field guide to the amphibians and reptiles of … · of arusha national park (tanzania) field guide...

Charles Andekia Msuya is currentlya Chief Laboratory Scientist at theUniversity of Dar es Salaam. Heholds a Ph.D in Zoology on theecology of Amphibians and a FullTechnological Certificate from theInstitute of Science and Technology.He has a long field experienceand conducts field surveys onAmphibians, Reptiles, Birds andSmall Mammals in Tanzania.

Edoardo Razzetti graduated inBiological Sciences at the Universityof Pavia (Italy) in 1993. He devotesmost of his time to the study ofherpetology, including distribution,habitat preferences and taxonomy.He has published over 25 scientificpapers and is presently working onseveral zoology research andconservation projects with IstitutoOikos, the University of Insubria andthe University of Pavia.

EDOARDO RAZZETTI & CHARLES ANDEKIA MSUYA

FIELD GUIDE TO THE AMPHIBIANSAND REPTILES

OF ARUSHA NATIONAL PARK(TANZANIA)

FIE

LD

GU

IDE

TO T

HE

AM

PH

IBIA

NS

AN

D R

EP

TIL

ES

OF

AR

US

HA

NA

TIO

NA

LPA

RK

(TAN

ZA

NIA

)

EuropeanCommission

Lombardy Region(Italy)

TanzaniaNational Parks

University of Insubria,Varese branch (Italy)

University of Dar es Salaam(Tanzania)

Istituto Oikos(Italy and Tanzania)

(Pho

to P

aola

Mar

iani

)

C M G N


OF ARUSHA NATIONAL PARK (TANZANIA)

European Commission

Lombardy Region (Italy)

Tanzania National Parks

University of Dar es Salaam (Tanzania)

University of Insubria, Varese branch (Italy)

Istituto Oikos (Italy and Tanzania)

The publication of this book has been made possible through contributions from:

Primary forest on the slopes of Ngurdoto crater.

EDOARDO RAZZETTI & CHARLES ANDEKIA MSUYA



Acknowledgements:The study of the amphibian and reptile populations of the Arusha National Park hasbeen carried out in the framework of the Mount Meru Conservation Project (2000-2002), a joint effort of the University of Insubria, Varese branch, Tanzania NationalParks (TANAPA) and Istituto Oikos, funded by the European Union. The publicationof this guidebook, that includes the results of two field campaigns, has been possiblethanks also to financial contributions by the Lombardy Region (Italy). The authors aretherefore grateful to all the above mentioned institutions for their support. They extend especial thank to TANAPA for granting permission to work in the park, toArusha National Park’s staff, for its guidance, cooperation and assistance in the field,Istituto Oikos and University of Insubria staff for co-ordination and logistic.We are also grateful to the many people who have contributed to the completion ofthis book, and particularly to: Prof. Kim M. Howell for confirming the identifications;Rossella Rossi for the co-ordination and enthusiastic support; Paola Mariani forpreparing the distribution maps and carrying bags full of venomous snakes in her car;Paola Codipietro, Valeria Galanti, Silvia Porrini, Cesare Puzzi, Stefania Trasforini andArchiebald Temu for their assistance during fieldwork; Donald G. Broadley, JohnPoynton and Guido Tosi for the invaluable help on the revision of the text and scien-tific support; Franco Andreone, Michael Lambert, Ronald A. Nussbaum, JensBoedtker Rasmussen, Sebastiano Salvidio, Stefano Scali, Roberto Sindaco, RichardWahlgren and Martin Whiting for their useful suggestions and the bibliographic mate-rial, Stephen Spawls and Michele Menegon for providing some of the excellent slidesand finally Josephine Driessen for the editing.

Front cover: Hyperolius viridiflavus

© Copyright 2002 TANAPA

Text by: Edoardo Razzetti and Charles Andekia Msuya

Photography: All photographs by Edoardo Razzetti with the following exceptions:pages 42 and 47 by Michele Menegon, pages 43, 44, 64 and 70 byStephen Spawls.

Maps by: Paola Mariani

All rights reserved - Printed in Varese, Italy

5

CONTENTS

FOREWORD by Lota Melamari(Director General Tanzania National Parks) . . . . . . . . . . . . . . . . . . . . . 7

INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Aim of the booklet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Arusha National Park . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Arusha National Park amphibians and reptiles . . . . . . . . . . . . . . . . . . . 13Data collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

LIST OF SPECIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

Bufo gutturalis (Guttural Toad) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19Xenopus muelleri (Mueller’s Clawed Frog) . . . . . . . . . . . . . . . . . . . . . . 20Ptychadena mascareniensis (Mascarene Grass Frog) . . . . . . . . . . . . . 22Rana angolensis (Common River Frog) . . . . . . . . . . . . . . . . . . . . . . . . 24Strongylopus fasciatus merumontanus (Striped Long-toed Frog) . . . . . . 26Phrynobatrachus keniensis (Puddle Frog) . . . . . . . . . . . . . . . . . . . . . . . 28Hemisus marmoratum (Mottled Shovel-nosed Frog) . . . . . . . . . . . . . . . 30Kassina senegalensis (Bubbling Kassina) . . . . . . . . . . . . . . . . . . . . . . . 32Hyperolius viridiflavus ommatostictus (Painted Reed Frog) . . . . . . . . . . 34Hyperolius nasutus (Long Reed Frog) . . . . . . . . . . . . . . . . . . . . . . . . . 36Geochelone pardalis babcocki (Tropical Leopard Tortoise) . . . . . . . . . . 37Hemidactylus mabouia (Tropical House Gecko) . . . . . . . . . . . . . . . . . . 38Pachydactylus turneri (Bibron’s Thick-toed Gecko) . . . . . . . . . . . . . . . . 39Keys for the identification of the chameleonsof Arusha National Park . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40Bradypodion tavetanum (Kilimanjaro Two-horned Chameleon) . . . . . . . 41Chamaeleo dilepis (Common Flap-necked Chameleon) . . . . . . . . . . . . 42Chamaeleo gracilis (Gracile Chameleon) . . . . . . . . . . . . . . . . . . . . . . . 43Chamaeleo rudis (Ruwenzori Side-striped Chameleon) . . . . . . . . . . . . 44Chamaeleo jacksonii merumontanus(Meru Three-horned Chameleon) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46Agama agama (Rock Agama) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48Mabuya striata (Eastern Striped Skink) . . . . . . . . . . . . . . . . . . . . . . . . . 50Mabuya varia (Variable Skink) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51Lygosoma afrum (Peter’s Writhing-skink) . . . . . . . . . . . . . . . . . . . . . . . 52Panaspis wahlbergii (Wahlberg’s Snake-eyed Skink) . . . . . . . . . . . . . . 53

6

Adolfus jacksoni (Jackson’s Forest Lizard) . . . . . . . . . . . . . . . . . . . . . . 54Nucras boulengeri (Boulenger’s Scrub-lizard ) . . . . . . . . . . . . . . . . . . . 55Leptotyphlops scutifrons merkeri (Merker’s Worm-snake) . . . . . . . . . . . 56Python natalensis (Southern African Python) . . . . . . . . . . . . . . . . . . . . 58Bitis arietans (Puff Adder) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60Elapsoidea loveridgei loveridgei (Loveridge’s Garter Snake) . . . . . . . . . 62Naja haje (Egyptian Cobra) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63Dendroaspis angusticeps (Green Mamba) . . . . . . . . . . . . . . . . . . . . . . 64Lamprophis fuliginosus (Brown House-snake) . . . . . . . . . . . . . . . . . . . 65Lycophidion capense jacksoni (Jackson’s Wolf-snake) . . . . . . . . . . . . . 66Psammophis phillipsii (Olive Grass Snake) . . . . . . . . . . . . . . . . . . . . . . 67Natriciteres olivacea (Olive Marsh-snake) . . . . . . . . . . . . . . . . . . . . . . . 68Crotaphopeltis hotamboeia (White-lipped Snake) . . . . . . . . . . . . . . . . . 69Thelotornis mossambicanus (Mozambique Twig Snake) . . . . . . . . . . . . 70Dasypeltis scabra (Common Egg-eater) . . . . . . . . . . . . . . . . . . . . . . . . 71

OTHER SPECIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

CHECK LIST OF AMPHIBIANS AND REPTILESAT THE ARUSHA NATIONAL PARK . . . . . . . . . . . . . . . . . . . . . . . . . . . 75

REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

7

FOREWORD

The breathtaking scenery of Mount Meru and Kilimanjaro is not the only rea-son for visiting Arusha National Park. Rather, the extraordinary quality of thepark lies in the variety of its landscapes and habitats, ranging from opensavannah to montane forests, lakes, marshes and rocky peaks, hosting ahighly diverse wildlife community. Therefore visitors, during their walksthrough the park, besides enjoying the view of an elephant roaming in the for-est, have the possibility to discover the enchanting world of the minorspecies: birds, butterflies, frogs, snakes. The Arusha National Parks her-petofauna seems to be particularly interesting and diverse.In the last 50 years many species of amphibians throughout the world havedeclined markedly in numbers, also within apparently pristine habitats, suchas national parks and nature reserves. Concern is so high that the SpeciesSurvival Commission of the International Union for the Conservation ofNature (IUCN) established the Declining Amphibian Populations Task Forceto collect and monitor data on amphibian populations and to assess theirgeographic distribution, their decline and possible causes. In order to con-tribute to this international effort, TANAPA decided to gather updated infor-mation on the Arusha National Park herpetofauna and to keep monitoring itspopulations. This booklet is the result of the joint effort of two naturalists: Charles Msuyafrom the University of Dar es Salaam and Edoardo Razzetti from theUniversity of Pavia. They have been working within the framework of theMount Meru Conservation Project (2000-2002), a joint effort of the Universityof Insubria, Varese branch, Tanzania National Parks (TANAPA) and IstitutoOikos, funded by the European Union and aimed at studying and preservingthe Arusha National Park biodiversity. The authors’ competence, commit-ment and enthusiasm, together with their ability to take beautiful photo-graphs, resulted in this fine work, that is expected to gently guide visitors intothe fascinating world of amphibians and reptiles, two groups of animals,which have been on earth so much longer than man.

LOTA MELAMARI

(Director GeneralTanzania National Parks)

11

INTRODUCTION

Aim of the booklet

Tanzania is internationally recognised as a key country for the conservationof African biological diversity. Its herpetofauna numbers about 130 amphib-ians and over 275 reptiles, many of them strictly endemic and included in the“IUCN” Red lists of different countries. This unique resource is still relativelyunknown even if amphibians and reptiles are ideal subjects for zoologicalinventories and biogeographical analysis as they are relatively easy to sur-vey and often strictly related to a particular environment. Moreover, since1989 the scientific community has started to realise that amphibians aredeclining in many areas of the world and that they are more sensitive thanother species to diverse environmental modifications. This is probably due tothe fact that their larval and adult stages occupy different habitats and havelimited vagility (Stebbins & Cohen, 1995; Houlahan et al., 2000). Despite its importance, the Arusha National Park herpetofauna has neverbeen completely studied up to now, even if some scientific papers showedalready its peculiarity and importance. The aim of this booklet is to fill the existing gap in the literature and to pro-vide a stimulus that will strengthen ecological tourism in the park. Visitors willbe encouraged to appreciate also this fascinating and a bit mysterious com-ponent of the ecosystems.

Arusha National Park

Arusha National Park is situated on the eastern slopes of Mt. Meru inTanzania. The area lies on the eastern edge of the Great Rift Valley. Thegeology and soils dominating much of the park and Mt. Meru area are vol-canic by origin, resulting from the activity of the mountain. The volcanicnature of Mt. Meru began during the Pleistocene, forming the Meru calderaand several minor craters including Ngurdoto Crater. About six thousandyears ago the eastern part of Meru caldera collapsed forming an extensivelahar and the closed alkaline lakes. The only lake which has an outflow watersystem is the Small Momela, which empties into Big Momela Lake.Continued volcanic activity built an ash cone in Meru Crater, an attractive fea-ture on Mt. Meru. A combination of climatic changes and river flows haveinfluenced the concentration of alkali in the lakes and therefore the biologicaldiversity and distribution of organisms. The highest biological diversity isfound in Lake Longil, which has relatively low alkaline levels.

12

The vegetation of Arusha National Park follows an altitudinal zonation(Hedberg, 1951). The lower altitude (1440-1700 m) vegetation cover variesfrom shrubland, thicket and bushland to dry evergreen forest, whereDiospyros abyssinica (Hiern) and Olea hochstetteri Baker are common. The mid altitude (1700-1800 m) vegetation on Mt. Meru is dominated by anevergreen mist fed forest, with Olea hochstetteri, Assearis, Croton, Ficus andNuxia sp. On the walls of Ngurdoto Crater Cassipourea malossana (Baker)dominates.The higher altitude (1800-2100 m) forest is dominated by Juniperus,Podocarpus, Ilex, Xymalos, Afrocrania sp. and several epiphytes. Plant com-munities around Meru caldera are mainly pioneers.Most of the lakes are very alkaline and open with Cyperus leavigatus domi-nating at the edge. Lake Longil has a less alkaline environment and littoralvegetation, with Cyperus, Papyrus and Typha sp. dominate. The lake is alsocovered with Nymphaea caerulea, Ceratophyllum demersum and Pistia stra-tiorates.

Pitfall traps at Lokie swamp; many species of amphibians can be found in this area including:Xenopus muelleri, Ptychadena mascareniensis, Phrynobatrachus keniensis, Kassina sene-galensis, Hyperolius viridiflavus and Hemisus marmoratum.

13

Arusha National Park amphibians and reptiles

The Arusha National Park herpetofauna has never been completely studiedalthough some specimens were collected in the Mt. Meru area during theSwedish scientific expedition in East Africa at the beginning of the last cen-tury (Andersson, 1911; Lönnberg, 1910, 1911). Later (1956-1957) somechameleons and a few other reptiles were collected by the hunters and snakeexperts C.J.P. Ionides and Lt. Col. J. Minnery (Loveridge, 1959; Rand, 1958;1963). Finally, a paper about the most common snakes was posthumouslypublished by the Scientific Officer of Tanzania National Parks, DesmondFoster Vesey-FitzGerald (1975).

The Arusha National Park is particularly interesting for amphibians and rep-tiles because (1) there are still large areas of montane forest, (2) the moun-tain systems of Meru, Kilimanjaro and Kenya are quite varied and host manyendemic species, (3) there are many different habitats.Last but not least, Arusha National Park is regularly visited by many touristsinterested not only in large mammals, but also in smaller animals such asbirds or butterflies. The opportunity to watch some brightly coloured endem-ic chameleons (such as Chamaeleo jacksonii merumontanus) or listen to the

Lokie swamp after heavy rainfalls, hundreds of Xenopus muelleri can be found in a single pit-fall trap.

14

melodic calls of beautiful frogs (like the endemic Hyperolius viridiflavusommatostictus or the mountain frog Strongylopus fasciatus merumontanus)could add value to the Park and help people to appreciate a different aspectof this beautiful protected area.

