Kinixys Conservation Blueprint

Transcription

I
A Comprehensive Assessment to Ensure the Future of the Genus
Herpetological Resource & Management, LLC
ii
Copyright 2014 © Herpetological Resource and Management, LLC.
Photographs by Herpetological Resource and Management, LLC. unless otherwise noted. Kinixys art by
Tricia Green Brockman. Document layout and format design by Hailey R. Brown.
ISBN: 978-0-9915169-1-9
Suggested Citation: Mifsud, David A., and Maegan M. Stapleton. 2014. Kinixys Conservation
Blueprint: A Comprehensive Assessment to Ensure the Future of the Genus. Herpetological Resource
and Management Technical Publication 2014.
Cover photo credits from top left: HRM, Sean Zera, Hutter Auctions, Lisa Call, Sean Zera, Clive Reid, HRM, Olivier Pauwels, Nathan
Gibbs. Inside cover photo: Victor Loehr.
Thomas Leuteritz
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Acknowledgements
This project was supported by the Mohamed bin Zayed Species Conservation Fund (Project
Number 12055390), the Conservation International Foundation and Turtle Conservation Fund
(Project Number 63321), the Tortoise and Freshwater Turtle Specialist Group, and Minnesota
Society of Herpetologists. This work does not necessarily represent the views of our project
funders.
The authors would like to express our gratitude for the support and encouragement of Dr. Brian
Horne, Dr. Russell Mittermeier, Dr. Anders Rhodin, and Dr. Peter Paul van Dijk. We hope work
conducted in Togo as part of the IUCN Red List Assessment will be a start of great things for
chelonian conservation in Africa.
A special thank goes to Dr. Donald Broadley who has contributed greatly to the pioneering
work on Kinixys and was a great inspiration for the development of this document. Dr. Broadley
has been a helpful resource along the way and an asset to Kinixys conservation. We are also very
grateful to Jim Harding for providing access and use of his personal library of books on African
reptiles and turtle conservation.
We are very grateful for the wonderful resources and information those working with Kinixys have
provided along the way. Thanks goes to Dr. William Ahrens, Dr. Phil Allman, Dr. Laurent Chirio,
Tomas Diagne, Dr. Adrian Hailey, Dr. Margaretha D. Hofmeyr, Dr. John Iverson, Dr. Dwight
Lawson, Dr. Luca Luiselli, Dr. Olivier S.G. Pauwels, Dr. Fabio Petrozzi, Dr. Gabriel Hoinsoude
Segniagbeto, and Dr. Jean-François Trape as well as others who have contributed to Kinixys
research and helped expand our knowledge and understanding of these incredible tortoises. This
work will build on your incredible efforts. Thanks also goes to all those who paved the way and
inspired us all to be passionate about turtles and tortoises.
Numerous individuals have contributed information, translation of manuscripts, photographs,
and resources instrumental in the creation of this publication including, Dr. Ernst Baard, James
Badman, Dr. Patrick Baker, Richard Bartlett, Ruth Basham, Chris Bednarski, Anat Belasen, Mary
Bohling, Gary Brown, Karol Chlopecki, Mallory Clark, Andre Coetzer, Gehard Diedericks, Kurt
Edwards, Megan English, Eric Fouchard, Chris Gertiser, Piet Grobler, Russ Gurley, Cris Hagen,
Ed Hammer, Jennifer Hammer, Dr. Michael W. Hance, Curtis Hart, Toby Hibbitts, Richard Hay,
Chris Hobson, Brian Horne, Dr. Chris Howe DVM, Rick Hudson, Kelly Hull, Neels Jackson,
Jovita JovskiRush-Morgan, Dr. Ross Kiester, Terry E. Kilgore, Sheena Koeth, Barry Lambert,
Dwight Lawson, Christine Light, Dr. Day Ligon, Dr. Thomas Leuteritz, Victor Loehr, Mark M
Lucas, Dr. Bruce G. Marcot, Liana May, Mick Mittermeier, Matt Muir, Jacob Mueti Ngwava, Paul
Rabiega, Reginald T. Mwaya, Dr. Terry M. Norton DVM, Michael Ogle, Paul Rabiega, Wayne
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Rawlinson, Dr. Sam Rivera, Ben Santhouse DVM, Simone Swiel, The Turtle Conservancy, Ryan
Van Huyssteen, Chris Woodley, Sean Zera and the participants of the IUCN/SSC Tortoise and
Freshwater Turtle Specialist Group’s workshop on Conservation Status of the Tortoises and
Freshwater Turtles of Sub-Saharan Africa.
We would also like to thank the Turtle Survival Alliance and the IUCN Tortoise and Freshwater
Turtle Specialist Group for their support in providing a venue for the First Kinixys Conservation
Taskforce meeting at the 11th Annual Symposium on the Conservation and Biology of Tortoises
and Freshwater Turtles. This forum allowed for Kinixys researchers, conservation groups, zoos,
hobbyists, and concerned citizens to meet to discuss the current state of knowledge on Kinixys and
develop objectives for things to accomplish. We are also appreciative to the Turtle Survival Alliance
for their help in setting up assurance colonies for K. homeana and K. erosa and their dedication to the
protection of Kinixys in Africa.
