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 Published by Struik Nature
(an imprint of Penguin Random House South Africa (Pty) Ltd)
Reg. No. 1953/000441/07
The Estuaries No. 4, Oxbow Crescent, Century Avenue, Century City, 7441
PO Box 1144, Cape Town, 8000 South Africa
Visit www.penguinrandomhouse.co.za and join the Struik Nature Club for updates, news, events and special
offers.

First published in 2021

1 3 5 7 9 10 8 6 4 2
Copyright © in text, 2021: Darren Pietersen, Luke Verburgt, John Davies
Copyright © in photographs, 2021: As indicated alongside individual images or at Photographic Credits on p.
371
Copyright © in illustrations, 2021: Gary Nicolau
Copyright © in maps, 2021: Darren Pietersen, Luke Verburgt, John Davies
Copyright © in published edition, 2021: Penguin Random House South Africa (Pty) Ltd

Publisher: Pippa Parker

Managing editor: Roelien Theron
Editor: Heléne Booyens
Designer: Dominic Robson
Proofreader: Emsie du Plessis

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted,
in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior
written permission of the copyright owner(s).
ISBN 978 1 77584 737 3 (Print)
ISBN 978 1 77584 738 0 (ePub)

Front cover: Katanga Bush Viper (Frank Willems), Anchieta’s Chameleon (Darren Pietersen),
O’Shaughnessy’s Thick-toed Gecko (Gary Brown), Leopard Tortoise (Errol Pietersen)
Half-title page: Speckled Thick-toed Gecko (Luke Kemp)
Title page: Marbled Tree Snake (Luke Verburgt)
Contents page: Rhombic Egg-eater, Flap-necked Chameleon (Luke Verburgt)
Spine: Rhombic Egg-eater (Luke Verburgt)
Back cover: Olive Sand Snake, Nile Crocodile (Luke Verburgt)
DEDICATIONS
To my parents, Errol and Michèle, and my brother Ryan. Thank you for all your love and for always
supporting my ideas, no matter how crazy they were.
– Darren Pietersen

To Peter Snyman, who fuelled my passion for herpetofauna and shared his immense knowledge with me. And
to my mom, Janine, my wife, Ursula, and my best friend, Sam – none of this would have been possible
without you, and I am forever grateful for the love, support and opportunities you’ve given me.
– Luke Verburgt

To my parents, Peter and Ona, who inspired so much of my passion for nature; my sister Lynne, for putting up
with my obsessive birdwatching; and my partner, Lindy, for all her love, patience and support.
– John Davies
ACKNOWLEDGEMENTS
A field guide of this nature would not have been possible without the assistance of numerous
individuals and organisations.
We gratefully acknowledge the following parties for sharing their data with us: Ditsong
National Museum of Natural History (South Africa), specifically Lauretta Mahlangu; Port
Elizabeth Museum, Bayworld (South Africa), specifically Werner Conradie; and the
Bulawayo Museum (Zimbabwe), specifically Shiela Broadley. The Biodiversity Institute,
specifically René Navarro and Les Underhill, are thanked for sharing the ReptileMAP data
with us. Werner Conradie, Johan Marais, Colin Tilbury, Norman Barrett, Gary Brown, Frank
Willems and Paul Lloyd are also thanked for sharing their personal data, and many
photographs, with us. Thanks to the many people who took the time to share their reptile
observations on ReptileMAP and other public citizen science platforms: you have greatly
contributed towards shaping the geographic distributions that we present in this field guide.
We are very grateful to the team at the Vegetationmap4Africa project
(vegetationmap4africa.org) for allowing us to use their vegetation map in the predictive
modelling of the species geographic distributions.
Gary Nicolau is thanked for drawing the excellent scale diagrams included throughout
this field guide.
For allowing us to use their photographs, we thank André Coetzer, André van Hecke, Bill
Branch, Carl Huchzermeyer, Caspian Johnson, Colin Tilbury, Derek Solomon, Errol
Pietersen, Fabio Pupin, Frank Willems, Gary Brown, Gary Nicolau, Harald Hinkel, Johan
Marais, Joseph Zulu, Laurent Chirio, Lesley Reynolds, Liam Baisley, Luis Ceriaco, Luke
Kemp, Matej Dolinay, Michele Menegon, Mike McLaren, Nick Evans, Norman Barrett, Paul
Lloyd, Ronald Auerbach, Stephen Spawls, Tomáš Mazuch, Tyrone Ping, Warren McCleland
and Werner Conradie. If we have missed anyone, we sincerely apologise – it was not
intentional.
Aaron Bauer, Bill Branch, Werner Conradie and Colin Tilbury are thanked for assisting
us in obtaining many obscure scientific publications.
We are extremely grateful to Pippa Parker, Heléne Booyens, Dominic Robson and the
entire team at Struik Nature for their help and guidance.
Finally, we thank our families, friends and partners for all their love, support and
understanding.
Contents
PREFACE