Data collection

This guidebook includes the results of a field campaign carried out in April-May 2001. Some scattered data were also collected by one of the authorsduring the ichthyological and limnological survey of the park in October-November 2000.The methods used to collect data on amphibians and reptiles agree with thestandard ones proposed by Heyer et al. (1994), Blomberg & Shine (1996)and Halliday (1996). Two trained persons were active for at least 6 hours aday (day time and night time) for 17 days (April-May 2001) always assistedby three more biologists.Photographs were taken of all taxa to document their natural coloration andpattern variation. As a further aid to taxonomic identification the acousticrepertoire of some amphibians was recorded with a Marantz professionaltape recorder. Voucher specimens were deposited at the University of Dar esSalaam to confirm identification but this was, if possible, limited to specimensoccasionally killed by ants or drowned in the traps.Two main survey techniques were adopted: (1) Drift fences & pitfall traps andthe (2) Systematic Sampling Surveys (time-constrained). Both techniqueswere applied in all the major natural habitats available in Arusha NationalPark.Drift fences & pitfall traps. Drift fences intercept amphibians and reptilesmoving on the ground and redirect them into pitfall traps. Four drift fenceswere located in different habitat types. Each fence was made from a 60 cmwide plastic sheeting placed in a 10 cm trench, backfilled with soil and fas-tened every three meters to a staple. The pitfall traps were made from largeplastic buckets (diameter 30 cm, high 40 cm) buried in the ground, with theopening flush with the surface. Fifty meters of fencing with 10 traps wereplaced near to possible amphibian breeding sites (swamps, ponds, streams)and 75 meters of fencing with 10 traps in suitable reptile habitats. The trapswere checked every day in the morning for seven days and then moved toanother location. Pitfall traps are extremely useful to obtain information about ground dwellingamphibians and reptiles, but some species are captured more easily thanothers: amphibians that are strong jumpers or climbers (like Ptychadena orHyperolius) or large reptiles (large snakes) are more difficult to trap. Systematic Sampling Surveys (time-constrained). This is an opportunisticsearch for amphibians and reptiles with the goal of finding as many speciesas possible. Before each search, the exact locality, latitude and longitude,

15

date, number of observers, weather conditions, temperature, habitat type,vegetation, slope and starting time were recorded. When a habitat had beenadequately sampled in the judgement of the investigator (i.e. when the wholearea had been thoroughly investigated or when no new species had beenlocated within a given period of time), the finishing time was recorded and theobservers moved to another location. This technique is very useful, making itpossible to obtain quantitative values as percentage composition of speciesand numbers seen per man-hour of searching.Secretive species were sought in their refuges (e.g., under stones, tree barksor fallen logs, in leaf litter or among the branches of trees). Night searcheswere carried out with the aid of head-lamps and flashlights. The calls ofamphibians at breeding sites were used to detect different species (some-times they can be heard up to 2 km away) and traced to their source when a“different” call was heard. Specific searching techniques were applied to findsome taxa (Caecilians, Chameleons). Different kinds of stake nets were usedto catch adult amphibians and tadpoles; fishing rods with slip knots wereused to noose lizards, agamas and skinks; thick leather gloves and boots,hooks, tongs and “T” shaped sticks helped to catch snakes.

Lake Longil during the wet season with Kilimanjaro on the background.

Results

During the herpetological survey of the Arusha National Park 10 species ofamphibians and 24 of reptiles were found. Analysis of the data collectedshows that the survey allowed us to do a complete (or almost complete)check list of the amphibians, but the accumulation graphs for the reptiles indi-cate that a few species are still lacking and more research is needed to com-plete the list. This is due to the limited time of the survey and also becausethe rainy season is optimal for the amphibian census, but is also the worstperiod to look for reptiles due to cold weather and high grasses. In particularmost of the large snakes were probably hibernating. We were unable toobserve any large pythons, for example, during the survey, whereas inOctober and November many specimens had been found.

16

LIST OF SPECIES

18

The species accounts are based on the following references except wherenoted:Common names for Reptiles are taken from Broadley & Howell (1991),Loveridge (1957) and Branch (1994); for Amphibians from Passmore &Carruthers (1995), Lambiris (1989b) and, for the species not listed, fromFrank & Ramus (1996). Systematics and Nomenclature are based on Frost, 1985, 2000, Duellman,1993, Meirte, 1992 and Uetz, 2001 except where noted.Relevant data about identification, geographic range, ecology and repro-duction were taken from (Amphibians): Andersson, 1911; Bowker &Bowker, 1979; Channing & Griffin, 1993; Duellman & Trueb, 1994; Frost,2000; Lambiris, 1989a, 1989b; Loveridge, 1925, 1953; Passmore &Carruthers, 1995; Passmore et al., 1995; Poynton, 1964; Poynton &Broadley, 1985a, 1985b, 1987, 1988, 1991; Rödel, 2000; Schiøtz, 1999;Stewart, 1967. (Amphibians and Reptiles): Barbour & Loveridge, 1928b;Bauer et al., 1993; Lambert, 1985, 1987; Laurent, 1964; Largen, 1997;Loveridge, 1935; 1957; Rose, 1962. (Reptiles): Broadley, 1990; Chippaux,1999, FitzSimons, 1943; Lönnberg, 1911, Loveridge, 1936, 1959; MacKay &MacKay, 1985; Marais, 1992; Neças, 1999, Pitman, 1974; Schleich et al.,1996; Uetz, 2001; Vesey-Fitzgerald, 1975.

Notes: Due to graphic necessities the order in which the species are pre-sented have been slightly modified but a complete systematic check-list hasbeen added at the end of specie accounts. The synonymies are limited to those names that can be found in field guideson African amphibians and reptiles published in recent years.

Bufo gutturalisPower, 1927

Common namesGuttural Toad, Greater Cross-markedToad

Synonyms Bufo regularis gutturalis Power, 1927

IdentificationBufo gutturalis, as it is common in thespecies of the same genus, is stout withshort strong limbs and reduced webbingon the feet. The skin is rough and warty,granular below; there are two largeprominent paratoid glands just behindthe eyes. The top of the snout is typical-ly marked by four dark patches with alight cross between them. The groundcolour is usually brown with symmetri-cally arranged irregular dark blotchesand often a light vertebral stripe. Someindividuals show a reddish tinge in theback of the legs. This species can growup to 98 mm of length but the biggestanimal we found in Arusha National Parkwas just 57.8 mm.

Geographic RangeEastern and southern Africa: from Kenyasouthward to South Africa includingBotswana, northern Namibia and east-ern Angola.

Local distributionThe guttural toad is apparently confinedto the lowlands and we found it up to thePark Rest House (less than 1700 m). Itis quite common in the bushlandbetween Momela Gate and the shore ofthe Ngare Nanyuki river but can befound also in the bushland Uwanja waMomela and between Big and SmallMomela lakes.

Ecology and general behaviourThis species lives in open country bush-lands and grasslands often quite farfrom wet areas and it is not unusual tofind it on roads, in gardens and near tohuman habitations. The diet is wide,they will eat almost any animal of a suit-able size. The call is a deep vibrantcroak.

ReproductionBreeding usually takes place in perma-nent shallow waters; the eggs are char-acteristically united in paired strings andare laid among submerged vegetation.During our survey in April and May weheard the call of a few males only onenight in the Serengeti Ndogo. We neverobserved toads in the water and wenever caught any Bufo gutturalis in thepitfall traps that we put close to thewater. All the guttural toads we found inthe pitfall traps were caught in openbushlands and over 75% of them werejuveniles (less than 50 mm of length).

19

Xenopus muelleri(Peters, 1844)

Common namesNorthern Platanna, Mueller’s Clawed Frog

Synonyms Dactylethra mülleri Peters, 1844

TaxonomyWhile Xenopus muelleri has a widerange of distribution in Tanzania andKenya, there is confusion in geographicdistribution with Xenopus laevis. Thespecies is monotypic.

IdentificationThe head is small with upwardly directedeyes, the pupil is circular and there is ashort tentacle under each eye; tympa-num and tongue are lacking. The body isflattened and there are sensory laterallines organs on the sides made by manytubercles; the skin is very slippery.Fingers lack webbing while toes are fullywebbed and the inner three terminate ina black claw. The back is usually darkbrown or grey with irregular dark patch-es, the belly is usually greyish white.Females can be distinguished by thelarger skin folds around the vent and areusually larger than males. In ArushaNational Park Xenopus muelleri canreach 82.5 mm of body length.

Geographic RangeAll southern Africa: from Burkina Faso toKenya and Uganda, southward to theRepublic of South Africa.

Local distributionXenopus muelleri is one of the mostcommon and widespread species inArusha National Park. from the opengrassland of Serengeti Ndogo up to thewetlands of Kilimanjaro view point andthe big pond near Njeku Camp (2519 m).It can be found both in temporary andpermanent waters even in some soda

lakes like Lekandiro and Small Momela.Using a beach seine we caught someplatannas even in the muddy waters ofEl Kekhotoito pond, a place that isorganically enriched by a large herd ofbuffalos and a few hippos. The highestdensity population is probably located inthe Lokie swamp where, using a driftfence, on a few occasions we caughtover 100 platannas in a single pitfalltrap. Many authors reported the pres-ence of Xenopus muelleri in streamsand rivers, but we never found any in thewatercourses of Arusha National Park.

Ecology and general behaviourPlatannas are usually restricted toaquatic habitats, they move on land dur-ing rainy nights. If the weather is wetenough they sometimes wander into theforest or bushland; we observed some ofthem over half a kilometre from the near-est wet zone. During the day they areusually difficult to spot, but in pools withpoor oxygen it is possible to detect theirpresence by circles in the water whenindividuals come to the surface to takeair. In the night with a lamp it is possibleto observe them as they float motionlessin the shallow water. Xenopus can feedboth in the water and land; a wide rangeof arthropods are preyed on but alsosmall fish and even small tadpoles. Thecall is a soft buzzing sound utteredunder water by both sexes.

20

ReproductionThe mating begins immediately after thestart of the rainy season and amplexusoccurs under water. Several thousandsof eggs are laid on the aquatic vegeta-tion. The tadpoles are plankton feedersand swim with the head directed down-ward. The body is almost transparentwith a long tail and two sensory tenta-cles in the mouth region. They some-what resemble the glass catfishKryptopterus bicirrhis, a commonspecies of aquarium fish.

21

22

Ptychadena mascareniensis(Duméril and Bibron, 1841)

Common namesMascarene Grass Frog

TaxonomyDespite the fact that species of thegenus Ptychadena are common andwidespread in most of Africa they areoften very difficult to identify.

IdentificationA “green frog” with six longitudinal ridgeson the back, and only the outer onesmay be interrupted. This species ismedium sized reaching a snout-ventlength of 51 mm (average size of adultsin Arusha National Park 25.5 - 30 mm).Fingers lack webbing and on the toeswebbing is present between the outermetatarsals. The back is usually brownor green with rounded green or brownblotches usually smaller than the size of

the eye. There is a light creamy vertebralband and a longitudinal light colouredline on the upper surface of the tibia.Males have paired gular slits on thesides of the throat.

Geographic RangeWidespread in most of Africa: fromSierra Leone to Egypt through Eritreaand Ethiopia to South Africa; alsoMadagascar and Seychelles Islands.Introduced into Mascarene Island.

Male of Mascarene Grass Frog from Kilimanjaro View Point; the opening of the vocal sac foldcan be spotted under the tympanum.

23

Local distributionWidespread and abundant in manyareas of the park, the Mascarene grassfrog is the most common amphibianaround the brackish waters of theMomela lakes (Big and Small Momela,Lekandiro). Walking on the banksamong the reeds it is possible to see ahundred frogs leaping away in the waterin less than ten minutes. This speciesalso inhabits most of the ephemeralponds in the grasslands, for example, inSerengeti Ndogo and the small pondbetween Lekandiro and Tulusia lake.Some specimens were found on theshore of the fast flowing stream NgareNanyuki. In Arusha National ParkP. mascareniensis can be found as highas Kilimanjaro view point and the ArchedFig tree (about 1900 m).

Ecology and general behaviourLives in grasslands, wooded grasslandsand forest not too far from water. This

species is extremely common in most ofthe wet areas as long as it can findrefuge among the vegetation. Accordingto Inger and Marx (1961) the diet con-sists mainly of terrestrial prey: beetles,grasshoppers, dragonflies, ants, butter-flies and small amphibians althoughaquatic invertebrates are preyed on aswell. The voice of the male can be heardboth during the day and the night, ashort low pitched nasal “quack” oftenassociated by a series of cluckingsounds. The males call from a con-cealed position in grass or just floatingon the surface with open legs.

ReproductionDuring the rainy season, small pigment-ed eggs are laid in a series of smallclumps among vegetation in shallowwater. We were not able to observe anyoviposition site but at the beginning ofMay we found a few females that lookedalmost ready to lay.

Female of Mascarene Grass Frog from Kilimanjaro View Point almost ready to lay the eggs.

24

Rana angolensisBocage, 1866

Common namesCommon River Frog, Angola River Frog

SynonymsRana fuscigula angolensis Bocage,1866

IdentificationA large “green frog” that can attain insome areas (Malawi) 90 mm of snout-vent length but usually no more than 70mm. Skin with incomplete longitudinalridges variable in development (cf.Ptychadena mascareniensis), long legs(length of the tibia is 55-72% of thesnout-vent length). Toes extensivelywebbed (cf. Strongylpus fasciatus), fin-gers not webbed. Ground colour on theback usually green or brown with blotch-es about the size of the eye, a lightgreen or yellow vertebral line usuallypresent.

Geographic RangeUpland areas from Ethiopia to Angola,eastward to Mozambique, includingmost of South Africa.

Local distributionThe common river frog in ArushaNational Park can be found both in

brackish and fresh water, at low altitude(Maksoro river springs, about 1400 m)and medium altitude (Kilimanjaro ViewPoint, arched fig tree wet area) up to theMaio falls (1926 m).

Ecology and general behaviourThe typical habitat of this species areslow flowing streams with permanentwater. In Arusha National Park most ofthe frogs can be found in forested areasthough many can also be observedamong the aquatic vegetation of theMaksoro river. Rana angolensis has twodistinct calls, a sharp rattle of about onesecond followed after a short pause by ashort “croak” that resemble the call ofthe European green frogs Rana synk.esculenta.

ReproductionBreeding may occur throughout theyear; several thousands of small pig-mented eggs are laid in shallow waterwith a very slow current. The tadpoles ofRana angolensis reach a length of 80mm at Gosner’s stage 40. We observedsome of them close to metamorphosis atKilimanjaro view point at the end of April.

Adult River Frog fromKilimanjaro View Point.

25

The wet area at Kilimanjaro View Point; in the area it is easy to spot: Xenopus muelleri, Ranaangolensis, Strongylopus fasciatus, Ptychadena mascareniensis, Phrynobatrachus kenien-sis, Kassina senegalensis and Hyperolius viridiflavus.

Only few amphibians can survive in the soda waters of the Small Momela lake: Xenopusmuelleri, Hemisus marmoratum and Ptychadena mascareniensis.

26

Strongylopus fasciatus merumontanus (Lönnberg, 1910)

Common namesStriped Stream Frog, Striped Long-toedFrog

Synonyms Strongylopus fasciatus (Smith, 1849)

TaxonomyThree subspecies are actually consid-ered valid (Poynton, 1964): the nominalform, S.f. fuelleborni and S.f. merumon-tanus. This latter subspecies wasdescribed by Einar Lönnberg from a sin-gle specimen collected on Mt. Meru at3000 meters during the first Swedishexpedition in 1905 (Lönnberg, 1910).

IdentificationSnout-vent length up to 50 mm (46 mmin S.f. merumontanus), very similar to ariver frog but with extremely long slenderlegs and toes. Webbing absent from fin-

gers and very reduced on the toes.There is a dark stripe on each leg fromthe knee to the ankle. The dorsal surfacelacks the skin ridges of Ptychadena. Theground colour is usually buff or goldenyellow with conspicuous dark longitudi-nal stripes. Some specimens of the Mt.Meru lack the dorsal stripes and have abrown-red back.

Geographic RangeStrongylopus fasciatus forms isolated

Strongylopus fasciatus from Kilimanjaro View Point, individual with striped pattern.

27

populations in the mountains from north-ern Tanzania to South Africa, westwardup to Zambia and eastern Zimbabwe.This scattered distribution is a clear relictof the cooler periods during thePleistocene when these populationswere linked together. Strongylopus fas-ciatus merumontanus is endemic in theuplands of northern Tanzania includingMt. Meru, Uluguru and Usumbara Mts.S.f. fuelleborni occurs in southernTanzania, eastern Zambia and Malawi.S.f. fasciatus is widespread in SouthAfrica and Zimbabwe.

