak River(Limpopo Province, Republic of South Africa).

Abb. 4: Sandsteppe mit offenen Trockengehölzgemeinschaften, südöstlich des Flusses Brak(Provinz Limpopo, Republik Südafrika).



and irregular activity period and the solitarylivestyle. The Spotted Sandveld Lizard is afast-moving terrestrial species with an un-usal activity cycle. Based on own observa-tions from other field studies in the LimpopoProvince and long-term studies in captivespecimens, the main activity period seems tobe in the summertime, where it appearsirregularly for some days to search for food,disappearing afterwards again for someweeks. During unsuitable conditions nospecimen can be seen for several months.Because of this short and irregular „surfaceactivity" records of this species are general-ly rare.

Nucras holubi (STEINDACHER, 1882).Ten specimens were observed during therain season of 1998. The records wererestricted to four localities and to a fourweeks period from mid January to midFebruary. An additional specimen wasrecorded on January 11, 2000. All speci-mens were adults except for one juvenile(February 16). Nucras holubi prefers hardersubstrates and is restricted to sandy clayloam or sandy loam soil in the reserve.Twice the species was observed in groups(six and two specimens, respectively) duringthe middle of the day feeding on alate ter-mites which were swarming after rainfall.These Nucras specimens demonstrated thetypical themoregulatory behaviour of climb-ing into low bushes from time to time.

Typhlosaurus lineatus subtaeniatusBROADLEY, 1968. One specimen of the rareStriped Blind Legless Skink was recordedon January 11, 2000. The skink was foundunder rotten logs in a deep sand area southof the Brak River.

Typhlosaurus lineatus subtaeniatus isa localized form of the north-western part ofthe Limpopo Province, first described fromthe Great Saltpans north of the Soutpansberg(BROADLEY 1968). The typical form T. I. lin-eatus BOULENGER, 1887 is widespread in theKalahari and has presumably extended itsrange eastwards during an interpluvial peri-od. During the following pluvial period theranges shrank leaving isolated populationsin northern Venda (7̂ /. richardi JACOBSEN,1987), north-western Transvaal (T. I. sub-taeniatus BROADLEY, 1968) and in westernZambia (T. l.jappi BROADLEY, 1968) (BROAD-LEY 1968; JACOBSEN 1987b). The species is

listed in the South African Red Data Book asrestricted (BRANCH 1988).

Scelotes limpopoensis albiventrisJACOBSEN, 1987. Two specimens of the rarefossorial Limpopo Dwarf Burrowing Skinkwere recorded in a deep sand area south ofthe Brak River (fig. 5). One was caught inthe middle of March 1998 in a pitfall trap,the other one was found under rotten leaveson January 11, 2000 in the same area. Therare and endemic subspecies albiventris hasa very local distribution in just two adjacent1/4° squares in the Limpopo Province andwas first described from the Langjan N. R.(JACOBSEN 1987a).

The subspecies S. I. albiventris inhab-its a small area within the middle of the dis-tribution range of S. I. limpopoensis FITZ-SIMONS, 1930. So far, both forms were notfound in sympatry although they occupysimilar habitats (JACOBSEN 1987a) suggest-ing that S. I. albiventris is probably a fullspecies (BROADLEY 1994). The species islisted in the South African Red Data Book asrestricted (BRANCH 1988).

Colopus wahlbergii wahlbergii PETERS,1869. Fourteen specimens of the endemicKalahari Ground Gecko were recorded be-tween February and March 1998 in the sand-veld south of the Brak River. They were col-lected during night drives (between 19.30and 22.00) in a deep sand area. Juvenilesand adults co-occurred from Februaryonwards. During a night drive in January2001, two specimens were observed in theabove area and three more specimens northof the Brak River in the central part of thereserve, also in a deep sand area.

The record of this species is remark-able with regard to aspects of zoogeographyand taxonomy. The collected specimensshow high individual variation of the dorsalpattern (figs. 6 a,b,c). Some specimens havea vertebral row of dark-edged spots (fig. 6a).In other specimens the vertebral spots tendto form a pale band with dark, irregularlywavy edges (fig. 6b) as mentioned byHAACKE (1976). In one juvenile specimen,the dorsal colour pattern is very similar tothe pattern of the subspecies C. w. furciferHAACKE, 1976 described from the dune areaof the western and south-western Kalahari.This juvenile has a light vertebral band,extending from the root of the tail to the


128 A. D. SCHMIDT

Table 1: Reptiles detected by the author in the Langjan Nature Reserve (Limpopo Province, Republic ofSouth Africa) with remarks on their abundance, habitat requirements and distribution in the reserve. D - daily inhigh numbers; R - regularly in camps and on buildings; S - Sandveld; H - Hardveld.