FOREWORD

INTRODUCTION
What are reptiles?
Taxonomy, classification and nomenclature
Habitat types in Zambia and Malawi
Biogeography
Venom types and snakebite treatment
Regional endemics
Reptile identification using scales
How to use this book

SCALED REPTILES – SQUAMATA

Snakes – Serpentes
Slender Blind Snakes
Blind Snakes
Pythons
Adders and Vipers
Sand Snakes and relatives
Shovel-snouts
Mole Snakes and relatives
African Burrowing Snakes
Old World Snakes
Malagasy Snakes and relatives
Mambas, Cobras and relatives
Old World Water Snakes
Typical Snakes

Lizards – Sauria
Geckos
Plated Lizards
Girdled Lizards
Legless Skinks
Old World Skinks
 Pacific Skinks
Advanced Skinks
True Skinks
Worm Lizards
Old World Lizards or Lacertids
Monitors
Chameleons
Agamas

CHELONIANS – TESTUDINES
Side-necked Terrapins – Pleurodira
African Terrapins

Modern Chelonians – Cryptodira

Soft-shelled Terrapins
Land Tortoises

CROCODYLIANS – CROCODYLIA
Crocodiles

Glossary
References
Photographic credits
Useful contacts
Index to scientific names
Index to common names
Preface
Zambia and Malawi are somewhat of a herpetological enigma, at least in the public sphere.
They are too far north to be included in southern African field guides, yet too far south for
East African field guides. As such, Zambia and Malawi have remained veritable
herpetological ‘black holes’. Until now, no detailed field guide that covers the entire region
has been published.
The idea behind this book originated during a trip to Zambia, when we became
increasingly frustrated at the difficulty of identifying reptiles without a suitable consolidated
resource. Once back home, we started digging, and realised that although there was a wealth
of information regarding the rich reptile diversity of Zambia and Malawi, it was scattered
across dozens – if not hundreds – of often obscure publications. Most casual nature
enthusiasts wouldn’t know where to start, and many of these publications are difficult to
obtain.
We decided to consolidate as much of the available information as we could. So began an
endeavour that was to last several years. We have done our utmost to ensure that the
information contained in these pages is as accurate and up to date as possible. It often
necessitated us to track down the original descriptions of species or reviews of genera, and to
update this information if anything additional has been published subsequently. Through the
field trips we undertook and the data and records we reviewed, several new discoveries were
made for both Zambia and Malawi. Some of these discoveries have already been
documented, while some newly discovered species remain to be formally described.
It is our hope that this field guide will enable people to identify the wealth of reptiles that
surround them in Zambia and Malawi, dispel the myths about all (or even most) snakes being
venomous, and generally instill a deeper appreciation for reptiles, which play important roles
in the ecosystem. We also hope that this book will stimulate further research into the reptiles
of Zambia and Malawi by professionals and amateurs alike, and consequently assist in the
conservation of these wonderful creatures.
Although we have made every effort to ensure that the information in this field guide is
error-free, any mistakes or oversights contained in these pages are our own. If you notice any
inaccuracies or have any questions, you are welcome to contact us.

Darren Pietersen
[email protected]

Luke Verburgt
[email protected]

John Davies
[email protected]
Foreword
In 1910, FW Fitzsimons wrote The Snakes of South Africa, the first guide of its kind. Some
three decades later, it was followed by Vivian F Fitzsimons’s comprehensive The Lizards of
South Africa. Other classics on African herpetology include Captain Charles RS Pitman’s A
Guide to the Snakes of Uganda (1938), Margaret Stewart’s Amphibians of Malawi (1967)
and RCH Sweeny’s Snakes of Nyasaland (1961).
Today, there are plenty of social media pages where snakes (and other reptiles, to a lesser
extent) feature daily. Much of what is posted is inaccurate, however, and websites circulate a
great deal of misinformation.
I was delighted to hear that Darren Pietersen, a well-known and respected South African
herpetologist, was working on a book on the reptiles of Zambia and Malawi with the help of
John Davies, who works with avifauna at the Endangered Wildlife Trust. Luke Verburgt,
another respected South African herpetologist, soon joined the team. It was an ambitious
project, as much of the information on the herpetofauna of Malawi and Zambia was scattered
across various scientific papers.
With their combined skills, and over many years, they have produced a comprehensive
book that will not just appeal to the layman wanting more information on the interesting
reptile diversity of Zambia and Malawi, but will also become the go-to reference for experts
in the field and required reading for those studying African herpetology.
The authors chose well when they teamed up with Struik Nature, one of the world’s
finest natural history publishers. The design and layout of the book is exceptional and makes
for easy reading, while the contents are well researched and accurate. Distribution maps are
vitally important in any field guide, and here the authors used their vast experience and
knowledge of reptile distribution in the region as well as a clear understanding of the diverse
habitats to produce maps that will be of great use to herpetologists.
The book covers 117 snake species, 108 lizard species, 13 terrapins and tortoises and two
crocodiles. It is lavishly illustrated with over 380 colour photographs. In an age of mediocrity
and misinformation, it is great to see a herpetological book of this calibre. It will fit in well
with other classics on African herpetology.