Local distributionLimited to the upper meadows and openforested areas of Mt. Meru fromKilimanjaro view point upwards, and thewet areas near the arched Fig tree(about 1900 m) up to Njeku camp in thecaldera (over 2500 m) and Kitoto forest.The species probably occurs also inhigher zones since the type specimen ofS.f. merumontanus has been collectedat 3000 meters.

Ecology and general behaviourStream frogs are generally found nearopen grasslands within the forest, butduring the wet season they move intothe forest quite far from wet areas. Weobserved several young individualsStrongylopus along the road fromKilimanjaro view point to Kitoto forestview point. Taking photographs of thisspecies is quite difficult as they are fastmoving and can jump long distances.The call of Strongylopus fasciatus is aclear high-pitched “pip” uttered singly orin a short burst of three or four; it is quitedifficult to distinguish from the call ofHyperolius viridiflavus.

ReproductionThe eggs are laid singly among vegeta-tion in shallow waters. During April andMay on Meru we found many juvenilesof about 20 - 25 mm body length. Thereproduction peak probably occurs dur-ing the small rains of October toDecember.

Strongylopus fasciatus from the same locality, individual with plain reddish back.

28

Phrynobatrachus keniensisBarbour and Loveridge, 1928

Common namesPuddle Frog, Cricket Frog

TaxonomyThis species has been described byThomas Barbour and Arthur Loveridgein 1928(a) from a specimen collected in“a marsh on the northeast slope of Mt.Kenya, Kenya Colony”. The systematicsof puddle frogs is still quite confusedespecially in some African regions: “Aslong as we lack a thorough revision ofthis genus, these frogs cannot be deter-mined for certain” (Rödel, 2000).

IdentificationIn Arusha National Park Phrynobatrachuskeniensis is, along with Hyperolius nasu-tus, the smallest amphibian species; itmay attain a body length that ranges

from 14.6 to 26 mm (29 individualsexamined). The body is rather stockywith short limbs, the head is small andpointed. The pupil is horizontal and thetympanum quite small. The most impor-tant diagnostic feature of this genus is atubercle in the middle of the tarsus.Colour and markings are very variablewithin the Arusha National Park with atleast three different patterns. Ground

Adult Puddle Frog from Mbuga Za Raiden pond.

29

colour is usually brown, grey or beigewith a golden tinge. The back can beuniform, faintly mottled or marked withdark blotches; some individuals have ayellow vertebral line extending fromsnout to vent. Most of the frogs observedhad a dark lateral band on both sides ofthe head and on the flanks.

Geographic RangePhrynobatrachus keniensis is endemicof the upland meadows of Kenya(Kikuyu, Molo, Mt. Kinangop, Mt. Kenya)and Mt. Meru.

Local distributionCommon and widespread in all the wetareas of the Park especially in grass-lands but also in some forested areas(Kitoto) from Serengeti Ndogo (1400 m)up to Njeku Camp (over 2500 m). Thisspecies can be observed especially in

the small temporary ponds of SerengetiNdogo and near Lokie swamp. Wenever found any Phrynobatrachus nearsoda lakes and brackish streams.

Ecology and general behaviourPuddle frogs usually live on the banks ofswamp, pools and streams and they areready to seek refuge in the water whendisturbed. They usually move away fromthe wet areas only after rainfall, but inthe meadows inside the caldera of Meruthe average humidity is so high that it iscommon to find many individuals wan-dering around. The voice of the males isa quick series of ticks that resembles thesound of a coin falling on the ground.

ReproductionBreeding occurs in shallow standingwaters. The small eggs float in a singlesurface layer.

Phrynobatrachus keniensis from Kilimanjaru View Point.

30

Hemisus marmoratum(Peters, 1854)

Common names Mottled Shovel-nosed Frog

SynonymsEngystoma marmoratum Peters, 1854Kakophrynus sudanensis Steindachner,1863

TaxonomyThe systematic position of Hemisusmarmoratum has been tentativelyrevised by Laurent (1972) that recog-nised a few subspecies: H.m. marmora-tum, H.m. ingeri, H.m. loveridgei andH.m. sudanese.

IdentificationA small amphibian with short, fat body;the limbs are powerful and short, thehead is small with transverse fold andhas a pointed snout hardened for dig-ging. The eyes are small with a verticalpupil. In Arusha National Park femalesreach a snout vent length of 35.5 mmand males 29.5 mm (30 individualsmeasured). The dorsal colour is usuallybrown with a pattern of darker reticula-tion and yellow patches. The throat ofmales is usually grey.

Geographic RangeSub-Saharan Africa excluding rainforestfrom southern Somalia to northernSouth Africa.

Local distributionThe Mottled shovel-nosed frog is quitecommon in most grasslands and openwooded areas from 1400 to 1670 m (wefound it up to the big fig tree nearLeopard Hill View Point on Ngurdotocrater). Usually lives near watercourses(Maksoro, Ngare Nanyuki rivers) andmost of the ponds, swamps and evenbrackish water lakes (like Big and SmallMomela lakes).

Ecology and general behaviourThis species is rarely seen as it spendsmost of its time underground and can befound above the surface only during thenight or in wet weather. Unlike mostof the other burrowing amphibiansHemisus burrows headfirst using theforelimbs and pointed snout to penetratethe soil. The prehensile tongue ofHemisus marmoratum has a peculiarstructure that allows it to be protractedslowly (increasing capture success) andalso to be elongated hydrostatically todouble its length during feeding(Nishikawa et al., 1999). The diet con-sists mainly of ants and termites. Malescall from the mouth of a small burrownear water, the voice is a repetitive high-pitched buzz that can be confused withthe sound produced by crickets.

ReproductionEggs are laid in an underground cham-ber near water and the female remainswith the brood (Van Dijk, 1997). The tad-poles develop inside the chamber andthey react very quickly to the first rainsgoing into temporary ponds before anyother species of amphibians. In ArushaNational Park during April and May wefound some tadpoles that were on theedge of metamorphosis and we couldhear very few males calling, so thebreeding season probably occurs duringthe small rains period.

31

Hemisus marmoratum from Momela gate.

Shovel-nosed Frogs call from concealed positions on the banks of Lokie swamp.

32

Kassina senegalensis(Duméril and Bibron, 1841)

Common namesBubbling Kassina, Senegal Running Frog

Synonyms Cystignathus senegalensis Duméril andBibron, 1841

TaxonomyPossibly composed of various crypticspecies or at least a number sub-species; Schiøtz (1975) discussed thedifferences in eastern African materialand observed four different “forms”based mainly on dorsal pattern, butrejected the recognition of subspecies.Poynton & Broadley (1987) concludedthat: “...the material appears to provideno clear grounds for the separation oftaxa within the senegalensis complex”.Examination of K. senegalensis in theArusha National Park shows both speci-mens with Schiøtz’s “Form 1” (patternsenegalensis) and “Form 3” (patternargyreivittis).

IdentificationBubbling kassinas are medium sizedfrogs reaching a length of 44 mm inArusha National Park (34 individualsexamined) with short hind legs. Fingerslack webbing and do not bear terminaldiscs. The pupil is vertical. The back isusually bright yellow, khaki or darkbrown (darker individuals are more com-mon at Njeku camp) with a disruptivepattern of longitudinal dark bands thatcan be continuous or broken into streaksand oblong spots. Males have a gulardisc and a large dark subgular sac divid-ed into paired lateral pouches.

Geographic RangeAfrican savannas south of the Sahara,from Senegal and southern Mali toEritrea, Ethiopia and Somalia, south-ward to Namibia and South Africa(excluding the western Cape province).

Local distributionIn the Arusha National Park bubblingkassinas avoid the brackish waters ofsoda lakes but are quite common both ingrasslands and forested areas from thelowland temporary ponds in SerengetiNdogo (1414 m) up to the Njeku camppond (2519 m). Kassinas are good walk-ers and sometimes single individualscan be found quite far from wet areas.We found a few specimen on theNgurdoto crater rim and a subadult instony bushland about one kilometre fromthe nearest pond. Lokie swamp, the wetareas near Kilimanjaro View Point andthe rest house ponds are the best placesto observe this species in the Park.

Ecology and general behaviourEven though Kassina senegalensisbelongs to the Hyperoliidae family (thesame as the reed frogs) it is a slow mov-ing ground dwelling species that prefersto walk rather than jump. During the dryseason the species seeks refuge underlogs and stones. The voice is an unmis-takable “quoip!” that resembles the pop-ping sound of bubbles coming to the sur-face. Males usually call from submergedvegetation in shallow water during lateafternoon and night. Large choruses canbe heard over great distances; during thewet season for example the large aggre-gations of Kassinas calling from swampsand ponds inside Ngurdoto crater can bedistinctively heard from the rim.

33

ReproductionIn Arusha National Park we observedsmall clumps of eggs on submergedgrass during April and May. The tadpoles

are often brightly coloured and growquite big (usually about 50 mm); theyhave broad fins and a pointed tail.

Bubbling Kassina from Kilimanjaro View Point.

Cluster of eggs laid by a single kassina at Lokie swamp.

Hyperolius viridiflavus ommatostictusLaurent, 1951

Common namesPainted Reed Frog

TaxonomyThe taxonomy of the Hyperolius viridi-flavus group is extremely complex with28 subspecies recognized by Schiøtz,(1999). Wieczorek et al. (2001) split H.viridiflavus into 10 species and accord-ing to this paper the subspeciesof Arusha National Park should beincluded, along with seven more taxain Hyperolius glandicolor (Peters,1878). Most of the books publishedbefore Schiøtz (1999) considered theommatostictus subspecies as membersof the large group of Hyperolius mar-moratus. Christina M. Richards (1981)discussed the pattern variation of differ-ent subspecies of Hyperolius viridi-flavus including H.v. ommatostictus.

IdentificationA medium sized treefrog with a snout-vent length up to 30 mm, the shaperesembles somewhat the Europeantreefrog Hyla arborea or the Americanbarking treefrog Hyla gratiosa. Fingersand toes bear terminal adhesive discsand are webbed. The snout is truncate.

As in all the species of the genusHyperolius, the pupil is horizontal and thetympanum concealed. The colour patternof H.v. ommatostictus is extremely vari-able usually the dorsum is dark brownwith small white rings or white spots thatcan be completely absent in some indi-viduals. During the day frogs seen rest-ing on the vegetation can be almostwhite. A few adult males and most juve-niles are beige or brown with undulatingdorsolateral stripes. The limbs are oftenred especially on the underside. Malespresent a large vocal sac on the throatprotected by a gular disc.

Geographic RangeThe distribution range of Hyperoliusviridiflavus complex includes most of thetropical Africa, while the subspecies ofArusha National Park Hyperolius viridi-flavus ommatostictus is an endemic ofthe Kilimanjaro and Meru areas.

Local distributionThe painted reed frog is widespreadnear the freshwater ponds of ArushaNational Park; it is quite easy to seebecause it reaches high population den-sities. In Arusha National Park it can befound in all wet areas from 1400 to 2400m (the highest point we found thespecies is the Giraffe pond between theKitoto view point and Miriakamba huts).It is not present near the Momela lakesas it usually avoids brackish waters.

34

Spotted individual from the rest house pond.

Ecology and general behaviourThis species can be found nearswamps, ponds, or slow flowing streamsin different habitats both in savannas(like Serengeti Ndogo) or in small openareas in the forest (Kilimanjaro viewpoint). The males call from reeds andsedges at the edge of ponds but alsofrom bush and trees that they are able toclimb up easily. The males spend a lot ofenergy during reproduction (Grafe,1996) and despite small body sizeare able produce very loud calls: ashort “weep!”, resembling that ofStrongylopus. Loud choruses can beheard during the night throughout theyear, but single frogs call often duringthe day especially in the dry season.

ReproductionThis species lays small clusters of eggs(up to 12 per season according to Grafe)on submerged water plants.Tadpoles have a long tail with a pointedtip. Grafe and Linsenmair (1989) report

that some females of H.v. ommatostictusare able to change into males. This isthe only known case of sex changeoccurring among amphibians.

35

Calling male from rest house pond.

Brown individual from Mbuga Za Raiden pond.

Hyperolius nasutusGünther, 1864

Common namesLong Reed Frog

TaxonomyThe systematics of the Hyperolis nasu-tus group is still not clear and manysubspecies of uncertain systematicvalue have been described. Poyntonand Broadley (1987) proposed thename Hyperolius benguellensis for thesouthern Mozambique and Natal popu-lations. Channing (in press) states thatthe South African populations shouldbe separated in a different species:Hyperolius poweri.

IdentificationA very small sharp-nosed reed frog withgreen translucent elongated body andwhite belly; the dorsum could be com-pletely uniform or finely stippled withdark spots occasionally forming a mid-dorsal stripe. A bright white dorsolateraloften black bordered band is frequentlypresent in males and sometimes also infemales. The size of breeding males isusually 19-22 mm, the maximum snout-vent length is 24 mm. Males have a yel-low or white gular disc.

Geographic RangeThis species is common in most of thesavanna areas south of the Saharaboth in west Africa (from Ivory Coast toCameroon) and in east Africa (fromEthiopia to South Africa).

Local distributionIn Arusha National Park Hyperoliusnasutus is localized at lower altitudes(1400-1600 m) and not common; it canbe found around the ponds and swampsin the open areas of the south easternpart of the Park like the pond Mbuga ZaRaiden, the edges of the Lokie swamp(near lake Longil) and in a few shallowpans inside Ngurdoto crater.

Ecology and general behaviourThe long reed frog lives in grasslandsand open wooded grasslands. Usually itcan be found, in the wet season only,well concealed on grass stems in thedense vegetation bordering ponds andwetlands. During the afternoon and theevening the males call from sedges andreeds above the water, a harsh highpitched chirp about 0.3 seconds long.

ReproductionThe small egg masses of about 20 eggsare laid on submerged vegetation justbelow the surface, usually the clutchsize ranges from 60 to 292. Tadpoleshatch in five days, are light brown withdark spots and resemble those ofPhrynobatrachus. Larger tadpoles usu-ally have a dark-tipped tail.

36

37

Geochelone pardalis babcocki(Loveridge, 1935)

Common namesTropical Leopard Tortoise

SynonymsTestudo pardalis babcocki Loveridge,1935

TaxonomyThe species is probably monotypic.

IdentificationA large tortoise that can grow up to 35-40 cm and exceptionally 70 cm of totallength. Neck hidden when the head iswithdrawn into the shell. Carapace with-out hinge, convex and humped (not flat-tened). Shell colour yellow or lightbrown, speckled with black. Leopard tor-toises from the Arusha area are usuallysparsely coloured while the ones fromSerengeti are buff with radiating spots;this could be related to the drier habitatin the Arusha area (Kabigumila, 2000).

Geographic RangeGeochelone pardalis is distributedthroughout the savannas of Africa

from southern Sudan, Kenya, Uganda,Tanzania, Rwanda, Burundi toSwaziland, including Zaire and Angola.

Local distributionDuring our survey this species was neverobserved inside the Park but large tor-toises have been found in different areasby park rangers and by some biologistsof Oikos Institute (usually betweenMomela gate and Serengeti Ndogo). Theleopard tortoise is probably the mostendangered reptile in Arusha NationalPark since it is a conspicuous, slow mov-ing, long living species very vulnerableto collecting. Two specimens have beencollected in Arusha N. P. by YngveSjöstedt in 1905 (Lönnberg, 1910).

Ecology and ReproductionThe growth rate is higher in immature ani-mals (6.9 mm per month) than in adults(2.9 mm per month) in northern Tanzania(Kabigumila, 2000). Sexual maturity isusually reached at 15 years. The femalesare usually much bigger than the males(usually 1.7 times) but are also lessnumerous; this could be because annualmortality is higher in females (Hailey &Coulson 1999). During the breeding sea-son males engage in combat. Femaleslay clutches of 6-15 eggs in a small holein the ground. Food consists of a varietyof plants and grasses.