Though recognized above we would like to thank those individuals who were willing to review
and provide comment for the document including Donald Broadley, Jim Harding, Brian Horne,
Dwight Lawson, Christine Light, and Sean Zera. Your comments and edits were incredibly helpful
and much appreciated.
Every attempt has been made to integrate feedback from the professionals and organizations listed
above. Their participation however does not suggest the endorsement of every recommendation
or professional opinion in this manual. This document was created using the best available
information available at the time of publishing and as new literature and research becomes
available, modification may be necessary.
We would like to recognize our HRM team members for their help on this report. Thank you Tricia
Green Brockman for taking your personal time to make such descriptive and colorful depictions of
these tortoises. Special thanks also go to Hailey R. Brown for her time, organization, and technical
skill in processing data and document layout. We would also like to thank Brittany Price for her
help and patience in compiling resources and helping develop range maps for the genus.
On a personal level, David would like to thank his mother Mary Mifsud and wife Rachel Mifsud
for whom their great love, support, and understanding of having a son and husband passionate
about tortoises conservation. You have helped me succeed in life and profession.
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Table of Contents
Executive Summary..................................................................................1
Overview...................................................................................................3
Objective ...............................................................................................3
Data Analysis and Approach ..................................................................4
Kinixys Tortoises.....................................................................................5
Taxonomy and Phylogenetics..............................................................6
Population Estimates...........................................................................7
Critical Areas and Zones of Sympatry..................................................7
Threats ...............................................................................................7
Habitat loss......................................................................................7
Agricultural conversion.................................................................9
Fire . ............................................................................................9
Fragmentation............................................................................10
Mineral and Oil Extraction .......................................................11
Deforestation and Timber Harvesting.........................................12
Animal Trade.................................................................................13
Kinixys homeana................................................................................. 18
Kinixys erosa....................................................................................... 26
Kinixys belliana.................................................................................. 36
Kinixys nogueyi................................................................................... 42
Kinixys spekii...................................................................................... 48
Kinixys zombensis............................................................................... 55
Kinixys natalensis............................................................................... 63
Kinixys lobatsiana.............................................................................. 68
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Conservation, Management, and Recovery......................................... 73
Habitat Protection ...............................................................................73
Trade ...................................................................................................74
Education and Outreach.......................................................................76
Adaptive Management .........................................................................76
Assurance Colonies...............................................................................77
Husbandry and Captive Breeding ........................................................78
Reintroduction Programs......................................................................78
Genetic Management............................................................................79
Research Needs.....................................................................................79
Appendix A:Species Table................................................................... 81
Appendix B: Maps.............................................................................. 84
Appendix C: Data Tables.................................................................. 106
Appendix D: Assurance Colony Husbandry Recommendations....... 118
Housing..............................................................................................118
Seasonal or Permanent Outdoor Enclosures........................................120
Diet....................................................................................................120
Health................................................................................................122
Reproduction......................................................................................124
Maintenance .....................................................................................126
Appendix E: Kinixys Questionnaire.................................................. 127
Appendix F: Kinixys Activity Chart.................................................. 128
References......................................................................................... 130
1
Executive Summary
Status
The genus Kinixys, commonly known as the Hingeback Tortoises, is a historically
understudied and monitored group of chelonians in need of greater conservation
attention and management. This genus occupies a vast region across sub-Saharan Africa;
however, population densities and distributions are currently unknown for many species.
A recent taxonomical revision of the genus which elevated two subspecies to full species
has resulted in the recognition of eight species: K. homeana, K. erosa, K. nogueyi, K. belliana,
K. spekii, K. natalensis, K. lobatsiana, and K. zombenis. The latter includes 2 subspecies: K.
z. zombensis and K. z. domerguei. There are numerous gaps in data regarding population
viabilities, threats, habitat use, natural history, and conservation needs. Based on current
literature it is highly likely that all Kinixys are in decline throughout the range. Until
recently, over 75% of Kinixys were considered Not Evaluated (NE) or Data Deficient
(DD). The data from an International Union for Conservation of Nature (IUCN) Red
List assessment workshop conducted in 2013 for sub-Saharan African turtles and tortoises
resulted in the draft ranking of all species as Vulnerable (K. nogueyi, K. spekii, K. belliana, K.
natalensis, K. lobatsiana, and K. zombensis), Endangered (K. erosa), or Critically Endangered (K.
homeana).
Threats
Hingeback tortoises are faced with multiple stressors that continue to reduce their range,
population size, and overall species viability. Habitat degradation and loss are likely major
causes of declining Kinixys populations. Factors contributing to the loss of high quality
habitat across Africa include fragmentation caused by infrastructure (i.e., roads, fences),
climate change, unsustainable land management (i.e., agricultural conversion, intentional
use of fire), mineral and oil extraction, and timber harvesting. Additionally, significant
numbers of Hingeback tortoises are unsustainably collected (both legally and illegally)
from the wild. Kinixys face extensive pressures from collection as they are hunted for
consumption and trade in bushmeat and fetish markets while others are exported for sale
in both the legal and illegal international pet trade.