Johan Marais
African Snakebite Institute
Introduction
Zambia and Malawi are landlocked African countries in the southern hemisphere. Perhaps
their best-known geographical features are the Zambezi River (the longest east-flowing river
in Africa), which arises in Zambia, and Lake Malawi (Nyasa), an East African rift lake that
covers more than 20% of Malawi. The region’s tropical climate, diverse landscape features
(ranging from towering inselbergs to lake shores) and wide range of vegetation types are
among the primary driving forces behind a high reptile diversity. To date, 240 species have
been recorded in these two countries: 117 snakes, 108 lizards, 13 terrapins and tortoises, and
two crocodiles. New species are continually ‘discovered’ using genetic techniques, as
widespread species in fact often comprise various closely related and similar-looking species.

Eastern Bark Snake

 O’Shaughnessy’s Thick-toed Gecko
Gary Brown

Eastern Hinged Tortoise

Tyrone Ping

WHAT ARE REPTILES?

Reptiles are a diverse group of vertebrates that belong to the class Reptilia. They come in an
extraordinary array of shapes and sizes, with extant reptiles including serpentes (snakes),
saurids (lizards), chelonians (tortoises, terrapins and turtles), crocodylians (crocodiles) and
sphenodonts (tuataras). Reptiles are characterised by a dry, keratinous skin, usually modified
into scales or plates, although in some species, such as the Zambezi Soft-shelled Terrapin
(Cycloderma frenatum), the scales are covered in leathery skin. Scales consist of a horn-like
structure called keratin, and provide protection from predators, injuries and moisture loss.
All reptiles are ectothermic, relying on outside energy sources (such as the sun) to heat
up. This is why they are often seen basking on cold mornings, and why reptile activity
decreases during cold periods. Because they rely on external heat sources rather than
metabolic heat, reptiles also eat less than mammals or birds of a similar size.
Most reptiles reproduce sexually through internal fertilisation and lay fluid-filled, self-
contained eggs that usually incubate unguarded outside the mother’s body. The eggs are
either soft- or hard-shelled, and in most species are abandoned after being deposited. Eggs
are usually laid in a sheltered, warm, moist spot, and in many species the sex of the offspring
is determined by the incubation temperature. When they hatch, the young are smaller replicas
of their parents – hatchlings of venomous snakes, for example, are just as venomous as their
parents.

The Nile Crocodile is one of two crocodile species found in the region.
André van Hecke

TAXONOMY, CLASSIFICATION AND NOMENCLATURE

Taxonomy is the branch of science that assigns names to organisms and documents how they
are related to each other. Organisms are grouped by traits that unify them and distinguish
them from all other organisms. These traits can be morphological (i.e. external), internal,
genetic, or even behavioural. Taxonomic classification is a hierarchical system, meaning that
each group higher up on the taxonomic ladder is more inclusive than the one below it. As
such, several species form a genus (in a few instances, a genus may contain a single species),
several genera are grouped together to form a family, several families form a class, and so
on.
Scientific (or Latin) names are those by which an organism is known to the scientific
community. Binomial nomenclature is the convention of assigning two-part names consisting
of a genus and species name, developed by the Swedish naturalist Carl von Linnaeus in 1735.
This approach forms the basis of modern taxonomic classification. Common (or vernacular)
names do not follow any specific convention and generally differ between languages and
sometimes between regions.
Taxonomy is dynamic, and the scientific names of species continually change as new
data become available. This can take several forms: a simple name change; one species being
split into two or more species; or forms thought to be different species being synonymised
(treated as the same species).
This book has been arranged to largely reflect the taxonomic relationships of families and
genera within each order or suborder, with families and genera that are most closely related
following each other, using the most recent large-scale phylogenetic studies as a baseline. In
a few instances, we feel that the presented evidence does not sufficiently support the
proposed taxonomic changes and elect to use the previously accepted taxonomy. Within each
genus, species accounts are arranged alphabetically.
Several ongoing studies will affect the taxonomy of reptiles in the region, and more
studies are sure to follow. For the most recent taxonomy of the species featured in this book,
we direct the reader to The Reptile Database (www.reptile-database.org), a useful online
platform that captures updated reptile taxonomy.
Where possible we standardised our use of vernacular names based on the most recent
regional reptile guides. We also consulted The Reptile Database for vernacular names of
species not occurring in existing guide books, and to see additional vernacular names in use.
We generally settled on the most frequently used names, modifying some where appropriate
and to reflect recent taxonomic changes.