Tab. 1: Die im Langjan Naturreservat (Provinz Limpopo, Republik Südafrika) von der Autorin nachgewiese-nen Reptilien mit Angaben zur relativen Häufigkeit, ihren Habitatansprüchen und der Verbreitung im Reservat.D - täglich in hoher Anzahl; R - regelmäßig an Häusern und festen Campinghütten; S - "Sandsteppe" (gekennzeich-net durch Sandböden); H - "Hartsteppe" (gekennzeichnet durch relativ harte Lehmböden); arboreal - baumbewoh-nend, arenicolous - sandbewohnend, (semi)fossorial - (mehr oder weniger) grabend, terrestrial - bodenbewohnend.

TaxonObservation frequency Special require- Distribution in the

Beobachtungs- Habitat ') ments/ besond. Reserve / Verbrei-häufigkeit Ansprüche ') tung im Reservat

Geochelom pardalis (BELL, 1828)Psammobates oculiferus (KÜHL, 1820)Pelomedusa subrufa (LACÉPÈDE, 1788)

Rhinotyphlops Mandel SCHLEGEL, 1844Leptotyphlops longicaudus (PETERS, 1854)Leptotyphlops conjunctus incognitus

BROADLEY & WATSON, 1976Python sebae natalensis A. SMITH, 1840Pseudaspis cana (LINNAEUS, 1754)Prosymna bivittata WERNER, 1903Psammophis subtaeniatus subtaeniatus

PETERS, 1882Psammophis brevirostris brevirostris

PETERS, 1881Philothamnus semivariegatus

semivariegatus (A. SMITH, 1840)Dasypeltis scabra (LINNAEUS, 1758)Dispholidus typus typus (A. SMITH, 1829)Elapsoidea sundevallii longicauda

BROADLEY, 1971Naja annulifera annulifera PETERS, 1854Naja mossambica PETERS, 1854Dendroaspis polylepis (GÜNTHER, 1858)Bitis arietans arietans (MERREM, 1820)Bitis caudalis (A. SMITH, 1839)

Typhlosaurus lineatus subtaeniatusBROADLEY, 1968

Scelotes limpopoensis albiventrisJACOBSEN, 1987

Lygosoma sundevallii sundevallii(A.SMITH, 1849)

Mabuya striata striata (PETERS, 1844)Mabuya varia (PETERS, 1867)Mabuya variegata punctulata (BOCAGE, 1872)Heliobolus lugubris (A. SMITH, 1838)Ichnotropis squamulosa PETERS, 1854Nucras intertexta A. SMITH, 1838Nucras holubi STEINDACHER,1882Pedioplanes lineoocellata lineoocellata

DUMÉRJL & BlBRON, 1839Gerrhosaurus flavigularis WIEGMANN, 1828Varanus albigularis (DAUDIN, 1802)Varanus niloticus (LINNAEUS, 1766)Agama armata PETERS, 1854Colopus wahlbergii wahlbergii PETERS, 1869Hemidactylus mabouia

(MOREAU DE JONNES, 1818)Homopholis wahlbergii (A. SMITH, 1849)Lygodactylus capensis (A. SMITH, 1849)Pachydactylus capensis capensis A. SMITH, 1846Pachydactylus turneri GRAY, 1864Pachydactylus punctatus PETERS, 1854Ptenopus garrulus garrulus (A. SMITH, 1849)

>305

>35011

4

4132

23

2

4

12

2

512

>3011

1

terrestrialterrestrialterrestrialfossorialfossorial

fossorial

terrestrialsemifossorial

fossorial

terrestrial

terrestrial

arboreal

terrestrialarboreal

semifossorial

terrestrialterrestrialterrestrialterrestrialterrestrial

fossorial

arenicolous

arenicolous

arenicolous

arenicolous

S /HSHSS

S

HSS

S/H

S

S/H

HS/H

S

S/HHH

S/HS

S

12

fossorial arenicolous

fossorial arenicolous

DD4DD410

8

221419

R

RR3R211

arborealterrestrialterrestrial (arenicolous)terrestrialterrestrialterrestrialterrestrial

terrestrial

terrestrialterrestrialterrestrialterrestrialterrestrial arenicolous

arboreal

arborealarboreal

terrestrialarboreal

terrestrialterrestrial arenicolous

S/HS/H

SS/HS/H

SH

S

SS/H

HS

C/3

S/H

HS/H

SS/H

GO

S

*> Übersetzungen zu den Spalteneinträgen siehe Tabellenlegende.