ProtectionIncluded in CITES appendix II.

Individual from Tarangire National Park.

Hemidactylus mabouia(Moreau de Jonnès, 1818)

Common namesTropical House Gecko

TaxonomyLoveridge, 1947

IdentificationA medium sized gecko with a flattenedhead which is longer than it is broad anda little broader than the neck. Toesdilated with paired adhesive lamellaebelow with a free distal digital joint risingfrom the end of digital expansion(cf. Pachydactylus). Thumb clawed;enlarged tubercles on tail and body (12-18 rows). Distance from anterior borderof the eye to the tip of the snout longerthan the distance from posterior borderof the ear opening to posterior border ofthe eye, 7-10 transverse dorsal scalerows in a caudal verticil. 22-40 pre-anofemoral pores in males. Pupil verti-cal. Colour very variable as in most ofthe geckos, usually brown or grey(sometimes almost white) with scattereddark spot, often four to five dark trans-versal bands on the body and 10-12bars on the tail.

Geographic RangeThis is a widely distributed reptile: mostof sub-Saharan Africa from Senegal

to Ethiopia down to South Africa.Elsewhere also on Madagascar,Seychelles, Antilles, Comoro and otherislands, Mexico, Panama, Trinidad,Puerto Rico, Colombia, Bolivia, Brazil,Guyana, French Guyana, Suriname andArgentina. The species has been intro-duced into Florida and Honduras and isstill expanding its distribution (Meshaka,2000).

Local distributionThis species has been found exclusivelyon the wall of the rest house where welived and on the building nearby but fewattempts has been made to check itspresence on other buildings inside thePark. Medium search time 0.084 geckosper hour.

Ecology and ReproductionThis species is usually found on thewalls and roofs of buildings but can befound also on cracked rocks, in the hol-lows of trees, on baobabs and the crownof palms. As with most geckos, thisspecies is able to emit sounds whilecommunicating with conspecifics: a soft“tik-tik-tik” repeated 7 or 8 times. Thebreeding period has its peak fromSeptember to January (Moodley &Biseswar, 1997); sometimes the femaleslay their eggs in a communal depositoryof 50-60 eggs. Loveridge reports H.mabouia seizing a small white-headeddwarf gecko (Lygodactylus picturatus).

38

Pachydactylus turneri (Gray, 1864)

Common namesBibron’s Thick-toed Gecko

TaxonomyThe origin and nomenclatural position ofgeckos of the Pachydactylus group(especially the endemic forms) is actual-ly under study (Bauer et al., 1997,Bauer, 1999). Gerald Benyr (1995) sep-arated P. laevigatus from P. bibronii andplaced turneri and pulitzerae as sub-species of it. However, the name turneriGray 1864 has priority over laevigatusFischer 1888, so Pachydactylus turneriis the correct name for the species.

IdentificationA stocky gecko with large keeled tuber-cles on the neck, back and limbs. Toesdilated throughout with a single row ofadhesive lamellae, distal joint not com-pressed (cf. Hemidactylus) and thumbclawless or with tiny claws. Pupil verti-cal; lower eyelids vestigial or absent.The back has minute granules and largetubercles. Rostral not bordering nostril,preanal pores in males absent. Theback is usually grey or brown with 4-5curved dark bands on the back and 8-10on the tail.

Geographic RangeFrom South Africa to Rwanda, Botswanaand Tanzania including Angola,Mozambique and Swaziland.

Local distributionUsually found in rocky areas especiallyon lake shores (Big Momela andTulusia) but also under stones in thegrasslands near Kusare post. Number ofanimals per hour of search: 0.320.

Ecology and ReproductionThis species is gregarious and many indi-viduals can be found under the samestone. P. bibronii usually feeds in theevening and early morning, eating mainlyants, termites, grasshoppers, beetles andflies. The female usually lays two eggs(16 x 14 mm) hidden in a rock crack.

39

40

Keys for the identification of the chameleonsof Arusha National Park(Simplified from Broadley & Howell, 1991)

1a. A single series of enlarged granules forms a gular crest on the median line of thethroat, often extending along the belly as a ventral crest; a white line from chinto vent ................................................................................................................. 2

1b. Gular crest absent ............................................................................................... 3

2a. Body scalation homogeneous ............................................................................. 4

2b. Body scalation heterogeneous, scattered largetubercles present ........................................................................ Chamaeleo rudis

3a. Body scales homogeneous, or at most a few slightly enlargedtubercular scales .............................................................. Bradypodion tavetanum

3b. Body scales heterogeneous, granular scales interspersedwith large tubercles ............................................................... Chamaeleo jacksonii

4a. Occipital lobes merely indicated, not moveable ...................... Chamaeleo gracilis

4b. Occipital lobes small to large, mobile, in contact on the median lineor narrowly separated ............................................................... Chamaeleo dilepis

Bradypodion tavetanum from Seneto Post.

Bradypodion tavetanum(Steindachner, 1891)

Common namesKilimanjaro Two-horned Chameleon

Synonyms Chamaeleo tavetanus Steindachner,1891

TaxonomyThe genus Bradypodion is endemic toSouth Africa and the East African speciesshould be returned to Chamaeleo for themoment. Several people are trying tosort out the problematic chameleon phy-logeny (Broadley, pers. comm.).

IdentificationCasque raised on the median line of thehead, males (and some females) havepaired rigid scaled horns extending for-ward from preorbital region. Canthalcrest which is not shovel shaped on thesnout. Body scalation heterogeneous,small granules with scattered largetubercles, gular crest absent.

Geographic RangeKenya (Teita Hills), Tanzania (Arushaarea, Kilimanjaro, south to North PareMountains).

Local distributionObserved during our survey on the treesaround Seneto Post. Loveridge (1959)reports about a series of specimen col-lected on the Meru for C.J.P. Ionides byCol. J. Minnery.


41

Individual from Seneto Post.

42

Individual from Micumi (Tanzania).

gular crest on the median line of thethroat and on the belly.

Geographic RangeMost of the savannas of tropical Africa.

Local distributionThis species has never been observedin the park but it is listed here as it iscommon in different areas in northTanzania and could be present as well.

Ecology and ReproductionFound in savannah and bushland, usu-ally feeds on grasshoppers and beetles.Breeding time is usually in the earlyrains. Egg development takes 3-4months and finally in the dry season thefemale lays 25-50 small eggs that willhatch in approximately 150 days.


Chamaeleo dilepisLeach, 1819

Common namesCommon Flap-necked Chameleon

Synonyms Chamaeleo petersii var. kirkii Gray, 1865

TaxonomyThe subspecific status of Chamaeleodilepis is controversial with five doubtfulsubspecies that urgently need a revi-sion: dilepis, idjwiensis, isabellinus,martensi and petersii.

IdentificationA large species (20-24 cm) with occipitallobes small to large, mobile, in contacton the median line or narrowly separat-ed, body scalation homogeneous. A sin-gle series of enlarged granules forms a

43

neous, a single series of enlarged gran-ules forms a gular crest on the medianline of the throat.

Geographic RangeFound in most of equatorial Africa.

Local distributionReported for Mt. Meru by Loveridge(1957).

Ecology and ReproductionThis widespread species can be found indifferent habitats including wet and dryforest, forest borders and bushlands buthumid areas are usually avoided. Aftermating, the female digs a small hole inthe ground and lays 20-30 eggs that willhatch after 240-300 days.


Chamaeleo gracilis Hallowell, 1842

Common namesGracile Chameleon

SynonymsChamaeleo granulosus Hallowell, 1856Chamaeleo burchelli Hallowell, 1856Chamaeleo simoni Boettger, 1885

TaxonomyThis species is monotypic sinceChamaeleo gracilis etiennei has beenelevated to species rank.

IdentificationA large chameleon (up to 40 cm) thatgreatly resembles Chamaeleo dilepisbut can be distinguished by the occipitallobes that are merely indicated and notmoveable. Body scalation is homoge-

44

Geographic RangeMountain areas in south westernUganda, eastern Zaire, Rwanda andBurundi. The relict populations on Mt.Kilimanjaro and Meru belong to theChamaeleo rudis sternfeldi subspecies.

Local distributionRecorded on Mt. Meru at “Laikinae”(7500 ft. alt.) by C.J.P. Ionides on August1957 and by B. Cooper on the easternslope at 9000 ft. (Rand, 1963).

Ecology and ReproductionA mountain species usually found over2500 m.


Chamaeleo rudisBoulenger, 1906

Common namesRuwenzori Side-striped Chameleon

Synonyms Chamaeleo rudis sternfeldi Rand, 1963

TaxonomyKlaver and Böhme (1997) list only onesubspecies Chamaeleo rudis sternfeldi,but they mention also that this taxa couldbe considered a full species.

IdentificationA small chameleon with snout-ventlength 62-85 mm. Body stocky, headshort and broad. No cranial horns, nosail-like dorsal crest, nostril laterallydirected, body squat with scalation het-erogeneous: large scattered tubercleson the flanks.

45

Chameleons can be observed mainly on the trees at the edge of the forest.

46

Adult male from unknown area on Mount Meru.

the subgenus Trioceros. Finally it mustbe stressed that the genus Chamaeleois masculine, thus the correct spelling ofRand’s subspecies is merumontanus.

IdentificationA small chameleon (up to 16 cm snoutvent length), with a low casque and adorsal crest formed by 17-20 large coni-cal twin scales. The males have threeforward oriented horns, the central onestarts from above the mouth, the othertwo from the orbital crest; the femalesusually lack horns and if they are presentthey are smaller than in the males.According to Howell & Broadley thesecharacteristics allow the identification ofthe species: no gular crest, body, tail andlimbs without soft spines, no occipitalflaps, body scales heterogeneous, gran-ular scales and scattered large tubercles.

Chamaeleo jacksonii merumontanusRand, 1958

Common namesMeru Three-horned Chameleon

SynonymsChamaeleo jacksonii merumontanaRand, 1958

TaxonomyChamaeleo jacksonii has three sub-species: C.j. merumontanus small sizedand limited to Mount Meru, C.j. xan-tholophus the largest subspecies (up to35 cm) from the eastern slopes of MountKenya and the nominal species C.j.jacksonii (medium sized: up to 25 cm) inthe rest of the geographic range. Klaver& Böhme (1986) placed C. jacksoni in

47

Geographic RangeHigh mountain areas of Kenya, Ugandaand Northern Tanzania. The merumon-tanus subspecies is strictly endemic toMount Meru. There are naturalized pop-ulations on Hawaii islands.

Local distributionThe type locality indicated by Rand(1958) of merumontanus is the farmLaikinoi on Mt. Meru, 7500 ft. alt. (mis-spelled Laikinae in Rand, 1963) butnobody in Arusha National Park has anyknowledge of this location. C.J.P.Ionides in a letter dated 5th December1957 wrote about the specimen collect-ed by Lt. Col. J. Minnery “they are foundon bushes and in low small trees.Laikinoi is a farm on the very edge of therain forest”. As far as we know, therehave never been any buildings at 7500ft. alt. except Miriakamba Huts on the

eastern slopes, and other specimens ofC. jacksoni have been collected on theeastern slope at 9000 ft. alt. It seemslikely therefore that Miriakamba is a newname for the old Laikonoi. No MeruThree-horned Chameleon was foundduring the limited time of our research(little time was spent searching at alti-tudes above 2500 m).

Ecology and ReproductionC. jacksoni is a mountain species that isparticularly common up to 2800 m. It canbe found in humid mountain forestsbut also in coffee plantations, gardensand in the bushes around paths.Ovoviviparous gestation lasts for 6-7months and finally the female gives birthto 7-51 youngs.


Adult individual from Nairobi.

48

Agama agama(Linnæus, 1758)

Common namesRock Agama

Synonyms Lacerta agama Linnæus, 1758

TaxonomyThe genus Agama includes about 60species of the 317 species of Agamids ofthe world; the biogeographical affinities,dispersal models and biochemical phylo-genetics have been studied by Moody(1980 fide Jacobsen, 1997) and Joger(1991). Agama agama includes ninesubspecies, four of which are present inTanzania: A.a. usambare, A.a. elgonis,A.a. dodomae, A.a. ufipae. The sub-species of Arusha National Park is prob-ably the Elgon rock agama (Agamaagama elgonis Loveridge, 1923) that is

distributed from Mt. Elgon (Kenya) southto Usandawi in central Tanzania.

IdentificationTop of the head covered with small irreg-ularly arranged scales and with the inter-parietal larger than the adjacent scales.Eyelids completely moveable (the eyescan be closed). Dorsal scales uniform,keeled and imbricate except for a small

Adult male from Momela gate with regenerated tail.

49

vertebral crest limited to the neck.Agama agama elgonis can be distin-guished from the other subspecies bythe colour of the throat of the males:brick red with a black transverse mark atthe base.

Geographic RangeFrom Senegal to Ethiopia and south-ward to Angola, Democratic Republic ofthe Congo and Tanzania.

Local distributionIn the Park Agama agama is usuallyfound in open areas: at the sides ofroads, on concrete buildings (like theweather station on Ngurdoto crater) ornear to the water (Ngare Nanyuki river,Lekandiro and Tulusia lakes). The indi-viduals on Momela gate live togetherwith Mabuya striata and are quite tame;by slowly moving towards them, it is pos-

sible to get very close. Medium searchtime 0.708 agamas per hour.

Ecology and ReproductionThis species is usually much more com-mon in habitats that have been modifiedby man: piles of brush, bridges, build-ings, but also on solitary trees. The aga-mas like the sunlight and they can befound in exposed situations during thehottest hours of the day. Males show abrighter colouration especially on thehead and the neck while females aremuch duller. The breeding season is notconfined to a single period. Growth isfast especially during the first year, juve-niles can double their length in 12 month.Sexual maturity is usually reached dur-ing the second year when the snout-ventlength is about 80 mm (Daniel, 1961).Diet consists mainly of ants, termites,beetles and grasshoppers.

Female or young individual from Ngare Nanyuki river.

50

Mabuya striata(Peters, 1844)

Common namesCommon Striped Skink, Eastern StripedSkink

SynonymsTropidolepisma striatum Peters, 1844Euprepes punctatissimus Peters, 1854

TaxonomyDonald G. Broadley (2000) has recentlyreviewed the taxonomy of the genusMabuya in south-eastern Africa. Hehas given specific status to all the previ-ously recognised subspecies: Mabuyastriata punctatissima, M.s. wahlbergii,M.s. sparsa and has revived Mabuyamlanjensis from synonymy. According tothis paper the species should be consid-ered monotypical and the geographicrange restricted.

IdentificationA medium sized skink with body lengthup to 107 mm (males) and 113 mm(females). Eyelids movable, the lowerone has a large transparent disc. Dorsalscales with 3-7 keels, midbody scalesrow 32-43; limbs well developed (cf.Panaspis and Lygosoma). No white lat-eral stripe (cf. Mabuya varia). The backis usually red-brown with yellow dorso-lateral stripes; the belly is white.

Geographic RangeMost of eastern Africa, from Ethiopia toCongo, south to central and north east-ern South Africa. The species is alsopresent on the Comoro Islands.

Local distributionThe common skink is a savanna specieseven if its anthropophilous habits make iteasy to observe in Arusha National Parkbasking on stone walls, every kind ofconcrete building and on the sides ofroads. We found it as high as the ParkRest house (1686 m). One of the bestplace to observe this species is Momelagate, but it is quite common also on theshores of Big Momela and Tulusia lakes.Systematic sampling surveys: 0.792skinks per hour.

Ecology and ReproductionAlthough this species is consideredarboreal, it is also quite common onrocks. M. striata is viviparous and thereproduction can occur throughout theyear (Patterson, 1990). In ArushaNational Park many new-borns werefound at the end of April. The growth isfast and sexual maturity is reached in15-18 months. The diet consists of dif-ferent invertebrates including insects(termites, beetles) and snails. Lambert &Dewhurst (1998) observed a M. striatahunt down and eat a dwarf gecko(Lygodactylus luteopicturatus).