2
Conservation Objectives and Recommendations
The Kinixys Conservation Blueprint (KCB) provides information on species distribution,
habitat and ecology, global and regional status, domestic and international threats,
conservation and management, and husbandry and captive breeding. The primary objective
of the KCB is to provide a broad/holistic approach to implement conservation measures
aimed at reducing the impacts and pressures on this genus and develop achievable
objectives to help maintain viable populations and species protection throughout Africa.
The long-term conservation strategy to successfully preserve Hingeback tortoises requires
a multifaceted approach and international partnerships utilizing adaptive management.
Several recommended strategies that should be considered to provide immediate and future
conservation of this genus include:
• Conduct additional in situ research on all Kinixys to better understand species
ranges and distributions, with an emphasis placed on K. belliana.
• Conduct comprehensive surveys targeting the Malagasy endemic K. z. domerguei
assessing natural history, threats, and conservation needs.
• Focus research to better understand Kinixys ecology and natural history
including reproductive behavior, nesting, incubation and associated diapause,
juvenile diet, and habitat selection.
• Collect genetic samples to better determine the number of species within the
genus and better define their geographic distribution and relatedness among
populations.
• Increase habitat protection through enforcement of stricter regulations.
• Protect and restore corridors of protected habitat to combat fragmentation
of populations within species ranges. Priority should be given to zones of
sympatry that support multiple species.
• Reduce export quotas to minimize collection pressure.
• Eliminating quotas for wild collected or “ranched” animals for K. homeana and
K. erosa is strongly recommended.
• Improve trade monitoring for Kinixys and other African illegally collected
wildlife.
• Within range countries, enact and enforce regulations to protect Kinixys from
sale in food or fetish markets.
• To meet the demand and interest in these species, placing a greater emphasis on
captive bred animals is encouraged.
• Establish both in situ and ex situ assurance colonies to help safeguard species
and increase public outreach and learning opportunities.
• Captive programs should be used as a means to develop a better understanding
of Kinixys behavior and husbandry needs including incubation, developmental
diapause, and TSD.
Richard Penryn
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2. Introduction
Overview
Gabriel Segniagbeto
Objective
1. Deforestation and
conversion of habitat to
agriculture is a common
sight throughout west
Africa as seen here in
Adélé, Togo.
Although research has been conducted with
select species, little is known about this genus
in many parts of its range. Numerous data
gaps exist regarding species viability, threats,
habitat use, natural history, reproduction,
longevity, population structure, husbandry,
and conservation needs by species within range
country and range-wide. This genus as a whole
is highly vulnerable to continued decline and
potential extinction if proactive conservation
measures are not implemented quickly. To best
2
Thomas Leuteritz
2. Endemic to
Madagascar, K. z.
domerguei is potentially
vulnerable to heavy
illegal collection,
incompatible land
management, and
habitat loss. The
subspecies is known
to inhabit secondary
growth forests as seen
here in Nosy Faly.
Despite Africa’s rich assemblage of tortoises, conservation
and management efforts for most species have been limited,
inadequate, or ineffective. Over the last 50 years tortoises of
the genus Kinixys have undergone significant declines from
various threats including substantial habitat loss, degradation
and fragmentation, massive collection for bushmeat and the pet
trade, and poor land management. Though this genus is afforded
some protection through the Convention on International
Trade in Endangered Species (CITES) with Appendix II status
and protected from legal collection in some countries, decline
throughout its range continues. Until recently, over 75% of the
species within the Kinixys genus were considered Not Evaluated
(NE) or Data Deficient (DD) by The International Union for
Conservation of Nature (IUCN) Redlist. Based on results of the
1
Redlist assessment focused on sub-Saharan African chelonians
conducted in August of 2013, all species of Kinixys were evaluated
as being provisionally Vulnerable, Endangered, or Critically
Endangered with forest species identified as those in greatest immediate need for conservation
action.
Richard Stupart
4
1
manage and conserve this genus, research to collect qualitative
and quantitative baseline data will be required to allow assessment
of population trends over time. To date, no comprehensive effort
has been made to compile and synthesize available research
and resources on the entire genus. The Kinixys Conservation
Blueprint (KCB) can be a critical step in developing a
comprehensive framework for conservation and best management
of the Kinixys genus. The goal of the KCB will be to serve as a
conservation and resource tool summarizing the current collective
knowledge on this genus to provide general recommendations
to help best protect and manage this genus. Through proactive
planning, science-based conservation, and adaptive management,
this genus of unique and ancient tortoises can continue to be a
common sight on the African landscape.
Data Analysis and Approach
Meg English
Multiple data sets were used in the analyses of this assessment.
Results compiled in Appendix B include maps of species’ ranges,
species potential habitat, and fire density and frequency within
species ranges, as well as human population density centers and
roads, land cover type, distribution of mining operations and
type, and elevation for Sub-Saharan Africa. Tables and figures
representing trade information are included in Appendix C.
1.Land management
in Africa frequently
involves burning to
convert the land for
agricultural use.
2. Nearby controlled
bush fires have created
a haze of smoke
through the Olifants
West Nature Reserve in
South Africa.
Range maps are based on best available data and should be used
as a starting point for further refinement of the ranges for Kinixys
species. Evaluation and depiction of species’ ranges were based on
2
published literature, museum records, and unpublished, but often
photo-vouchered observations from researchers and volunteers.