Table 2: Amphibians detected by the author in the Langjan Nature Reserve (Limpopo Province, Republic ofSouth Africa) with remarks on their abundance, habitat requirements and distribution in the reserve. S - Sandveld;H - Hardveld.

Tab. 2: Die im Langjan Naturreservat (Provinz Limpopo, Republik Südafrika) von der Autorin nachgewie-senen Amphibien mit Angaben zur relativen Häufigkeit, ihren Habitatansprüchen und der Verbreitung im Reservat.S - "Sandsteppe" (gekennzeichnet durch Sandböden); H - "Hartsteppe" (gekennzeichnet durch relativ harteLehmböden); arboreal - baumbewohnend, arenicolous - sandbewohnend, fossorial - grabend, semiaquatic - mehroder weniger aquatisch; terrestrial - bodenbewohnend.

Observation frequency Special require- Distribution in theTaxon Beobachtungs- Habitat " ments / besond. Reserve / Vorkom-

häufigkeit Ansprüche" men im Reservat

Bufo garmani (MEEK, 1897)Breviceps adspersus PETERS, 1882Phrynobatrachus natalemis (SMITH, 1849)Ptychadena anchietae (BOCAGE, 1867)Tomopterna mormorata (PETERS, 1854)Tomopterna cryptotis (BOULENGER, 1907)

Chiromantis xerampelina (PETERS, 1854)

' ) Übersetzungen zu den Spalteneinträgen siehe Tabellenlegende.

4261228

6 foam nests /6 Schaumnester

terrestrialfossorial

semiaquaticsemiaquatic

fossorialfossorial

arboreal

arenicolous

(arenicolous)(arenicolous)

HSHHSSH

Fig. 5: The rare endemic Limpopo Dwarf Burrowing Skink Sceloles limpopoensis albiventris JACOBSEN, 1987,has a very local distribution in the Limpopo Province and was described from specimens of the Langjan N. R.

(specimen without actual collaction number, kept alive in the Transvaal Museum, Pretoria).Abb. 5: Der seltene endemische Skink Scelotes limpopoensis albiventris JACOBSEN, 1987, besitzt eine sehr lokale

Verbreitung in der Provinz Limpopo und wurde nach Exemplaren aus dem Langjan Naturreservat beschrieben(Exemplar derzeit ohne Sammlungsnummer, Lebendhaltung im Transvaal Museum, Pretoria).


130 A. D. SCHMIDT

Fig. 6: The rare endemic Kalahari Ground Gecko Colopus wahlbergii wahlbergii PETKRS, 1869 found in theLangjan Nature Reserve represents the second South African record of this species.

a - Adult specimen with a vertebral row of dark-edged spots (TM 82185).b - Subadult specimen. The vertebral spots form a pale band with dark, irregularly wavy edges (TM 82I87).

c - Juvenile specimen. A light vertebral band extends from the root of the tailto the nape where it bifurcates (TM 8219I).

Abb. 6: Die seltene endemische Geckoart Colopus wahlbergii wahlbergii PETERS, 1869 aus dem LangjanNaturreservat repräsentiert erst den zweiten Nachweis dieser Art für Südafrika.

a - Adultes Individuum mit einer vertebralen Reihe von dunkel umrandeten Flecken (TM 82185).b - Subadultes Individuum. Die vertebralen Flecken sind zusammengeflossen

und bilden ein wellenförmiges Band mit dunkler Umrandung (TM 82187).c - Juveniles Tier. Ein helles, dunkel gesäumtes Vertebralband verläuft

von der Schwanzwurzel bis zum Nacken, wo es sich gabelt (TM 82191).



nape, where it bifurcates (fig. 6c). Furtherinvestigation of this highly isolated C.wahlbergii population is necessary to eluci-date the systematic relationship. A series often specimens from this locality has beendeposited in the Transvaal Musum, Pretoria(TM 82184-82193).