Leopard Tortoise

Common Tropical House Gecko

 Southern African Python

The Leopard Tortoise, Common Tropical House Gecko and Southern African Python all belong to the same
phylum (Chordata) and class (Reptilia). They are known to the scientific community by the binomial names
Stigmochelys pardalis, Hemidactylus mabouia and Python natalensis.

CLASSIFICATION OF REPTILES WITHIN THE ANIMAL KINGDOM

PHYLUM The animal kingdom is divided into approximately 35
phyla, with reptiles belonging to the phylum Chordata.
Members of this phylum have, during some period of
their development, a notochord, a dorsal nerve chord,
pharyngeal slits and a post-anal tail. Most chordates
belong to the subphylum Vertebrata, which is
characterised by organisms having a vertebral column
(backbone comprising bony vertebrae) and a distinct
skull.

CLASS There are seven classes of chordates, including Reptilia

(reptiles), Aves (birds), Amphibia (amphibians),
Mammalia (mammals) and three classes of fish.

ORDER Reptiles are divided into four orders, namely Testudines

(tortoises, terrapins and turtles), Crocodylia (crocodiles),
Squamata (scaled reptiles) and Sphenodontia (tuataras),
the latter found only in New Zealand. The order
Squamata is further divided into the suborders Sauria
(e.g. lizards, chameleons and agamas) and Serpentes
(snakes).

FAMILY There are 92 reptile families globally, 32 of which occur

in the region.

GENUS At present, some 1,206 reptile genera are recognised

worldwide, 97 of which have been recorded in the
region.

SPECIES Currently, 11,341 reptile species are recognised

globally, of which 240 have been recorded in the region.
Over time, the number of recognised species will
continue to increase both regionally and globally.

Evolutionary relationships within the class Reptilia

From an evolutionary perspective, birds ought to be

included in the class Reptilia, but are traditionally placed
in a separate class, Aves, a convention followed in this
 guide. Crocodylia is most closely related to Aves, while
Sauria and Serpentes are more closely related to each
other than to any of the remaining reptiles.

HABITAT TYPES IN ZAMBIA AND MALAWI

Forests
Forests consist of tall, thick trees with interlocking crowns. They are usually rich in plant
species, but owing to geological and hydrological characteristics the species composition and
height of the forest can vary over short distances. Forests are broadly categorised into two
types: moist forests and dry forests. In the region, moist forests are most prevalent in the east
where the higher moisture content (from frequent rain and/or mist) sustains them. They
usually have a cool, moist and relatively dark interior and the mid- and lower storeys
(including climbers or lianas) may be poorly to well developed, depending on the forest type.
Moist forests are generally fire-sensitive and thus often occur in isolated patches in gullies,
along rivers or other landforms providing protection from fire. They are also generally
intolerant of frost. Of the moist forests, Afromontane rainforests 1 are restricted to
high-elevation, high-rainfall regions, particularly on mountain slopes and plateaus. Swamp
forests or mushitu’s 2 occur mainly at the headwaters of larger rivers and along smaller
streams in north-western Zambia, thinning out to riparian forest in the middle and lower
stretches of these rivers. Swamp forests are characterised by tall, dense evergreen trees, often
with a well-developed understorey. Many of the trees and shrubs may be seasonally or
permanently submerged in shallow water, resulting in a very marshy habitat. Riparian
forests 3 occur in isolated pockets along some of the larger rivers, usually not extending
far into the surrounding vegetation and often forming a relatively abrupt boundary with it.
Dry forests are restricted to the drier western regions. In the region, they are represented by
the Cryptosepalum dry forest (also known as mavunda) in north-western Zambia and
adjoining Angola. This forest is nearly impenetrable, and many interesting discoveries
undoubtedly await the intrepid explorer. In many areas, forests are heavily impacted on by
human activities, which include felling for timber, charcoal production and clearing land for
agriculture.
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