An implied degree of spatial uncertainty is assumed with the range maps, as presently available
information is inadequate to accurately depict current ranges for the genus. These maps include
historic ranges and likely do not represent current distributions.
Spatial analysis was conducted using the most recent publically available land cover data for
Africa to assess potential available habitat within species ranges. GlobCover Land Cover v2 was
used for the analsyis. This dataset is a global land cover map with 300 meter resolution and land
cover classes defined within the United Nations Land Cover Classification System (LCCS). Data
developed by the ESA GlobCover 2008 GlobCover products are based on the ENVISAT satellite
mission’s MERIS sensor (Medium Resolution Image Spectrometer) The GlobCover Land Cover
v2 data was derived from a time series of MERIS composites covering the period December
2004 to June 2006. This data was queried to display specific land cover classification types
within Sub-Saharan Africa that represent potential community types associated with targeted
Kinixys species. These land cover types were grouped to provide an approximate available habitat
count and spatial distribution. This assessment was intended to serve as a coarse-level analysis
based on assumed historical distribution overlaid with land use data. These ranges and associated
potential habitats do not account for heavy collection pressure or other factors that are not
measureable in available GIS-based datasets. Based on trends and impacts across Africa in the
5
1. This juvenile K. spekii
was found on the road
in Kruger National
Park, South Africa.
Much of Kinixys range
is fragmented by roads
(Appendix B, Map 4).
2
A map displaying human population density
centers was generated using the African
Population Database by Nelson (2004). This
dataset contains population information from
2000 and was created in 2011. A topographic map
3
of Africa was created using 30 arc-second DEM
of Africa by U.S. Geological Survey’s Center for
Earth Resources Observation and Science (1996).
The dataset Mineral Operations of Africa and the Middle East by Eros and Candelario-Quintana
(2006) was used to display the distribution of mining activities and type within Africa.
Liana May
Analyses of trade data were conducted using CITES trade
statistics derived from the CITES Trade Database, UNEP World
Conservation Monitoring Centre. From this information the
leading export countries, top ten wildlife species traded, and total
number of exported Kinixys and specifically K. belliana, K. erosa, K.
homeana, and K. spekii between 1975 and 2012 is depicted. A table
of all reported trade data for the species of Kinixys is also included.
4
Wayne Rawlinson
4. Savanna Kinixys
prefer more open
habitats such as coastal
forests and miombo
woodlands. For this
document, spatial
analysis was performed
for each species to
better estimate the
amount of suitable
habitat present within
their range (Appendix
B).
1
Shane Boylan, S.C. Aquarium
3. Omnivorous Kinixys
have an extremely
variable diet compared
to other tortoises and
are known to consume
a substantial amount of
live prey.
Maps depicting fire density were based on
information collected by the Moderate Resolution
Imaging Spectroradiometer (MODIS) on NASA’s
Terra Satellite. This dataset shows the locations
of actively burning fires around the world on a
monthly basis and was clipped to display portions
of Kinixys ranges where fire occurred between
May 2013 and February 2014 The colors are
based on a count of the number (not size) of fires
observed within a 1,000-square-kilometer area.
White pixels show the high end of the count with
as many as 100 fires in a 1,000-square-kilometer
area per day. Yellow pixels show as many as 10
fires, orange shows as many as 5 fires, and red
areas as few as 1 fire per day.
Curtis Hart
2. X-rays demonstrate
the unique physiology
of K. homeana.
last decade it is assumed that these values are
very likely lower than shown and this should be
considered when reviewing the data. Due to the
small scale of K. z. domerguei range and resolution
of data, spatial analysis was not performed for this
subspecies. On the ground research is needed to
confirm the amount and condition of remaining
habitat for this endemic Malagasy Kinixys.
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Tyrone Ping
Kinixys Tortoises
Neels Jackson
1
2
Kinixys is a unique genus within the Testudinidae family which
occurs throughout most of sub-Saharan Africa. Kinixys species
display a wide range of ecological specializations not found among
other groups of land tortoises. Hingeback tortoises are unique
among chelonians in having a hinged carapace, which they use
as a defense against predation and perhaps as a flexion point to
facilitate oviposition. There are eight species and one subspecies of
Kinixys, which can be broadly characterized as either savanna or
forest species. Occurring from arid scrubland to tropical forest, they
exhibit adaptations to a wide range of environments. Kinixys are also
unique as compared to most other tortoises due to their omnivorous
diets, which include an appreciable percentage of live food (Obst
1976). Forest Kinixys have even been observed hunting in floodplain
pools for fish and amphibian larvae. Though live food can make up
a seasonally large percentage of their intake, fruit is an important
part of the Kinixys diet and this group is recognized as important
seed dispersers in their habitats (Branch 2008). These tortoises are
also known to act as scavengers as they will opportunistically feed
on carrion. Kinixys are sentinel species and critical contributors to
healthy and functional ecosystems across the African landscape.
They are an excellent indicator species with a prominent midlevel
position in food webs and their conservation is a critical component
to preserving Africa’s natural heritage.