JACOBSEN (1989) simply gives anunconfirmed notice of C. w. wahlbergii inLimpopo Province based on a photo from„near Mopane". HAACKE (2001 pers. comm.)substantiated the occurrence of this species inthe Limpopo Province due to a collection ofnine specimens from the Farm Pylkop, Mes-sina district in 1991 (TM 7102-5, 81510).BRANCH'S (1998) record of C. w. wahlbergiiin the Limpopo Province goes back to thesespecimens deposited in the TransvaalMuseum. The record from the LangjanNature Reserve is the second observation ofthis species in the territory of South Africa.

Additional species recordedin the Langjan area

During the present study, a number ofadditional reptile species were recorded inthe Langjan area, however, not in thereserve proper but in its neighbourhood.Most of these taxa can be expected to occuralso inside the reserve, because of the pres-ence of suitable habitats: Leptotyphlops scu-tifrons scutifrons (PETERS, 1854), Atract-aspis bibronii A. SMITH, 1849, Aparallactuscapensis A. SMITH, 1849, Lamprophis fulig-inosus (BOIE, 1827), Lycophidion capernecaperne (A. SMITH, 1831), Psammophylaxtritaeniatus (GÜNTHER, 1868), Telescopussemiannulatus semiannulatus A. SMITH,1849, Acanthocerus atricollis (A. SMITH,1849), Panaspis wahlbergi (A. SMITH,1849).

DISCUSSION

During the field study in the LangjanNature Reserve, 50 taxa of the herpetofauna(43 reptiles, 7 amphibians) were recorded.Seventeen of these (14 reptiles and 3 am-phibians) were already listed by JACOBSEN(1989), who found 18 reptile and 6 amphib-ian species in the reserve during his reptilesurvey of the Transvaal. Four reptile species- Pelusios sinuatus (A. SMITH, 1838),Mabuya capensis (GRAY, 1830), Cordylustropidosterum jonesi (BOULENGER, 1891),Prosymna sundevallii lineata (PETERS,1871), and three amphibians - Cacosternumboettgeri (BOULENGER, 1882), Kassina sene-galensis (DUMÉRIL & BIBRON, 1841),Phrynomantis bifasciatus (SMITH, 1847)recorded by JACOBSEN (1989) were not ob-served during the present investigation.

As a result of the current survey, theknown herpetofauna of the area consists of 47reptile and 10 amphibian species (table 3).

Lizard taxa dominate (25 species), fol-lowed by snakes (18), amphibians (10) andchelonians (4 species). The dominance oflizard species is a commonly observed fea-ture of reptile species richness in aridregions, whereas in humid regions snakesare more diverse than lizards (JOGER &LAMBERT 1997). For example, HAACKE

(1984) found in the area of the SouthernKalahari Domain a similar relationship of 27lizards, 18 snakes, 3 chelonias, 5 amphis-baenia and 6 amphibians.

Among the lizard species, 16 are terres-trial, 3 fossorial and 6 arboreal. The highnumber of lizard species in the reserve can beattributed to the presence of large sandy areas.Seventeen lizard species occur in the Sand-veld and are mainly terrestrial, except Typhlo-saurus lineatus subtaeniatus, Scelotes limpo-poensis albiventris, Lygosoma s. sundevallii(A. SMITH, 1849) which are fossorial. Twoterrestrial lizard species Nucras holubi andVaranus niloticus (LINNAEUS, 1766), werefound only in the Hardveld.

Fourteen lizard species of the Sand-veld areas are terrestrial and separated intheir ecological niches by a number of para-meters, especially space, time (daily andannually), diet and foraging strategies, re-production and predator escape tactics. Thenocturnal niche is occupied by four geckospecies Pachydactylus c. capensis A. SMITH,1846, Pachydactylus punctatus PETERS,1854, Ptenopus g. garrulus (A. SMITH,1849) and Colopus w. wahlbergii. They areseparated among others in preferring differ-ent habitat conditions: Ptenopus g. garrulus


132 A. D. SCHMIDT

and Colopus w. wahlbergii are distributed inthe reserve only in deep sand and mainlyopen areas. On the other hand Pachy-dactylus c. capensis and P. punctatus favoura firmer compact sand structure with a moredense vegatation, extending their range alsoto sandy loam soil.