51

Mabuya varia(Peters, 1867)

Common namesVariable Skink

SynonymsEuprepes varius Peters, 1867Euprepes damaranus Peters, 1870

TaxonomyNo subspecies is actually consideredvalid. A recent taxonomical review(Broadley, 2000) has confirmed thatMabuya varia nykae do not represent avalid taxon.

IdentificationA medium sized skink with body lengthusually up to 70 mm and exceptionally117 mm (but individuals found ArushaNational Park usually around 50 mm).Eyelids movable, the lower one has alarge transparent disc. Dorsal scaleswith three keels, midbody scales row 30-36; limbs well developed (cf. Panaspisand Lygosoma). White lateral stripealways present (cf. Mabuya striata). Thecolouration can be very variable, theback is usually olive red-brown with orwithout vertebral and dorsolateral stripesand black blotches; the belly is white.

Geographic RangeSouth eastern Africa, from Eastern CapeProvince in South Africa north to Sudan,Ethiopia and Somalia. Westward up toCongo, Angola and Namibia.

Local distributionThe distribution data collected arescarce; most of the individuals havebeen observed in the bushland on thebanks of Ngare Nanyuki river (about twokilometres up from Momela gate) and inthe grasslands around Kusare post. Thedistribution and maximum altitude (1650m) here presented are probably veryunderestimated. Number of animalsobserved per hour 0.388.

Ecology and ReproductionMabuya varia is a terrestrial species thatcan be found around rocks, bushes andat the base of the trees in savannahsand montane grasslands. It preys main-ly on insects: beetles, crickets, caterpil-lars, termites and a few other inverte-brates. Reproduction usually occur dur-ing the winter, females give birth to 2-4up to 10 youngs.Head of Mabuya varia

modified fromFitzSimons (1943).

Adult from Ngare Nanyuki river.

52

Lygosoma afrum(Peters, 1854)

Common namesPeters’ Writhing-skink, MozambiqueWrithing Skink

SynonymsEumeces afer Peters, 1854Mochlus afer Bocage, 1867

TaxonomyThe genus Lygosoma has beenreviewed by Donald G. Broadley in1966a and1994; L. afrum has been dis-tinguished from Lygosoma (Riopa) sun-devalli by the different pattern of theback (speckled) and larger size. In EastAfrica four species has been distin-guished.

IdentificationMovable eyelids, the lower one scaly (cf.Panaspis). Dorsal scales smooth in 26-28 rows at midbody (cf. Mabuya), limbsshort. Dorsum light to dark brown,speckled with dark and white spots, ven-trum white. Total length 80-140 mm.

Geographic RangeFrom Somalia, Sudan and Ethiopiasouth to Mozambique (north of latitude24° S), west to northern Zambia andCongo. This species is also present onZanzibar.

Local distributionFound under stones only in open areasat low altitude (1400-1650 m); in grass-lands (around Kusare post), bushland(Uwanja wa Momela) and around lakes(Longil, Big Momela) and streams(Ngare Nanyuki). Search time: 0.118skinks per hour.

Ecology and ReproductionOviparous, lays 4-7 eggs in an under-ground chamber. The young measure 24mm (snout-vent length). Preyed itemsinclude caterpillars, grasshoppers, bee-tles, sandhoppers and tiny snails.Atractaspis bibroni and Lycophidioncapense have been reported to feed onL. afrum.

Adult from Uwanja wa Momela.

Head of Lygosomasundevalli modified from

FitzSimons (1943).

53

Panaspis wahlbergii(A. Smith, 1849)

Common namesWahlberg’s Snake-eyed Skink, SavannaSnake-eyed Skink

TaxonomyThe first complete review of this grouphas been done by the Rumanian her-petologist Iohan Fuhn (1969, 1970) whoassigned the European species to thegenus Ablepharus (with the exception ofone cosmopolitan species) and theAfrican species to the genus Panaspis.Subsequently Greer (1974), Perret(1975) and Broadley (1989) modified thenomenclature status of many African“ablepharine” skink, erecting new gen-era and assigning P. wahlbergii to thegenus Afroablepharus. Finally Jacobsen& Broadley (2000) recognized a newspecies: Panaspis maculicollis formerlyincluded in P. wahlbergii.

IdentificationA small dark skink with elongated cylin-drical body and limbs reduced but withfive digits. Eyelids fused and immove-able, the lower one with a large trans-parent disc which covers the eye (cf.Lygosoma and Mabuya). Interparietaldistinct and frontoparietal fused. Dorsalscales smooth, row 22-28 at midbody.Ground colour usually olive to darkbrown; the back can be uniform or may

present six longitudinal continuous orbroken dark lines. A white lateral line ispresent at least on the anterior part ofthe body. Breeding males are usuallypink to vermilion on the ventral parts ofthe body.

Geographic RangeFrom Democratic Republic of Congo toSomalia and Ethiopia; southward toZambia and Zimbabwe.

Local distributionIn Arusha National Park Panaspiswahlbergii is very common in all the low-land open areas and can be easilyobserved basking on stones especiallyalong the Ngare Nanyuki river, aroundLongil, Tulusia and big Momela lakes butalso at Asili post, Kinandia and the old“picnic” area. During our survey themedium search time for this species was0.742 skinks per hour.

Ecology and ReproductionThese skinks are diurnal and can befound in a variety of habitats includinggrassland and bushlands. They fre-quently take refuge under stones orinside grass tussocks. The diet consistsmainly of spiders, termites, homopteranand hemipteran bugs, beetles and ants.The females lay 2-6 small eggs understones or logs during November-January.

54

Adolfus jacksoni(Boulenger, 1899)

Common namesJackson’s Forest Lizard

SynonymsLacerta jacksoni Boulenger, 1899

TaxonomyAdolfus jacksoni kibonotensis, the sub-species of Kilimanjaro and Mt. Meru, isnot considered valid anymore.

IdentificationA brown-greenish lizard that resemblesthe European wall lizard Podarcismuralis. Frontoparietal present, dorsalscales small and granular, ventralssmooth, subdigital lamellae not keeled.Snout to vent length of five individualsmeasured in Arusha National Park up to51.5 mm.

Geographic RangeTanzania, Kenya, Uganda, Rwanda,Burundi and Democratic Republic ofCongo.

Local distributionMany Jackson’s forest lizards have beenfound under stones near a tree at Njekucamp (about 2500 m) and a single spec-imen in a pitfall trap on the slopes of

Ngurdoto crater near to Leopard hill(1670 m). Einar Lönnberg in 1911observed on Mt. Meru some Adolfus atthe “escarpment station” on the trunks ofbig cedar trees (this locality is probablyMiriakamba huts as the other station,Saddle huts, is well above the tree line).

Ecology and ReproductionThis species usually lives in trees at theedge of the forest where it feeds mainlyon beetles, moths and spiders. It is anarboreal species but can be found alsounder stones and debris. Loveridgereports of a female laying four eggsmeasuring 14 x 7 and 15 x 7 mm respec-tively.

Head of the same individual.

Adult male from the forest on southern sideof Ngurdoto crater.

Nucras boulengeriNeumann, 1900

Common namesBoulenger’s Scrub-lizard

SynonymsNucras kilosae Loveridge, 1922

TaxonomyBroadley and Howell (1991) examinedspecimens from the same locality ofdescription of Loveridge’s Nucras kilo-sae and could not find any diagnosticfeature, so they placed this species insynonymy with Nucras boulengeri.

IdentificationA lizard with rounded head and very longred tail; frontoparietal present, dorsalscales small and granular, ventralssmooth, subdigital lamellae not keeled,nostril bordered by 2 or 3 nasals andwell separated from the first labial (cf.Adolfus). Collar well marked, headshields smooth.

Geographic RangeUganda, Kenya and Tanzania, south tonorth-western Zambia.

Local distributionWe found a single juvenile specimen(SVL 26.5 mm) at the beginning of Mayin the open bushland sandy area along

the Ngare Nanyuki river (about 600meters west of Momela gate).

Ecology and ReproductionThe beautiful lizards of the genusNucras are usually found in open areaswith sandy soils; they are quite secretiveas they forage mainly in the early morn-ing and evening. If threatened by a pred-ator they are able to run away withnotable speed and agility. The mainbreeding period of this species is proba-bly the during the rainy season.

55

Young specimen frombushland aroundNgare Nanyuki river.

56

Leptotyphlops scutifrons merkeri(Werner, 1909)

Common namesMerker’s Worm-snake

SynonymsStenostoma scutifrons Peters, 1854Glauconia merkeri Werner,1909

TaxonomyThe worm snakes of south easternAfrica have been reviewed by Broadley& Watson (1976). Two subspecies wererecognised, the nominal form L.s. scu-tifrons (southern form, from South Africaup to central Tanzania) and L.s. merkeri(Kenya and Tanzania). Broadley (1990)and Webb et al. (2000) report thatLeptotyphlos conjunctus can be consid-ered a subspecies of L. scutifrons.

IdentificationSize and body shape resembling anearthworm except for the colour that isdark reddish brown to dark brown andthe shiny appearance. Blunt head withsnout prominent, no teeth on the upperjaw, the eyes are vestigial and coveredby scales. Ventral scalation similar to thedorsal one. Maximum size in ArushaNational Park 253 mm (body + tail).

Geographic RangeFrom South Africa northward up toKenya, Tanzania and Angola.

Local distributionUsually found by turning up stones ingrassland and bushland areas; five indi-viduals were observed along the banksof Ngare Nanyuki river. Other wormsnakes were found on the shores ofsmall and big Momela lakes and nearKusare post. Medium search time 0.152snakes per hour.

EcologyWorm snakes live underground and canbe observed only after heavy rains whenthey are flushed out or during the night.The females lay two or three eggs thatlook like a rice grain and the new-bornare just 55 mm long. Merker’s wormsnake feeds almost exclusively on antlarvae and pupae; they producepheromones that prevent them frombeing attacked by the ants (Webb et al.2000).

Individual from Lenganassa river.

57

The shore of the Big Momela lake is very poor in amphibian species but rich in reptiles:Lygosoma afrum, Mabuya striata, Pachydactylus turneri, Psammophis phillipsi.

Ngare Nanyuki river is a good habitat for amphibians like Hemisus marmoratum and Bufogutturalis but especially for reptiles: Panaspis wahlbergii, Mabuya varia, Agama agama,Psammophis phillipsi, Leptotyphlops scutifrons and Python natalensis.

58

Python natalensisA. Smith, 1840

Common namesSouthern African Python

Synonyms Python sebae natalensis A. Smith, 1840

TaxonomyPython sebae natalensis has been ele-vated to full species status by Broadley(1999).

IdentificationThis is, along with Python sebae, thelargest African snake with adults averag-ing 3-4 meters and exceptionally attain-ing 6 meters. The back is light brownwith black edged dark patches irregular-ly connected to form sinuous crossbars.On the sides there are irregular darkblotches, the underside is light greyspeckled with black spots and small darkpatches. Typical of the Boidae family arethe ventral scales narrower than body,

anterior supralabials scales with deeppits and vestiges of hind-limbs presentand visible externally. Python natalensiscan be distinguished from Python sebaeby the frontals broken up in 2-7 scales,no dark preocular patch and subocularpatch reduced to a dark oblique streak.

Geographic RangeFrom north eastern parts of South Africato central Kenya, in the west up to south-ern Angola including eastern Congo,Zambia and Burundi.

Local distributionThe Southern African python is fairlycommon in Arusha National Park andlarge snakes can be observed in manywet areas. The best places to meetpythons are the shores of the SmallMomela lake (they are usually hiddenamong the reeds or swimming in shallowwaters) but also walking on the banks ofthe Ngare Nanyuki river (starting fromthe Momela gate area) could be anexcellent way to observe them (forexample, the 3.70 meters long individualshown in the picture). The largest pythonwe saw was basking on the floating veg-etation of lake Longil at the beginning ofNovember. The best periods to observelarge snakes are during hot weather;during the rainy season, the weather isprobably too cold for this species and weobserved just a single juvenile along theSmall Momela lake.

Pythons can be easily observed along theNgare Nanyuki river as they are very fond ofwater.

59

EcologyPythons are non venomous snakes butdue to their large size and long recurvedteeth their bite can inflict deep wounds.The rock pythons are the only Africansnakes large enough to be potentiallyable to eat a man and even if attacks onman are probably mostly legend, a fewrecords in the past have been reported(Branch & Haacke, 1980). Pythons are very fond of water and usu-ally they do not live too far from it. Theyoften lay submerged with just the headon the surface waiting for prey like smallantelopes, wild pigs, monkeys, hares,cane-rats or sometimes fish. The femalelays 30-50 large eggs that she protectsby coiling around them. The newbornare 60 cm long and in the wild probablyneed 10-15 years to reach full maturity.The African python is protected in someCountries and has been included inCITES Appendix II.

Lower jaw of Python sebae: the longrecurved teeth of this snake can inflictnasty wounds.

The head of a Southern African Python from Small Momela lake.

60

Bitis arietans(Merrem, 1820)

Common namesPuff Adder

SynonymsEchidna arietans Wagler, 1828

TaxonomyTwo subspecies are actually consideredvalid: Bitis arietans arietans (Merrem,1820) is distributed in most of Africaincluding Tanzania and Bitis arietanssomalica Parker, 1949 in Somalia andNorthern Kenya.

IdentificationAn extremely stout, heavily built snakewith a broad flattened hornless triangu-lar head covered by small scales, snoutrounded, tail short. Eyes of moderatesize with vertical pupil. The scales arestrongly keeled. Colour yellow or brown

(pale, dark, orange or reddish) some-times greyish with a pattern of regularchevron-shaped bars on back and tail,venter yellowish or white. Total lengthusually up to 90 cm, exceptionally 150cm with a weight of 6 kg.

Geographic RangeOne of the most widespread Africansnake distributed from SouthernMorocco to Arabia and south to the

A Puff Adder perfectly camouflaged among the low grass of a garden.

61

Cape excluding the Sahara and the rainforest areas.

Local distributionThis species has been found in ArushaNational Park by Vesey-Fitzgerald(1975) and often seen by biologists ofOikos institute and Park rangers. Duringour survey, in the cold rainy season, wenever observed this species inside thepark. The best way to observe puffadders is to look for them on the roadsduring the night where they often restwhile hunting for prey or bask on thewarm soil. Puff adders are sluggish andcan be safely approached and pho-tographed from a few meters distance,but are also able to strike suddenly

extremely fast and they must be treatedwith extreme care and respect.

EcologyBitis arietans has long recurved fangs(12-18 mm in large individuals) at thefront of the upper jaw that are capable ofinjecting a large quantity of haematotox-in venom deep into the victim.Symptoms of bites include largeswelling, pain and necrosis. This speciesis responsible for most of the severesnake bites in Africa, even if only 5% ofthem prove fatal to man.The puff adder is a terrestrial snake andonly seldom climbs low bushes. The typ-ical movement is rectilinear and cater-pillar like but if annoyed it can movefaster sideways. It usually relies on itscryptic colour pattern to escape noticeand catch its prey. Diet consists mainlyof small terrestrial ground living mam-mals including rats and mice, but alsolizards, frogs and toads are often preyedupon. The species is ovoviparous withone litter per year. Litters of 20-40 arecommon but extremely large litters of147-156 neonates have been recorded.During cold periods the puff adder usu-ally hibernates or emerges for a fewhours to bask in the midday sun.

Mobbing of few superb starlings against alarge Puff Adder (same individual of thepicture on right).

Large Puff Adder found during the nightalong a road in Tarangire N.P.

The same individual of the opposite page;found about 20 km west of Mount Meru.