Terry E. Kilgore
Taxonomy and Phylogenetics
1-3. While the genus
Kinixys shares similar
traits such as a hinged
carapace, there is great
variability among its
species as seen here
between K. z. zombensis
(1), K. lobatsiana (2), and
K. erosa (3).
The KCB is not intended to be a comprehensive taxonomical
description of Kinixys species but can serve as a general guide for
identification purposes. This document includes a table (Appendix A)
and illustrations showing the basic distinctions between species, and
also provides references for more detailed descriptions.
The genus has undergone many taxonomical changes since it was
initially described by the English zoologist Thomas Bell in 1827.
3
For a complete description of historical Kinixys nomenclature and
taxonomy, refer to Fritz and Havas (2007). Until recently, a majority
of taxonomic classifications were based solely on morphological features (Broadley 1989,1993);
however, recent studies have also investigated the molecular phylogenetics of Kinixys. The latest
taxonomic revision to the genus has resulted in the elevation of a number of former subspecies
to full species status. As recently as 2012, six species were recognized within the genus; K.
erosa, K. homeana, K. lobatsiana, K. natalensis, K. spekii, and K. belliana, which included four
subspecies (K. belliana belliana, K. b. nogueyi, K. b. zombensis, and K. b. domerguei). Kindler et
al. (2012) supported the elevation of K. b. nogueyi and K. b. zombensis to full species status with
K. b. domerguei as a subspecies of K. zombensis. K. belliana no longer contains any subspecies
and eight species of Kinixys are now recognized (van Dijk et al. 2012). This conclusion is
supported by species morphology, habitat use, and range. As additional field work is conducted
7
a better sense of range-wide and population-level genetics can be
determined as well as the potential for identifying additional species
or documentation of intergrades in zones of sympatry.
Liana May
Population Estimates
1. Areas of continuous
open woodlands
provide critical habitat
for multiple Hingeback
species.
2. Deforestation
leaves large gaps
in the canopies
of mature forests
drastically altering these
ecosystems as seen here
on Mount Gorongosa,
Mozambique.
Few population estimates are available on this group of tortoises
and natural population densities are largely unknown. Based on
current available literature, trade data, and testimony from locals
and researchers in range countries it appears that these animals are
declining in numbers and likely experiencing collection rates that
1
exceed recruitment rates consistent with viable populations. Though
additional field surveys are warranted to accurately assess population
status, it can be said with a high degree of confidence that all Kinixys species are in decline and in
danger of local and even range-wide collapse due to several stressors impacting these populations.
Critical Areas and Zones of Sympatry
Conservation priority should be given to ecological “hotspots” as well as zones of sympatry and
multiple species overlap. Several species of Kinixys occur within areas that have been designated
as biodiversity hotspots by Myers et al 2000 (i.e., a region containing exceptional concentrations
of endemic species as well as exceptional loss of habitat). The coastal forests of Tanzania and
Kenya potentially support sympatric populations of K. spekii, K. belliana, and K. z. zombensis
while the West African forests are known to support K. homeana and K. erosa and to a limited
extent K. nogueyi. Although not identified as a hotspot, the forests of the Albertine Rift in
eastern Democratic Republic of the Congo,
southwestern Uganda, and northern
Rwanda have been identified as an area
of high conservation priority (Myers et al
2000) and populations of K. homeana, K.
erosa and K. belliana likely occur together
there. Additionally, in southern Africa,
multiple species share portions of their
range including K. natalensis, K. lobatsiana,
K. spekii, and K. z. zombensis.
Threats
Habitat loss
2
Philip Powell
Habitat degradation and loss are
considered the major factors threatening
wildlife populations throughout most of
the world and are a significant threat to
Africa’s natural resources. The continent
contains multiple biodiversity hotspots
with rich floral and faunal diversity and
is also experiencing one of the most rapid
Jacob Fahr
8
1
2
1. The range of K. spekii, one of the
largest among Hingebacks, will likely be
affected by climate change.
2. Variation in climatic patterns has
resulted in less productive crop yields
across Africa and more people are
converting tropical forests to agriculture
than ever before.
rates of habitat conversion and loss in the world.
No other region on earth houses a greater diversity
of tortoises than the continent of Africa and these
unique reptiles are especially vulnerable to loss
of habitat (Biggs et al 2008). Tortoises endemic
to this continent face greater risks of declines
and extinctions as many are highly specialized
in terms of habitat requirements, making them
particularly sensitive to environmental changes.
Hingeback tortoises span over a large portion of
sub-Saharan Africa and are known to occupy a
vast array of vegetation types that currently face
various threats due to human activities and climatic
variation. Slash and burn agriculture, logging,
fuel wood extraction, oil production, mining, and
infrastructure development are among the activities
contributing to rapid changes in land cover in
Africa (Weber et al 2001).