Ten terrestrial lizard species are diur-nal and separated by their different activitypatterns, as well as foraging strategies,reproductive behaviour, and microhabitatselection. Four of them are lacertid lizardsof the Sandveld areas, one of which being atypical sit and wait hunter [Pedioplanis I.lineoocellata (DUMÉRIL & BIBRON, 1839)],characterized by a long daily activity periodof up to 10 hours (NAGY et al. 1984). Theactive hunter Nucras intertexta has shortand irregular activity periods and a solitarylifestyle and, thus, is rarely seen. The othertwo active hunters Heliobolus lugubris (A.SMITH, 1838) and Ichnotropis squamulosaPETERS, 1854 are common and live syntopi-cally in dense populations. These twospecies are segregated from each other bydifferences in their activities, reproductiveand life cycles as well as anti-predatorbehaviour. Heliobolus lugubris is mainlyactiv in the morning during the rain season,whereas /. squamulosa shows a longer dailyactivity including an afternoon peak(SCHMIDT 2001). Heliobolus lugubris pro-duces up to three clutches of 4 to 6 eggs dur-ing one rain season and has a life periodbetween 3 and 4 years (SCHMIDT unpubl.),whereas /. squamulosa is an annual specieswith one clutch of 9 to 12 eggs at the end ofthe rain season (BROADLEY 1979; JACOBSEN1987c; SCHMIDT 2001). The juveniles of//.lugubris mimic noxious carabid beetles(HUEY & PIANKA 1977; SCHMIDT 1997a,b),and emerge between January and April.Ichnotropis squamulosa is a remarkably fastgrowing species, of which the juvenilesappear in November, reaching adulthood inMarch and April (BROADLEY 1979; JACOB-SEN 1987; SCHMIDT 2001). Thus, the juve-niles of these two species do not co-occur.

Between the two Mabuya species ofthe Sandveld area, a spatial separation dueto different preferences concerning the sub-strate and the degree of vegetation cover canbe observed. Mabuya varia (PETERS, 1867)is widespread in the reserve and can be

found in nearly every habitat, except in theopen and deep sand areas where the vegeta-tion cover is mainly restricted to tufts ofgrasses. There, Mabuya varia is replaced byMabuya variegata punctulata (BOCAGE,1872), which is restricted to this habitat typein the reserve.

The snake fauna of the reserve com-prises six fossorial species (Rhinotyphlopslalandei SCHLEGEL, 1844, Leptotyphlopslongicaudus (PETERS, 1854), L. conjunctusincognitus BROADLEY & WATSON, 1976,Prosymna bivittata WERNER, 1903, P. sun-devallii lineata, Elapsoidea sundevalliilongicauda BROADLEY, 1971), which isclearly more than in the lizard fauna (3).Concerning the habitat selection, terrestrialspecies (11) dominante among the snakes.Only two arboreal snake species wererecorded until now: Dispholidus t. typus (A.SMITH, 1829) and Philothamnus semivarie-gatus(A. SMITH, 1829).

As expected, amphibian diversity iscomparatively low in this highly arid envi-ronment in that the amphibian fauna is re-stricted to species which are independent ofopen water for reproduction (Brevicepsadspersus PETERS, 1882) or breed in tempo-rary [Tomopterna cryptotis (BOULENGER,1907), T. marmoratus (PETERS, 1854), Bufogarmani (MEEK, 1897), Chiromantis xeram-pelina (PETERS, 1854)] and shallow pools[Phrynobatrachus natalensis (SMITH, 1849),Ptychadena anchietae (BOCAGE, 1867)] andfeature a fast larval development.

The highest species diversity in thereserve was observed in the Sandveld habitatwith a total of 35 reptile species recorded. Bycontrast, the Hardveld holds fewer records(22 reptile species). The vegetation in theSandveld is much more diverse than in theHardveld, offering more ecological nichesand leading to a more diverse herpetofauna.Generally, a higher species richness can beaccommodated in structurally complex vege-tation than in structurally simple vegetation(HEATWOLE & TAYLOR 1987).

Most of the savanna reptiles are con-tinuously distributed within their range(POYNTON & BROADLEY 1978). Exceptionsinclude, among others, the arenicolous rep-tiles, which are most abundant in theKalahari sand areas and the coastal alluviumof the Mozambique plain (POYNTON &



Table 3: Preliminary numerical synopsis of rep-tile and amphibian taxa known from the LangjanNature Reserve (Limpopo Province, Republic of SouthAfrica) based on the author's observations and recordsof JACOBSEN (1989). The numbers of taxa exclusivelyrecorded by JACOBSEN (1989) are indicated in paren-theses.