62

Elapsoidea loveridgei loveridgeiParker, 1949

Common namesLoveridge’s Garter Snake

Synonyms Elapsoidea sundevalli loveridgei Parker,1949

TaxonomyElapsoidea loveridgei was originallyincluded in the Elapsoidea sundevalliand elevated to specie rank by Broadley(1971). Recently a revision has beenpublished by Jakobsen (1997). Foursubspecies are actually recognized: E.l.colleti, E.l. multicincta, E.l. scalaris andthe nominal form E.l. loveridgei, this lat-ter is the subspecies of Arusha NationalPark and is limited to central Kenya andnorthern Tanzania.

IdentificationA stout, medium sized dark snake (totallength usually not exceeding 60 cm) with16-20 white or pink transverse lines (ornarrow bands); the body has a smooth“oily” appearance. As in all the membersof the Elapidae family this species hasone pair of enlarged, fixed, tubular poi-son fangs not enclosed in a membra-nous sheath, but this is not a useful char-acteristic unless you are handling theanimal. Internasal not bordering nostrils,dorsal scales in 13 rows at midbody.

Geographic RangeNorth eastern Congo, Burundi, Rwanda,Tanzania, Uganda and Kenya, Sudan,Ethiopia and Somalia.

Local distributionOnly one garter snake was caught dur-ing the survey; it was resting under alarge boulder on the shores of lakeTulusia.

EcologyA nocturnal and secretive species thatfeeds on geckos, skinks, lizards andsometimes amphibians. The area wherethe snake was found sheltered a largepopulation of Pachydactylus bibronii aswell as skinks and agamas, but noamphibians. The strange colour of theanimal is due to the fact that it wasalmost ready to shed. The venom is neu-rotoxic but the species is not aggressiveand very few human bites have beenrecorded.

Garter Snake from lake Tulusia.

Closer view of the same individual, theopaque eye is typical of the early sheddingprocess.

63

Naja haje(Linnæus, 1758)

Common namesEgyptian Cobra

SynonymsColuber haje Linnæus, 1758

TaxonomyThere are five races N.h. arabica, N.h.legionis and the nominal form N.h. haje.Naja haje annulifera has been elevatedto species rank and Naja haje anchietaeis now considered a subspecies of thistaxon (Broadley, 1995).

IdentificationA large snake (150-200 cm) with thickbody and stout head. Large dilatablehood on the neck. Young specimens areyellow-grey or brown to black largerones usually darker. Fangs as in all theelapid snakes, one preocular in contactwith nasal and separating prefrontalsfrom labial scales; supralabials not incontact with the orbit of the eye.

Geographic RangeSaudi Arabia, Oman and throughoutAfrica north of Angola, Zambia, and

southern Tanzania. The range of N.hajehaje is from Morocco to Egypt, south tocentral Tanzania and west to Senegal.

Local distributionThe presence of the species has beenconfirmed on the basis of a large (183cm) complete exuvia (shedded skin)found on the shores of Lekandiro lake.Local snake catchers affirm that alongthe rivers, near the eastern borders ofthe park, Naja nigricollis (another cobraspecies) is quite common but we per-sonally never found one in the park nordid Vesey-FitzGerald in the past.

EcologyA terrestrial species that often remains inthe same area for a long time. They arenormally not aggressive even if the poi-son is neurotoxic and extremely danger-ous. Despite the wide distribution of thisspecies the number of fatal bites is verylow. The Egyptian cobra feeds mainly onamphibians and other snakes. TheEgyptian cobra is not able to spit venombut the close species Naja nigricollis is.If annoyed they are able to stand up anddisplay a large hood. As defensivebehaviour Naja haje often shams death.

The Egyptian Cobra shedded skin foundnear lake Lekandiro.

Dendroaspis angusticeps(A. Smith, 1849)

Common namesGreen Mamba

SynonymsNaja angusticeps Smith, 1849Dendroaspis sjöstedti Lönnberg, 1910

TaxonomyEinar Lönnberg described Dendroaspissjöstedti from a specimen with aberrantscalation collected at Kibonoto; thespecies is not considered valid anymore.

IdentificationA large slender green snake (adultsaverage 180 cm, exceptionally up to 250cm), with coffin shaped head andsmooth scales. Fangs as all the elapids,3 preocular scales. Dorsal scales in 17-19 rows at midbody, inside of mouthwhite to bluish-white. This species is noteasily distinguishable in the wild fromPhilothamnus hoplogaster.

Geographic RangeKenya, Tanzania, Mozambique, Malawi,East Zimbabwe, Natal in South Africa.

Local distributionThis species was reported in Arusha

National Park by Vesey-FitzGerald, weobserved a green mamba among theacacia trees along the Ngare Nanyukiriver.

EcologyThis is an arboreal snake rarely seenoutside forests or dense bushlands; itfeeds on birds and eggs. Along with theblack mamba (that has never beenreported in the area) this is the mostfeared African snake due to its extremeagility, aggressiveness and the power ofits poison. Compared to the blackmamba the green mamba is lessaggressive and the venom is less toxic,but still capable of inflicting fatal enven-oming. Medical treatment is alwaysneeded for this species (Hodgson &Davidson, 1996).

64

Green Mamba from Arusha N.P.

65

Lamprophis fuliginosus(Boie, 1827)

Common namesBrown House-snake

Synonyms Lycodon fuliginosus Boie, 1827Boaedon fuliginosus Schmidt, 1923

TaxonomyFitzSimons (1962) recognized only onesubspecies L.f. mentalis (but consideredit questionable), but Broadley (1990) inthe revision of the same book concludedthat: “there is no justification for retainingmentalis as a subspecies of Lamprophisfuliginosus”.

IdentificationHead relatively flat with round snout,head slightly distinct from the neck, eyesrather small with vertical pupil. The backcolour in Arusha National Park is usuallyblackish grey (but in some areas thissnake can be brown). Typical of thisspecies are two light stripes on the sidesof the head and the high number (27-33)of midbody scales row. Total lengthbetween 60 and 120 cm.

Geographic RangeThroughout Africa but restricted to southwestern Morocco in North Africa.

Local distributionThe brown house-snake is consideredcommon in Arusha National Park byVesey-FitzGerald; we observed youngindividuals of this snake at Leopard hill(Ngurdoto Crater) and Uwanja waMomela (about 1 Km north of MomelaLodge).

EcologyThis harmless species, as the commonname suggests, is often found near vil-lages; in traditional communities it ismuch appreciated because it eatsrodents. Rodents are hunted inside theirburrows and killed by constriction. Dietincludes also lizards (incl. Heliobolusneumanni and Hemidactylus mabouia)and occasionally birds and bats.

All the Lamprophis that we found in ArushaN.P. were almost black.

Closer view the same individual fromUwanja wa Momela.

Lycophidion capense jacksoniBoulenger, 1893

Common namesJackson’s Wolf-snake, Jackson’s TigerSnake

SynonymsLycophidion jacksoni Boulenger, 1893Lycophidion irroratum Schmidt, 1923

TaxonomyLycophidion capense (A. Smith, 1831) isa polytypical species and the differentsubspecies show great variability in thecolouration patterns. The genus hasbeen reviewed by Laurent (1968) andlater by Broadley (1996) and threesubspecies are considered valid: L.c.capense, L.c. jacksoni, L.c. loveridgei;all these taxa, except the nominal form,occur in Tanzania.

IdentificationA small species measuring 35-40 cmwith short tail, head flattened and notvery distinct from the neck. L.c. jacksoniis usually dark grey or brownish, thedorsal scales are usually bordered withwhite at the apex and there is a paleband around the snout. No enlarged poi-son fang in the upper jaw, dorsal scales

smooth, nostril pierced in an entire nasalshield followed by a small postnasal,pupil vertically elliptic in strong light, dor-sal scales row reduced to 15 before thevent. In males 170-211 ventrals and 31-58 subcaudals; in females 178-221 ven-trals and 21-55 subcaudals.

Geographic RangeLycophidion capense is widely distrib-uted throughout Africa. The range of L.c.jacksoni includes southern Sudan,Ethiopia, north eastern Congo, Uganda,Western Kenya, Rwanda, Burundi andWestern Tanzania extending south-eastto Morogoro and the Uzungwa moun-tains.

Local distributionConsidered common in Arusha NationalPark by Vesey-FitzGerald. The speci-men in the picture was found on thewestern shores of Small Momela lake.

EcologyThe tiger snake is a nocturnal speciesthat feeds mainly on skinks: Panaspisand Mabuya. The prey is seized on theback of the neck and constricted.Lycophidion capense is a harmlesssnake, but in the field caution is neededas it can be confused with the stilettosnake Atractaspis bibronii.

66

Aggressive posture of Wolf-snake fromSmall Momela lake.

67

Psammophis phillipsii(Hallowell, 1844)

Common namesOlive Grass Snake

TaxonomyThe species included in thePsammophis sibilans complex (P. sibi-lans, P. phillipsii, P. subtaeniatus, P. bre-virostris, P. leightoni and P. rukwae) donot have a clear systematic position.The group has been reviewed byLoveridge (1940), Broadley (1966c,1977, 1990) and later by Brandstätter(1994, 1995 fide Hughes) criticallyreviewed by Hughes (1999). Accordingto Brandstätter the plain (unpatterned)specimen from Arusha should beascribed to Psammophis sibilans irregu-laris while Hughes does not justify theexistence of the subspecies. The sameunpatterned Tanzanian Psammophiswere regarded as P. phillipsii byBroadley & Howell (1991). Broadley(pers. comm.) probably will assign thespecimen from Arusha to Psammophismossambicus. Since the situation is stillconfused and a complete revision isurgently needed we decided to followBroadley & Howell (1991).

IdentificationA large robust brown long tailed snake(up to 190 cm) that can be not patterned

or with black edged scales forming thinblack longitudinal lines. 17 rows ofscales at midbody, 151-183 ventrals, 1preocular, 82-110 subcaudals, 8 upperlabials of which the fourth and fifth orfourth, fifth and sixth are in contact withthe eye, the first four infralabials in con-tact with anterior chin shield. The twospecimens examined from ArushaNational Park showed anal shield divid-ed and 10 lower labials.

Geographic RangeFrom Senegal to Kenya, south to north-ern Namibia, Botswana and part ofSouth Africa.

Local distributionObserved along the Ngare Nanyuki river(three individuals) and on the westernand southern shores of Big Momelalake.

EcologyThis snake is common in grassland andsavannas especially near water and it isoften encountered on the roads. It is notan arboreal species even though it mayclimb on bushes to bask. It moves veryquickly and when captured bites fiercely.The mild venom of P. phillipsii maycause pain and nausea that will passwithin 48 hours. The olive grass snakefeeds mainly on lizards, but also onmammals, frogs and small snakes(including venomous species such aspuff adder and black mamba).

Natriciteres olivacea(Peters, 1854)

Common namesOlive Marsh-snake

SynonymsCoronella olivacea Peters, 1854Natrix olivacea Cott, 1928

TaxonomyReviewed by Broadley (1966). This isactually a monotypical species, but inthe past Loveridge (1935) describedN.o. uluguruensis and considered validN.o. pembana that is actually regardedas closely related to the west Africanform N. variegata.

IdentificationA small harmless snake (up to 35-40 cm)with smooth scales. The body is grey,olive or brown with dark mid-dorsal band(4-5 scales row wide), ventral scalesusually orange or yellow. The pupil isrounded, anal shield divided, 19 scalesrow at midbody, 130-153 ventrals, 57-87subcaudals.

Geographic RangeSavannas and Forests from Sudansouth to Mozambique westward toGuinea and Angola.

Local distributionThis small snake was found often in thepitfall traps around wet areas includingMbuga Za Raiden pond and LokieSwamp. A specimen was found under astone at the Maksoro river springs andanother inside the Kambi ya fisi Forest(i.e. about 1 km from the nearest wetarea).

EcologyA diurnal savanna species that does notlive far from water. Swims well and feedsoften in the water on frogs and tadpoles,small fish and some invertebrates. Thissnake when first caught does not usual-ly attempt to bite.

68

Natriciteres is the most common snake inArusha N.P.

Detail of the head of a Olive Marsh-snakefrom Lokie swamp.

69

Crotaphopeltis hotamboeia(Laurenti, 1768)

Common namesHerald Snake, White-lipped Snake

SynonymsCoronella hotamboeia Laurenti,1768

TaxonomySince Broadley (1968) has elevatedCrotaphopeltis hotamboeia kageleri tospecies level (and moved it to a differentgenus Dipsadoboa shrevei) C. hotam-boeia is considered monotypic.

IdentificationA small snake (up to 60-75 cm, rarelymore) with short depressed roundedhead, broadened behind. The tail isshort. The body is dull grey or blackishgrey and unpatterned except for somescattered white dots. The head is usual-ly darker than the body and often irides-cent especially in young individuals, lipsare usually white as well as the belly.The identification can be based on thesecharacteristics: presence of poisonfangs in the upper jaw behind the eye,loreal shield present and excluded fromthe orbit by a single preocular scale,head quite large, pupil vertical, ventrals139-174, subcaudals 24-47, dorsalscales in 19 (rarely 21) rows at midbody.

Geographic RangeTropical Africa excluding a few areas inSouth Africa.

Local distributionRegarded as the commonest snake ofeast Africa (along with Lamprophis fulig-inosus) and common in Arusha NationalPark according to Vesey-FitzGerald. Wecollected many specimens around theamphibian breeding sites, including thepond at Mbuga Za Raiden, lake Longil,lake Tulusia up to Kilimanjaro view point(over 1900 m). The medium search timerecorded for this species during system-atic sampling surveys was 0.067 snakesper hour.

EcologyIt feeds at night on amphibians that areseized and held until the venom hasparalysed the prey and then swallowed.The poison is injected with a blade-likeback fang but is mild and has virtually noeffect on man. When first approached orcaptured this harmless terrestrial snakehas an aggressive reaction, it flattensthe body and the head to such a degreethat the white lips became evident on thedark/black head of this snake. It coilsand uncoils its body and attempts a fewfalse strikes. The origin of the strangecommon name “Herald snake” is due tothe fact that the presence of this speciesin South Africa was first published in theEastern Province Herald newspaper.

Young White–lipped Snake from Mbuga ZaRaiden.

70

Thelotornis mossambicanus(Bocage, 1895)

Common namesMozambique Twig Snake

TaxonomyBroadley (1979) reviewed the genusand recognized three subspecies ofThelotornis capensis: T.c. capensis (thesouthern race), T.c. oatesii (the westernrace) and T.c. mossambicanus in EastAfrica. In a recent paper by Broadley(2001) T. mossambicanus is recognisedas a good evolutionary species, which issympatric with T. capensis oatesii ineastern Zimbabwe. T. usambaricus isdescribed from coastal forests of theEast Usambaras, but it is also recordedfrom the West Usambaras, North Pare,Nguru and Uluguru Mts.

IdentificationA slender snake (up to about 140 cm)with long tail. The body is brown or greywith marbled and speckled with darkerblotches. The head is slender and dis-tinct from the neck, the top is uniformgreen and, typical of T. mossambicanus,the temporal region is always speckledwith black. Pupil horizontal “key”shaped, loreal shield present, a pair ofenlarged grooved poison fangs behindthe eye in the upper jaw.

Geographic RangeThelotornis mossambicanus rangesfrom southern Somalia south to centralMozambique, west to the shores of LakeTanganyika, Malawi and easternZimbabwe.

Local distributionVesey-FitzGerald reports that thespecies has never been recorded fromArusha National Park but we observed alarge individual among the acacia treesaround the Ngare Nanyuki river inNovember.

EcologyIt is considered an arboreal species butcatches a lot of its prey on the ground. Itis rarely seen in the field due to its cryp-tic colours and habit of remaining immo-bile for a long time. If disturbed can moveaway swiftly or inflate the neck. The dietconsists of small snakes and lizards(especially chameleons and arborealgeckos) but sometimes also few birds,frogs and toads are taken (Broadley,pers. comm.). The bite from thesesnakes is dangerous as no serum isavailable and a few fatalities have beenrecorded. It must be stressed, however,that bites are very rare and that this backfanged snake needs to hold on to its vic-tim for some time to inject poison.