Climate Change
Africa is experiencing recurring droughts, crop failures, and water
scarcity and has been identified in many climate change prediction
models as the most vulnerable populated region of the world. Ongoing
clearing and degradation of vegetation is likely contributing to increases
in temperatures and wind speeds and decreases in precipitation and
relative humidity thus creating warmer and drier climates (Pricope et
al. 2013). In southern Africa where populations of K. lobatsiana and K.
natalensis occupy small ranges, the clearing of vegetation contributes
greatly to desertification. This type of land degradation that occurs in
arid, semi-arid, and dry sub-humid areas is a large constraint on Africa
where only 14% of the continent is free of soil moisture stress (Reich et
al. 2004). Dry and windy conditions have left lands previously used for
agriculture unsuitable for crop production, forcing locals to further alter
their landscape in order to grow food. This will become a significant
problem as models are predicting more extreme rain and drought
events in the future that will make farming even more unreliable
(Conservation International 2014). Localized alterations in the rate and
frequency of precipitation have also been reported in tropical regions
where significant logging has occurred. Even portions of protected areas
will experience altered landscapes due to changes in precipitation if
significant portions of an area are cleared (Weber et al. 2001). Wildlife
in Africa faces major obstacles in the future as their habitats dwindle
not only because of climate change but also from the actions of humans
who are adapting to a shifting climate as well.
Simone Swiel
Flickr User - Floeschen
H. Sinica
9
1. Conversion of land
for grazing livestock is
a contributing factor to
habitat loss in Africa.
2. Savanna landscapes
are intentionally burned
for a variety of reasons.
3. The shell of this
Hingeback tortoise has
been damaged by a fire.
1
Agricultural conversion
A large portion of Africa’s economy is supplemented by agriculture
which employs 65% of the continent’s labor force and accounts
for 32% of the gross domestic product (The World Bank 2013).
As human population densities grow exponentially throughout
sub-Saharan Africa and climatic variation continues to contribute
to declining crop yields, communities are converting natural
landscapes for agricultural use at alarming rates (Maitima 2009).
Typical agricultural practices include the use of shifting cultivation
which is characterized by the shift between fields rather than
between crops on the same field. As a result, patches of land are
used for a short period of 1-3 years for crop production followed
by 15-30 years of a fallow period (Nasi et al. 2012). As demand
for food increases, less and less time is being allowed for fallow and
soil regeneration to occur resulting in soils that are degraded with
heavy weed infestations and depleted seed banks and unsuitable for
crop production. Even more damaging than the overuse of fallow
fields is the establishment of permanent agriculture (Rainforest
Conservation Fund 2014).
It is estimated that 38% of agricultural lands in Africa have been
converted from tropical rain forests (Rainforest Conservation Fund
2014). By 2050 it is expected that trends in cropland conversion
2
will include grasslands in the north and deciduous woodlands
in the south (Heubes et al. 2012). The loss of these unique and
productive ecosystems will have a large effect on the wildlife
that they support. The most extensive losses of biodiversity are
expected to occur in Angola, Democratic Republic of the Congo,
and Sudan, which are among the remaining areas on earth that
have high agricultural potential but have not yet been extensively
farmed (Biggs et al. 2008). Within this region multiple Kinixys
species may be present including, K. belliana, K. spekii, and K.
erosa. Additionally, it is predicted that within southern Africa,
grassland vegetation will be highly impacted in the future due to
its suitability for agriculture which may affect several Hingeback
tortoise species including K. spekii, K. z. zombensis, K. lobatsiana,
3
and K. natalensis (Biggs et al. 2008). The planting of monoculture
crops is known to cause significant declines of local biodiversity
and can be mitigated through the implementation of poly-culture crops containing numerous
types of produce (Maitima et al. 2009).
Fire
Across sub-Saharan Africa, fire is heavily used as a means to keep unwanted vegetation from
encroaching on crops or pastures, to divert grazing animals away from crops, remove crop
stubble, return nutrients to the soil, convert natural ecosystems to cropland, and to clear
vegetation for infrastructure. In West Africa, the burning season generally extends from January
through April or May and occurs between crop harvesting and the following growing season. On
Clive Reid
10
1. K. z. zombensis
crossing a road in
Imfolozi-Hluhlu,
South Africa. Roads
serve as population
sinks due to deaths
from vehicle collisions
as well as increased
exposure to predation
and collection.
2. In addition to
the direct mortality
of wildlife, roads
fragment the
landscape and limit
the movement of
animals.
the continent, the burning season sweeps from north to south
as the dry season progresses. While this practice may result in
land that is more suitable for use as agriculture, the process also
results in the loss of high quality habitat on which wildlife like
Kinixys depends (Giglio et al. 2003). Both naturally occurring
fires and those intentionally set are enhanced by drought, past
fires, and logging or other disturbances (Weber et al. 2001).
Although fires occur naturally throughout the continent,
especially in grasslands and savannas, man-made fires are
creating landscapes that are vulnerable due to excessive burning
in the past. The rate at which fire returns to an area determines
which plants and therefore which animals can survive there
and when fire returns too quickly, the land may become too
degraded for growing crops or grazing livestock, thus creating
a landscape that is unsuitable for both humans and wildlife
(Giglio et al. 2003). In southern Africa, tortoises are known to
experience direct mortality from fire (Boycott and Bourquin
2000) and it is highly likely that Kinixys do as well. To put
into perspective the extent of fire impacts, burn data from May
2013 through February 2014 was overlaid with Kinixys’ range
1
data to demonstrate the extent of area burned that is potentially
occupied by Kinixys. With the exception of K. erosa which
is typically associated with wet forest communities most species overlapped with substantial fire
density within their range during this time period (Appendix B). These maps indicate the location,
distribution, and number of deliberate fires used in an attempt to manage the landscape. (Giglio
et al 2003). Review of fire data from 2005 shows even greater fire densities across the landscape
indicating that burning seasons vary in fire density and some years may have greater potential
impacts to Kinixys and other declining wildlife.