Tab. 3: Vorläufige Anzahl der Reptilien- undAmphibien-Taxa des Langjan Naturreservates (ProvinzLimpopo, Republik Südafrika), zusammengestellt nacheigenen Untersuchungen und den Nachweisen vonJACOBSEN (1989). Die Anzahlen der Taxa, die nur vonJACOBSEN (1989) nachgewiesen wurden, sind inKlammern angegeben.

Taxon

CheloniaTestudinidaePelomedusidae

SerpentesTyphlopidaeLeptotyphlopidaeBoidaeColubridaeElapidaeViperidae

SauriaScincidaeLacertidaeGerrhosauridaeCordylidaeVaranidaeAgamidaeGekkonidae

I Reptilia

BufonidaeHyperoliidaeMicrohylidaeRanidaeRhacophoridae

E Amphibia

Number of formsAnzahl Formen

22(1)

1218(1)42

6(1)511(0219

11(1)2(1)5(1)1

4(1)

18(1)

25(2)

47(4)

10(3)

BROADLEY 1978). The Kalahari deep sandareas in the reserve facilitate the occurrenceof arenicolous species. Some of these spe-cies {Psammobates oculiferus, Prosymnabivittata, Typholosaurus lineatus subtania-tus, Lygosoma s. sundevallii, Helioboluslugubris, Pedioplanes l. lineo ocellata,Colopus w. wahlbergii, Ptenopus g. garru-lus) belong to the Kalahari group. Other

species are coastal arenicolous forms of theMozambique plains (Elapsoidea sundevalliilongicauda, Scelotes limpopoensis albiven-tris, see POYNTON & BROADLEY 1978).Arenicolous species originated in the eastextended their ranges westwards during thePleistocene. At the same time, species orig-inating from the west extended their rangeseastwards. The latter became possible dur-ing arid interpluvials when aeolic sandswere deposited over large areas in theTransvaal (KING 1967; POYNTON &BROADLEY 1978).

The Langjan Nature Reserve and adja-cent areas south to it is the type locality oftwo localised subspecies (Typhlosaurus lin-eatus subtaeniatus, Scelotes limpopoensisalbiventris) of otherwise widespread areni-colous lizard species (BROADLEY 1968;JACOBSEN 1987a,b, 1989). A further anom-aly from this area includes an ovoviviparouspopulation of Mabuya capensis (recordedby JACOBSEN 1989 for the Langjan N. R.),while the species is oviparous in northernTransvaal (BROWN WESSELS 1989). Thehighly isolated record of the rare endemicgecko species Colopus w. wahlbergii in thereserve is of faunistic interest, as it repre-sents the easternmost extension of its rangeand is only the second observation of thisspecies for South Africa. The above factssupport the opinion that the area was isolat-ed for a longer period (JACOBSEN 1989).

There is no doubt that our knowledgeof the reptile and amphibian fauna of theLangjan Nature Reserve is still incomplete.Common South African taxa such as Lam-prophis fuliginosus, Lycophidion capensecapense, Telescopus semiannulatus,Psammophylax tritaeniatus, Atractaspisbibronii, Aparallactus capensis, Lepto-typhlops scutifrons scutifrons, Panaspiswahlbergi, Acanthocercus africo I Us or Pyxi-cephalus edulis PETERS, 1854 can be expect-ed to occur in the reserve. A potential num-ber of about 70 species for the reserveseems possible.

ACKNOWLEDGEMENTS

I would like to thank Wulf HAACKE of theTransvaal Museum (Pretoria) who supplied informationand answered many inquiries. The Nature ConservationDepartment of the Limpopo Province (Johan BOTHA &

Lizanne NEL) kindly allowed me to pursue this study inthe reserve. Special thanks are given to the presentLangjan Nature Reserve Warden Anton VAN WETTENfor excellent accomodation and housing. I am most


134 A. D. SCHMIDT

grateful to Peter and Janine SNYMAN, Blouberg NatureReserve (Vivo, Limpopo Province), for their help,friendship and for all the valuable scientific discussion

and also to Dr. Michael GRUSCHWITZ (Bad Honnef), forreading and commenting on the manuscript.

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DATE OF SUBMISSION: September 16, 2002 Corresponding editor: Heinz Grillitsch

AUTHOR: Dipl. Biol. Almuth D. SCHMIDT, Institute of Ecology and Evolutionary Biology, University ofBremen / FB2, P.O. Box. 330440, D-28344 Bremen, Germany. < [email protected] >


the herpetofauna of the langjan nature reserve (limpopo

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