71

Dasypeltis scabra(Linnæus, 1758)

Common namesCommon Egg-eater

TaxonomyMany subspecies have been describedand then synonimized, the last one wasDasypeltis scabra loveridgei (Mertens,1954) described from specimens collect-ed in Namibia and then regarded as sim-ply colour phase by Broadley (1990).Gans (1959) revised the systematics ofthe genus.

IdentificationA medium sized snake (60-70 cm), witha small rounded narrow head barely setoff from the neck. Teeth rudimentary.Body slender with relatively short tail(longer in males). Scales strongly keeledwith apical pit. Dorsal ground colourslate grey, light brown or olive brownwith a median series of brown squareblotches. Like Causus rhombeatus thetop of the head and the neck has a nar-row “V” shaped mark.

Geographic RangeWidely distributed in Africa south of theSahara except in deserts and denserainforest; there is a relict population inwestern Morocco.

Local distributionVesey-FitzGerald considered thisspecies as common in Arusha NationalPark; we found only one individual thatwas probably looking for the nests ofspur-winged plovers (Vanellus spinosus)on the shores of the Small Momela lake.

EcologyFeeds exclusively on eggs especially ofbirds, but sometimes lizard eggs too.Eggs are swallowed whole and crackedin the snake’s neck by a series of bonyprojections. The shell is then crushedand regurgitated. When annoyed thisharmless species reacts with somespectacular defensive behaviour suchas head triangulation to mimic viperidsnakes, hissing, false strikes and gape.It is also able to coil and uncoil in aseries of serrated loops to produce arasping sound similar to that producedby Echis (Gans & Richmond, 1957;Young et al., 1999).

Common Egg-eater snake from SmallMomela lake.

73

OTHER SPECIES

Atractaspis bibronii A. Smith, 1849

Common namesBibron’s Stiletto-snake

IdentificationBrown to black small burrowing snake, with an average adult total length of 45 cm.The head is rounded and not distinguishable from the neck. Head and neck not asbroad as the body. Typical of this family are the long fangs extending backward pos-terior to the eye, supralabials five, infralabial five (rarely six). A. bibronii has 21-25midbody scales row, 212-246 ventrals in males and 238-260 in females, frontal large,longer than broad, the length is more or less the same as parietals.

NotesA single snake that probably belonged to the Atractaspis genus was found at KinandiaView Point but unfortunately escaped while we were attempting to take a picture of it.Vesey-FitzGerald regards Atractaspis bibronii as common in Arusha National Park.This species bites protruding one of its long fangs and stabbing with closed mouth.The poison causes local pain and nasty necrosis, no effective serum is produced.

Causus rhombeatus (Lichtenstein, 1823)

Common namesRhombic Night-adder

IdentificationMedium sized snakes (usually up to 60 cm), the back is pale grey, olive brown oralmost black and with a typical “V” shaped mark on the neck and dark rhombic whiteedged blotches, some individuals are completely unpatterned. Other characteristics:enlarged grooved poison fangs in the upper jaw folded into membranous sheath,head not larger than the body covered by large shields, pupil round, snout not turnedup at the tip.

NotesThis nocturnal species is quite similar to the common egg eater snake but has a short-er and thicker body and tail. Vesey-FitzGerald considered C. rhombeatus common inArusha National Park. No fatalities have been recorded from the bite of this species.

Duberria lutrix (Linnæus, 1758)

Common namesSlug Eater Snake

IdentificationA small snake (30-35 cm); the specimen we found was completely black above andbelow except for few scattered white dots on the lower side of the head. The head is

74

small with nostril pierced in an entire nasal, loreal shield small or absent, eye smallwith round pupil. No enlarged poison fang in the upper jaw.

NotesThis upland species is regarded by Vesey-FitzGerald as common in Arusha NationalPark, we collected a specimen freshly killed by a car near the Rest house of the Park.

Aparallactus capensis (A. Smith, 1849)

Common namesCape Centipede-eater

IdentificationAnother small snake (up to 25-30 cm) with slender brown body and a distinctive blackhead backed by a black collar; the belly is grey white. Each nostril is pierced in anundivided nasal shield, loreal shield absent, subcaudal scales single, first pair ofinfralabials widely separated by the anterior sublinguals.

NotesThis species has enlarged poison fangs in the upper jaw but is completely harmless.Reported from Arusha National Park by Vesey-FitzGerald.

Prosymna stuhlmannii (Pfeffer, 1893)

Common namesShovel Snout Snake

IdentificationA small snake with flat head and short tail that average 30 cm in length. The back isdark brown to metallic black. No enlarged poison fangs in the upper jaw, dorsal scalessmooth, anal shield entire, dorsal scales in 15-17 rows at midbody.

NotesA burrowing nocturnal snake of the savannah, often feeds on geckoes’ eggs.Common in Arusha National Park (Vesey-FitzGerald, 1975).

Philothamnus hoplogaster (Günther, 1863)

Common namesSoutheastern Green Snake

IdentificationA small slender bright green uniform snake, yellow green below (50-70 cm max 96cm). 73-106 subcaudal shields, anal divided, scales in 15 rows at midbody.

NotesA species that is usually found near water and that resembles the green mamba.Occurs in Arusha National Park (Vesey-FitzGerald, 1975).

75

CHECK LIST OF AMPHIBIANS AND REPTILESAT THE ARUSHA NATIONAL PARK

The symbol “*” means that the presence of the species is based on bibliographic dataonly (Loveridge, 1959; Rand, 1958; 1963; Vesey-Fitzgerald, 1975).

AMPHIBIANS

Order Anura

Suborder Opisthocoela

Family PipidaeSubfamily Xenopodinae

Xenopus muelleri (Peters, 1844)

Suborder Procoela

Family BufonidaeBufo gutturalis Power, 1927

Family RanidaeSubfamily Raninae

Ptychadena mascareniensis (Duméril and Bibron, 1841)Rana angolensis Bocage, 1866Strongylopus fasciatus merumontanus (Lönnberg, 1910)

Subfamily PhrynobatrachinaePhrynobatrachus keniensis Barbour and Loveridge, 1928

Family HyperoliidaeSubfamily Kassininae

Kassina senegalensis (Duméril and Bibron, 1841)Subfamily Hyperoliinae

Hyperolius viridiflavus ommatostictus Laurent, 1951Hyperolius nasutus Günther, 1864

Family HemisotidaeHemisus marmoratum (Peters, 1854)

REPTILES

Order Chelonii

Family Testudinidae* Geochelone pardalis (Bell, 1828)

76

Order Sauria

Family GekkonidaePachydactylus turneri (Gray, 1864)Hemidactylus mabouia (Moreau de Jonnès, 1818)

Family AgamidaeAgama agama (Linnæus, 1758)

Family ChamaeleonidaeBradypodion tavetanum (Steindachner, 1891)* Chamaeleo dilepis Leach, 1819Chamaeleo gracilis Hallowell, 1842* Chamaeleo jacksonii merumontanus Rand, 1958* Chamaeleo rudis Boulenger, 1906

Family ScincidaeSubfamily Lygosomatinae

Mabuya striata (Peters, 1844)Mabuya varia (Peters, 1867)Lygosoma afrum (Peters, 1854)Panaspis wahlbergii (A. Smith, 1849)

Family LacertidaeAdolfus jacksoni (Boulenger, 1899)Nucras boulengeri Neumann, 1900

Order Scolecophidia

Family LeptotyphlopidaeLeptotyphlops scutifrons merkeri (Werner, 1909)

Order Scolecophidia

Family PythonidaePython natalensis A. Smith, 1840

Family ColubridaeCrotaphopeltis hotamboeia (Laurenti, 1768)Dasypeltis scabra (Linnæus, 1758)Duberria lutrix (Linnæus, 1758) * Hemirhagerrhis hildebrandtii (Peters, 1878)Lamprophis fuliginosus (Boie, 1827)Lycophidion capense jacksoni Boulenger, 1893Natriciteres olivacea (Peters, 1854)* Philothamnus hoplogaster (Günther, 1863)* Prosymna stuhlmannii (Pfeffer, 1893)Psammophis phillipsii (Hallowell, 1844)Thelotornis capensis mossambicanus (Bocage, 1895)

77

Family Viperidae* Bitis arietans arietans (Merrem, 1820)

Family Viperidae* Causus rhombeatus (Lichtenstein, 1823)

Family Atractaspidae* Aparallactus capensis (A. Smith, 1849)* Atractaspis bibronii A. Smith, 1849

Family ElapidaeElapsoidea loveridgei loveridgei Parker, 1949Naja haje haje (Linnæus, 1758)Dendroaspis angusticeps (A. Smith, 1849)

The species listed below have been collected in a large area around the Meru craterby Yngve Sjösted during the Swedish Zoological Expedition in 1905 (Lönnberg, 1910);few of them come from lower altitude areas along the Ngare Nanyuki and are com-pletely absent from the Park, but some others could be confirmed with further fieldresearches.Reptiles (29 species): Geochelone pardalis, Pelomedusa subrufa, Hemidactylussquamulatus, Lygodactylus conradti, Agama doriae [?, this is a West African species],Agama mwanzae, Laudakia atricollis, Varanus albigularis, Nucras boulengeri, Latastialongicauda, Heliobolus neumanni [or Heliobolus speckii], Gerrhosaurus nigrolineatus,Gerrhosaurus flavigularis, Mabuya megalura, Mabuya varia, Mabuya striata,Lygosoma sundevalli, Leptosiaphos kilimensis, Panaspis wahlbergii, Chamaeleogracilis, Bradypodion tavetanum, Leptotyphlops scutifrons, Lamprophis fuliginosus,Lycophidion capense, Dasypeltis scabra, Crotaphopeltis hotamboeia, Psammophissubtaeniatus, Aparallactus jacksoni, Causus rhombeatus.Amphibians (8 species): Rana fasciata merumontana, Rana angolensis, Rana fus-cigula [probably again Rana angolensis], Ptychadena oxyrhynchus, Ptychadenamascareniensis, Phrynobatrachus natalensis [probably Phrynobatrachus keniensis],Bufo gutturalis, Xenopus laevis.

79

REFERENCES

Andersson L.G., 1911. – Batrachians. In: Andersson L.G. & Lönnberg E. Reptiles,Batrachians and fishes collected by the Swedish Zoological Expedition to BritishEast Africa 1911. Kungl. Svenska Vetenskapsakademiens Handlingar, 47 (6): 25-41 + 1 plate.

Barbour T. & Loveridge A., 1928a. – New frogs of the genus Phrynobatrachus fromthe Congo and Kenya colony. Proceedings of the New England Zoological Club,10: 87-90.

Barbour T. & Loveridge A., 1928b. – A comparative study of the herpetological faunaeof the Uluguru and Usambara mountains, Tanganyika territory with descriptions ofnew species. Mem. Mus. Comp. Zool., Harvard College, 50 (2): 87-265 + 4 plates.

Bauer A., 1999. – Evolutionary scenarios in the Pachydactylus Group geckos ofsouthern Africa: new hypotheses. African Journal of Herpetology, 48(1/2): 53-62.

Bauer A., Branch W.R. & Haacke W.D., 1993. – The herpetofauna of the Kamanjaband adjacent Damarland, Namibia. Madoqua, 18(2): 117-145.

Benyr A., 1995. – Systematik und Taxonomie der Geckos des Pachydactylus bibronii- laevigatus Komplexes (Reptilia: Squamata: Gekkonidae). Diplomarbeit:Universitat Wien, Naturwissenschaftliche Fakultat. 75 pp + figs.

Blomberg S. & Shine R., 1996. – 7 reptiles. In: W.J. Sutherland (Editor), Ecologicalcensus techniques: a handbook. Cambridge University Press, Cambridge, pp.218-226.

Bowker R.G. & Bowker M.H., 1979. – Abundance and distribution of Anurans in aKenyan pond. Copeia, 1979(2): 278-285.

Branch B., 1994. – Field guide to Snakes and other reptiles of Southern Africa. Struik,Cape Town, 330 pp.

Branch W. R. & Haacke W.D., 1980. – A Fatal Attack on a Young Boy by an AfricanRock Python Python sebae. Journal of Herpetology, 14 (3): 305-306.

Brandstätter F., 1995. – Eine revision der Gattung Psammophis mit berücksichtigungder Schwesterngattungen innerhalb der Tribus Psammophiini (Colubridae:Lycodontinae). Teil 1: Die Gattungen und Arten der Tribus Psammophiini. Teil 2:Rasterelektronemikroskopische Untersuchungen zur Schuppenultrastruktur beiden Arten der Tribus Psammophiini mit besonderer Bercksichtigung der Arten derGattung Psammophis. Dissert. Doktor Naturwiss. Math.-Naturwiss. Fak. Univ.Saarlandes.

Brandstätter F., 1996. – Die Sandrennattern. Westarp-Wiss., Magdeburg; SpektrumAkad. Verlag, Heidelberg.

Broadley D.G. 1966a. – A review of the Riopa sundevalli group in southern Africa.Arnoldia (Rhodesia), 2(34): 1-7.

Broadley D.G. 1966b. – A review of the genus Natriciteres Loveridge (Serpentes:Colubridae). Arnoldia (Rhodesia), 2(35): 1-11.

Broadley D.G. 1966c. – A review of the African Stripe-bellied sandsnakes of the genusPsammophis. Arnoldia (Rhodesia), 2(36): 1-9.

Broadley D.G. 1968. – A new species of Crotaphopeltis (Serpentes: Colubridae) fromBarotseland, Zambia. Fieldiana Zoology, 51(10): 135-139.

Broadley D.G., 1971. – A revision of the African snake genus Elapsoidea Bocage(Elapidae). Occ. Pap. natn. Mus. Rhod. Ser. B, 4(32): 577-526.

80

Broadley D.G. 1977. – A review of the genus Psammophis in southern Africa(Serpentes: Colubridae). Arnoldia (Rhodesia), 8(12): 1-29.

Broadley D.G. 1979. – Problems presented by geographical variation in the Africanvine snakes, genus Thelotornis. S. Afr. J. Zool., 14: 125-131.

Broadley D.G., 1989. – A reappraisal of the genus Panaspis Cope with the descrip-tion of a new species of Leptosiaphos (Reptilia, Scincidae) from Tanzania.Arnoldia Zimbabwe, 9(32): 439-449.

Broadley, D.G., 1990. – Fitzsimons’ snakes of southern Africa, revised and updatedby Donald G. Broadley. Jonathan Ball and Ad Donker publisher, Johannesburg,387 pp.

Broadley D.G. 1994. – A review of Lygosoma Hardwicke and Gray, 1827 (ReptiliaScincidae) on the East African Coast, with the description of a new species.Tropical Zoology, 7(1): 217-222.

Broadley D.G., 1995. – The snouted cobra, Naja annulifera, a valid species in south-ern Africa. J. Herpetol. Ass. Africa, 44: 26-32.

Broadley D.G., 1996. – A revision of the genus Lycophidion Fitzinger (Serpentes:Colubridae) in Africa south of the Equator. Syntarsus, 3: 1-33.

Broadley D.G., 1999. – The southern African python, Python natalensis A. Smith1840, is a valid species. African Herp News, 29: 31-32.

Broadley D.G. 2000. – A review of the genus Mabuya in southeastern Africa (Sauria:Scincidae). African Journal of Herpetology, 49(2): 87-110.

Broadley D.G. 2001. – A review of the genus Thelotornis A. Smith in eastern Africa,with the description of a new species from the Usambara Mountains (Serpentes:Colubridae: Dispholidini). African Journal of Herpetology, 50(2): 53-70.

Broadley D.G. & Howell K.M., 1991. – A Check List of the Reptiles of Tanzania, withSynoptic Keys. Syntarsus, 1: 1-70.