Fragmentation
In Africa as in other regions, large contiguous areas of pristine habitat are often fragmented by
roads, railways, and fences. While roadways are a useful tool for improving access to healthcare
and education, especially in more remote regions of the continent, they also disrupt the natural
landscape and cause harm to wildlife. Besides road-related mortalities, wildlife living in areas with
high densities of roads may experience
limited dispersal abilities resulting in
disrupted migratory movements and low
population viability due to a lack of genetic
input from neighboring populations.
2
Michael Ayers
New roads in Africa are typically built for
accessing previously remote areas as well
as for transporting the natural resources
obtained from them. Funding for road
construction is provided primarily by
foreign logging and mining companies
but there is also extensive effort in many
Lynette Knott Rudmann
11
1. Fences are a
common sight across
the African landscape.
Electric fences are
known to cause direct
mortality to tortoise
species including
Hingbacks.
2. These retention
ponds in Sudan
are used to contain
contaminants
produced during oil
extraction. Pools such
as these can infiltrate
groundwater and
pollute local water
sources.
African nations to upgrade and expand the region’s highway
network (Coghlan 2014). In addition to roads, fences significantly
contribute to the fragmentation of African landscapes. In South
Africa there are currently thousands of kilometers of fencing
used to divide farms, national parks, and individual properties
(Collinson 2014). Although these fences are known to limit the
number of road-related wildlife mortalities, they also severely
constrain the movement of animals across the landscape.
Additionally, fences that are electrified are known to kill wildlife
that come into contact with them including Hingeback tortoises
1
(Beck 2010, Collinson 2014). Due to their natural defensive
behaviors, tortoises are especially vulnerable to electrocution. Once
in contact with the fence, individuals typically withdraw their head
and limbs into the shell while remaining in contact with the live wire thus continuing to receive
electrical shocks. Direct mortality frequently occurs from the destruction of cells by electrolysis;
however, tortoises that are stuck along fences and exposed to direct sunlight for long periods may
also suffer from heat stress and dehydration (Beck 2010). Furthermore, the smooth metal wires
used for electric fences are known to provide snare material for poachers (Beck 2010). Negative
effects of roads and fences on wildlife can be mitigated through the use of modified drainage
culverts, underpasses, and overpasses (Collinson 2014). Further measures that can be taken to
reduce the risk of fences to tortoises include raising the height of the lowest strand to no less than
200 mm in order to allow these small animals to safely pass under while maintaining the integrity
and functionality of the structure.
United Nations Environment Programme
Mineral and Oil Extraction
The removal of minerals and oil from the African landscape is having a large effect on its once
pristine ecosystems and the wildlife that depend on them. Based on data analysis of the Mineral
Operations of Africa and the Middle East data set (Eros and Candelario-Quintana 2006), there are
currently over 650 known mines within Africa
(Map 3), and presumably many more unknown
illegal operations. The continent’s abundant
gold, diamonds, iron ore, uranium, and other
valuable resources are targeted largely by foreign
companies. The acquisition of Africa’s natural
resources directly threatens the landscape
with deforestation, degradation of the natural
resources, and pollution. Indirect threats include
those caused by infrastructure built to support
such projects (Nasi et al. 2012, Weber et al.
2001). Within Cameroon, a Chinese-owned
mining company has recently begun developing
the landscape to support an operation that will
include a 500 kilometer dedicated rail line that
will intersect 100 kilometers of forest in order
to reach a custom-built transport terminal at
the port of Lolabé (Downes 2013). Large scale
2
Jason Taylor
12
Allison M. Jones
Marko Laakkonen
1
1 -3. Clearing of forest
for timber harvest
results in loss and
degradation of habitat
for Kinixys and other
wildlife. In many cases,
even if forests are
allowed to regenerate
it can take decades
for the community
to resemble that of
unlogged areas.
Deforestation and Timber Harvesting
A large portion of the African continent is experiencing widespread
2
deforestation and the associated sale of materials for the timber
trade. These activities have been historically widespread with a high
concentrated in Central and West Africa. Approximately 0.2-0.6% of
forest in Central Africa is cleared annually and more than 0.9% on
average is cleared from West Africa (Weber et al. 2001). Within Ghana,
which has lost roughly 80% of its forest habitat since the 1920’s, 17
years of deforestation between 1955 and 1972 are estimated to have
removed 1/3 of the nation’s forests (McCullough et al 2005). In the
Congo sub-basin, industrial logging concessions are known to make up
at least 595,381 km2 of the region (Nasi et al. 2012). Across Central
Africa these concessions occupy 30-45% of all tropical forests and over
3
70% of forests in other countries (Poulsen et al. 2009). Unfortunately
the amount of data associated with forest extent, clearing, and
regeneration is alarmingly inadequate (Weber et al. 2001). Additionally, there is evidence of
corruption within the management of natural resources in some countries where large amounts
of timber are exported illegally. In 2012, 48% of China’s timber exports from Mozambique
were illegally obtained, costing the fourth least-developed country in the world tens of millions
of dollars in lost taxes (Environmental
Investigation Agency).