Broadley D.G. & Watson G., 1991. – A review of the worm snakes of southeasternAfrica (Serpentes: Leptotyphlopidae). Occ. Pap. natn. Mus. Rhod. Ser. B, 5: 465-510.

Channing A., (in press). – Field guide to the frogs of Central and Southern Africa.Channing A. & Griffin M., 1993. – An annotated checklist of the frogs of Namibia.

Madoqua, 18(2): 101-116.Chippaux J.P., 1999. – Les serpents d’Afrique occidentale et centrale. Faune et Flore

tropicales. IRD Editions, Paris.Daniel P.M. 1961. – Notes on the life history of Agama agama africana (Hallowell) in

Liberia. The Ohio Herpetological Society, Special Publication, 3: 1-5.Duellman W.E. 1993. – Amphibian species of the world: additions and corrections.

Univ. Kansas Mus. Nat. Hist., Special Publications No. 21, 372 pp.Duellman W.E. & Trueb L., 1994. – Biology of Amphibians. 2nd edition. The Johns

Hopkins University Press, London, XXI + 670 pp.FitzSimons V.F., 1943. – The Lizards of South Africa. Transvaal Museum Memoir No.

1, Pretoria, XV + 528 pp + 28 plates.FitzSimons V.F., 1962. – Snakes of southern Africa. Mac Donald, London, 423 pp.Frank N., & Ramus E., 1996. – A complete guide to scientific and common names of

reptiles and amphibians of the world. Reptile and Amphibian magazine, Pottsville,377 pp.

Frost D.R. 1985. – Amphibian species of the world, a taxonomic and geographical ref-erence. The Association of Systematic Collections, Lawrence, Kansas, 732 pp.

Frost D.R., 2000. – Amphibian Species of the World: An online reference.V2.20http://research.amnh.org/herpetology/amphibia/index.html (1 September 2000).

81

Gans C., 1959. – A taxonomic Revision of the African Snake Genus “Dasypeltis”(Reptilia: Serpentes). Ann. Mus. R. Congo Belge, Terv. (sér. 8) Sci. Zoo., 74: 1-237 + Xpp + 13 pl.

Gans C. & Richmond N.D., 1957. – Warning behaviour in snakes of the genusDasypeltis. Copeia, 1957(4): 269-274.

Grafe T.U., 1996. – Energetics of vocalization in the African reed frog (Hyperolius mar-moratus). Comparative Biochemistry & Physiology, 114(3): 235-243.

Grafe T.U. & Linsenmair K.E., 1989. – Protogynous sex change in the reed frogHyperolius viridiflavus. Copeia, 1989: 1024-1029.

Greer A.E. Jr., 1974. – The generic relationships of the scincid lizard genusLeiolopisma and its relatives. Aust. J. Zool., Suppl. Ser. 31: 1-67.

Hailey A. & Coulson I.M., 1999. – The growth pattern of the African tortoiseGeochelone pardalis and other chelonians. Canadian Journal of Zoology, 77(2):181-193.

Halliday T.R., 1996. – 6 Amphibians. In: W.J. Sutherland (Editor), Ecological censustechniques: a handbook. Cambridge University Press, Cambridge, pp. 205-217.

Heyer R.W., Donnelly M.A., McDiarmid R.W., Hayek L.A.C. & Foster M.S., 1994. –Measuring and monitoring Biological Diversity. Standard Methods for Amphibians,1. Smithsonian University Press, 364 pp.

Hodgson P.S. & Davidson T.M., 1996. – Biology and treatment of the mambasnakebite. Wilderness & Environmental Medicine, 7(2): 133-145.

Houlahan J., Findlay C.S., Schmidt B.R., Meyer A.H. & Kuzmin S.L., 2000. –Quantitative evidence for global amphibian population declines. Nature, 404: 752-755.

Howell K.M., 1982. – Geographic distribution – Sauria: Eremias cf. neumanni Tornier(Neumann’s Sand lizard). Herp. Review, 13(2): 52.

Hughes B., 1999. – Critical review of a revision of Psammophis (Linnaeus, 1758)(Serpentes, Reptilia) by Frank Brandstätter. African Journal of Herpetology,48(1/2): 63-70.

Inger R.F. & Marx H., 1961. – The food of amphibians. In: Mission G.F. de Witte:Exploration du Parc National de l’Upemba. Institute des Parc Nationaux du Congoet du Ruanda-Urundi, 64: 1-86.

Jacobsen N.H.G., 1997. – Sub-order Sauria, family Agamidae. Page 6 in: van Wyk(Ed.), Proceedings of the Fitsimons Commemorative Symposium, South Africanlizards: 50 years of progress and Third H.A.A. Symposium on African Herpetologyheld at the Transvaal Museum, Pretoria, South Africa 11-15 October 1993. H.A.A.,Cape Town, IX + 227 pp.

Jacobsen N.H.G. & Broadley D.G. 2000. – A new species of Panaspis Cope (Reptilia:Scincidae) from southern Africa. African Journal of Herpetology, 49(1): 61-71.

Jakobsen A. 1997. – A review of some East African members of the genus ElapsoideaBocage with the description of a new species from Somalia and a key for thegenus (Reptilia, Serpentes, Elapidae). Steenstrupia, 22: 59-82.

Joger U., 1991. – A molecular phylogeny of Agamids lizards. Copeia, 1991: 616-622.Kabigumila J., 2000. – Growth and carapacial colour variation of the leopard tortoise,

Geochelone pardalis babcocki, in northern Tanzania. African Journal of Ecology,38(3): 217-223.

Klaver C. & Böhme W., 1986. – Phylogeny and classification of the Chamaeleonidae(Sauria) with special reference to hemipenis morphology. Bonn. zool. Monogr., 22:1-64.

Klaver C. & Böhme W., 1997. – Chameleonidae. Das Tierreich, 112(14): 1-85.

82

Lambert M., 1985. – A-herping in Tanzania, but hardly in Loveridge’s footsteps. BritishHerpetological Society Bulletin, 12: 19-27.

Lambert M.R.K., 1987. – More of the herpetofauna in the Commonwealth (EthiopianZone). British Herpetological Society Bulletin, 21: 13-22.

Lambert M.R.K., & Dewhurst C.F. 1998. – Gecko predation by skink observed inTanzania., British Herpetological Society Bulletin, 62: 29-30.

International Commission on Zoological Nomenclature 1999. – International Code ofZoological Nomenclature. ICZN, Padova, 306 pp.

Laurent R.F., 1964. – Reptiles et Amphibiens de l’Angola, Subsídios para o estudo dabiologia na Lunda. Companhia de Diamantes de Angola, Serviços Culturais,Lisboa, 165 pp.

Laurent R.F., 1968. – A re-examination of the snake genus Lycophidion Duméril &Bibron. Bull. Mus. comp. Zool. Harvard. 136(12): 461-482.

Laurent R.F., 1972. – Tentative revision of the genus Hemisus Günther. Annls. Mus.R. Afr. Cent. Sci. Zool., (8)194: 1-67.

Lambiris A.J.L., 1989a. – A review of the amphibians of Natal, Lammergeyer, 39: 1-212.

Lambiris A.J.L., 1989b. – The frogs of Zimbabwe, Monografia X. Museo Regionale diScienze Naturali, Torino, 247 pp.

Largen M.J., 1997. – An annotated checklist of the amphibians and reptiles of Eritrea,with keys for their identification. Tropical Zoology, 10: 63-115.

Lönnberg E., 1910. – 4. Reptilia and Batrachia. In: Sjöstedt Y. (Ed.),Wissenschaftliche ergebnisse der schwedischen zoologischen Expedition nachdem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905-1906. Band 1, Abteilung 1-7, Stockholm: 1-28 + 1 plate.

Lönnberg E. 1911. – Reptiles In: Andersson L.G. & Lönnberg E. Reptiles, Batrachiansand fishes colleted by the Swedish Zoological Expedition to British East Africa1911. Kungl. Svenska Vetenskapsakademiens Handlingar., 47 (6): 1-24 + 1 plate.

Loveridge A., 1925. – Notes of East African batrachians, collected 1920-1923 with thedescription of four new species. Proc. Zool. Soc. London, 1925 (2): 763-791 + 2tav.

Loveridge A., 1935. – Scientific Results of an Expedition to rain forest regions inEastern Africa. I New Reptiles and Amphibians from East Africa. Bull. MusCompar. Zool., Cambridge, 79(1): 1-19.

Loveridge A., 1936. – Scientific Results of an Expedition to rain forest regions inEastern Africa. V Reptiles. Bull. Mus Compar. Zool., Cambridge, 79(5): 209-337 +9 pl.

Loveridge A., 1936. – Revision of the African snakes of the genera Dromophis andPsammophis. Bull. Mus. comp. Zool. Harvard, 87(1): 1-69.

Loveridge A., 1953. – Zoological results of a fifth expedition to East Africa IV:Amphibians from Nyasaland and Tete. Bull. Mus Compar. Zool., Cambridge,110(4): 325-406.

Loveridge A., 1957. – Check list of the Reptiles and Amphibians of East Africa(Uganda; Kenya; Tanganyika; Zanzibar). Bull. Mus Compar. Zool., Cambridge,117(2): 153-362 +XXXVI.

Loveridge A., 1959. – On a fourth collection of reptiles, mostly taken in Tanganyika ter-ritory by Mr.C. J. P. Ionides. Proc. zool. Soc. London, 133(1): 29-44.

MacKay A. & MacKay J. 1985. – Poisonous snakes of Eastern Africa and the treat-ment of their bites. A. & J. MacKay, Nairobi, 95 pp.

Marais J., 1992. – A complete guide to the snakes of Southern Africa. Krieger

83

Publishing Company, Malabar, Florida, 208 pp. + 210 plates.Meirte D., 1992. – Clés de determination des serpents d’Afrique. Musée Royal de

l’Afrique Centrale, Sciences Zoologiques, 267: 1-152.Meshaka W.E., 2000. – Colonization dynamics of two exotic geckos (Hemidactylus

garnotii and H. mabouia) in Everglades National Park. Journal of Herpetology,34(1): 163-168.

Moodley G.K. & Biseswar R., 1997. – Reproduction in male Hemidactylus mabouia(Sauria: Gekkonidae). page 201 in: van Wyk (Ed.), Proceedings of the FitsimonsCommemorative Symposium, South African lizards: 50 years of progress andThird H.A.A. Symposium on African Herpetology held at the Transvaal Museum,Pretoria, South Africa 11-15 October 1993. H.A.A., Cape Town, IX + 227 pp.

Moody S.M., 1980. – Phylogenetic and historical biogeographical relationships of thegenera in the family Agamidae (Reptilia: Lacertilia). Ph.D. thesis, University ofMichigan, Ann Arbour, USA.

Neças P., 1999. – Chameleons, nature’s hidden jewels. Chimaira, Frankfurt am Main,352 pp.

Nishikawa K.C., Kier W.M., Smith K.K., 1999. – Morphology and mechanics of tonguemovement in the African pig-nosed frog Hemisus marmoratum: A muscular hydro-static model Journal of Experimental Biology. 202(7): 771-780.

Passmore N.I. & Carruthers V.C., 1995. – South African frogs. A complete guide.Revised edition. Witwaterssrand University press, Johannesburg, 322 pp.

Passmore N.I., Carruthers V.C. & Zähringer J., 1995. – South African frog calls,Passmore & Carruthers CD guide. Megatone studios, Johannesburg, 4 pp + AudioCD.

Patterson J.W., 1990. – Female reproductive cycles in two subspecies of the tropicallizard Mabuya striata. Oecologia, 84: 232-237.

Perret J.L., 1975. – La différentiation dans le genre Panaspis Cope (Reptilia,Scincidae). Bull. Soc. neuchateloise, Sci. Nat., 98: 5-16.

Pitman C.R.S., 1974. – A guide to the snakes of Uganda. Revised edition. Whelon &Wesley, London, 290 pp.

Poynton J.C., 1964. – Amphibia of the Nyasa-Lungwa region of Africa. Senck. Biol.,45 (3/5): 193-225.

Poynton J.C. & Broadley D.G., 1985a. – Amphibia Zambesiaca 1. Scolecomorphidae,Pipidae, Microhylidae, Hemisidae, Arthroleptidae. Ann. Natal Mus., 26(2): 503-553.

Poynton J.C. & Broadley D.G., 1985b. – Amphibia Zambesiaca 2. Ranidae. Ann.Natal Mus., 27(1): 115-181.

Poynton J.C. & Broadley D.G., 1987. – Amphibia Zambesiaca 3. Rachophoridae andHyperoliidae. Ann. Natal Mus., 28(1): 161-229.

Poynton J.C. & Broadley D.G., 1988. – Amphibia Zambesiaca 4. Bufonidae. Ann.Natal Mus., 29(2): 447-490.

Poynton J.C. & Broadley D.G., 1991. – Amphibia Zambesiaca 5. Zoogeography. Ann.Natal Mus., 32: 221-277.

Rand A.S., 1958. – A new subspecies of Chameleo jacksoni Boulenger and a key tothe species of three-horned chamaeleons. Breviora, 99: 1-8.

Rand A.S., 1963. – Notes on the Chamaeleo bitaeniatus complex. Bull. Mus. com.Zool., Harvard, 130(1): 1-29.

Richards C.M., 1981. – A new color pattern variant and its inheritance in some mem-bers of the superspecies Hyperolius viridiflavus (Duméril & Bibron), (AmphibiaAnura). Mobit. Zool. Ital., n.s. 15 suppl. (16): 337-351.

Rödel M.O., 2000. – Herpetofauna of West Africa. Vol. I Amphibians of West AfricanSavanna. Edition Chimaira, Frankfurt am Main, 334 pp.

Rose W., 1962. – The reptiles and amphibians of Southern Africa. Maskew Miller,Cape Town, XXIX + 494 pp.

Schiøtz A., 1975. – The treefrogs of eastern Africa. Steenstrupia, Copenhagen, 232pp.

Schiøtz A., 1999. – Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.Schleich H.H., Kastle W. & Kabish, K., 1996. – Amphibians and Reptiles of North

Africa, biology, systematics, field guide. Koeltz Scientific books, Koenigstein, 630pp.

Snelson D. & Bygott D., 1987. – Arusha National Park. Tanzania National Parks &African Wildlife Foundation, Nairobi, 52 pp.

Stebbins R.C. & Cohen N.W., 1995. – A natural History of amphibians. PrincetonUniversity Press, Princeton, 316 pp.

Stewart M.M., 1967. – The amphibians of Malawi. State University of New York Press,New York, 164 pp.

Uetz P., 2001. – Classification of living reptiles, http://www.embl-heidelberg.de/~uetz/LivingReptiles.html (4th July 2001).

Van Dijk D.E., 1997. – Parental care in Hemisus (Anura, Hemisotidae). South AfricanJournal of Zoology, 32(2): 56-57.

Vesey-Fitzgerald D.F., 1975. – A guide to the snakes of the Tanzania and Kenya bor-derlands. Jl. E. Africa nat. Hist. Soc. & natn. Mus., 149: 1-26.

Wieczorek A. M., Drewes R. C. & Channing A., 2001. – Phylogenetic relationship with-in the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comment on tax-onomic status. Amphibia Reptilia 22(2): 155-165.

Webb J.K., Shine R., Branch W.R. & Harlow P.S., 2000. – Life history strategies inbasal snakes: reproduction and dietary habits of the African thread snakeLeptotyphlops scutifrons (Serpentes: Leptotyphlopidae). J. Zool. Lond., 250: 321-327.

Young B.A., Lalor J. & Solomon J., 1999. – The comparative biomechanics of anophidian defensive behaviour: head triangulation in hognose snake (Heterodon)and an egg-eating snake (Dasypeltis). Journal of Zoology, 248(2): 169-177.

Published in Varese (Italy) by Pubblinova Edizioni Negriand Istituto OIKOS in February 2002

field guide to the amphibians and reptiles of … · of arusha national park (tanzania) field guide...

Documents