The extraction of timber products reduces
the extent of mature forests and alters species
composition. In heavily exploited areas,
logging results in tropical forest ecosystems
that more closely resemble those of temperate
forests (Weber et al. 2001). Some forest
understories have been shown to remain
dissimilar from their uncut counterparts
4
Tomas Diagne
4. K. homeana are
often collected during
logging and brought
back for sale in the pet
trade or for food as
shown here in Ghana.
construction efforts such as this are not uncommon. A significant
portion of pollution caused by extraction activities is associated with oil
production. The highest levels of high quality habitat loss occur in areas
where such operations are concentrated. Nigeria is Africa’s main oilproducing country and a majority of extraction occurs within the Niger
Delta, one of the largest wetland areas in West Africa and important
habitat for both K. homeana and K. erosa (Luiselli et al. 2004). Oil
spills result from breaks in pipelines that extend across vast ecosystems
including forests, swamps, and mangroves. Within this region between
1976 and 2001 roughly 7,000 spills involving millions of barrels of oil
have occurred and over 70% of the oil was never recovered (Obaji Ori
2013). These spilled chemicals are known to cause short term effects
such as direct mortality to wildlife as well as long-term degradation
through its toxic persistence in the environment and bioaccumulation
in the food chain (Luiselli and Akani 2003, Luiselli et al. 2004, Luiselli
et al. 2006)
Russ Mittermier
13
1. In order to transport
natural resources to
local villages, vast
infrastructure such as
roads is constructed.
2. Extensive logging
opens up previously
inaccessible areas
to hunters and puts
species like K. nogueyi
at risk of overexploitation.
for up to forty five years after logging has occurred (Weber et al.
2001). Within these forests, timber trees are killed directly during
the cutting process and as the large timber fall, they indirectly kill
numerous smaller trees and saplings. Additional vegetation is also
removed during the construction of logging roads (Weber et al.
2001). Large gaps created by logging reduce canopy cover within
a forest and increase the amount of sunlight penetrating the forest
floor. These large gaps in the canopy encourage the establishment
of quicker-growing, light-dependent species which can outcompete
the seedlings and saplings of the resident tree species. In many
logging regions, selective logging is utilized as a means of obtaining
1
sustainable yields of timber and is often promoted as a compromise
between completely destroying a forest versus leaving it intact. This
method does not guarantee a forest will sufficiently regenerate; in the Kibale Forest in Uganda 20
years after selectively logging, there does not appear to be any evidence of regeneration in logged
areas, particularly where large gaps were made (Weber et al. 2001).
Similar to mining and oil operations, landscapes surrounding logging concessions are indirectly
threatened by the infrastructure built to support the industry. These developments generally
include large numbers of new roads and expanding urban centers built to support large influxes
of migrant workers. A study of five logging towns in the Congo basin revealed that development
of the logging industry in these areas led to a 69% increase in the towns’ populations and 64%
increase in the demand for bushmeat. The illegal exploitation of wildlife is greatly increased in
those areas where previously inaccessible forests have been opened to hunters (Bushmeat Crisis
Task Force 2009, Poulsen et al. 2009).
Animal Trade
For decades it has been well known among African conservationists and national governments
that their wildlife is facing significant threats from hunting and collection of animals for
bushmeat markets as well as the legal
and black market pet trade. Although
data has generally focused on megafauna (e.g., elephants, rhinoceros, and
gorillas), smaller animals including
Hingeback tortoises are also targets for
hunters and collectors alike.
2
Tomas Diagne
Wildlife occupying the tropical forests
of Africa has historically been a vital
resource to the indigenous people
as food, folk medicine, and other
traditional practices. In the last several
decades as human populations have
dramatically increased and vast areas of
forest have been removed and exposed,
this traditional practice has become
14
Tomas Diagne
more commercialized and unsustainable
for wildlife species. Over 24 million people
are living in the forested regions of Central
and West Africa, and most rely on the
meat from wildlife as their primary source
of animal protein (Bushmeat Crisis Task
Force 2009, Wilkie and Carpenter 1999).
Given the high diversity of wildlife in these
forests, hunters are not typically targeting
a single species but will instead roam the
forests in search of any animal that is
1. K. nogueyi in a Togo
valuable in terms of food or sale. While
village near Kpalme
1
larger species such as duikers, primates,
with a noticeable cut
and
pigs
are
preferred,
populations
of
in the front of its
these animals have dwindled from overshell. This provides a
location to attach rope exploitation. As a result, smaller species are becoming more heavily exploited and evidence of this
to carry tortoises from can be seen throughout local bushmeat markets (Luiselli et al. 2013). Reptiles including tortoises
are a popular market item because they can be easily collected and sold (Weber et al. 2001). In the
the field.
Congo basin it is estimated that 1 to 5 million tons of bushmeat is harvested annually (Poulsen et
2. Many Kinixys in
al. 2009). For many poor rural families, hunting for bush meat offers a much needed short-term
Africa await export for source of money (Kümpel et al 2010). When wildlife is abundant, hunters can make between $400
the pet trade or sale
and $1100US per year from bushmeat alone which is comparable if not greater than salaries of
for food markets in
guards paid to protect land from hunters (Bushmeat Crisis Task Force 2009). Although wildlife is
cramped and poorly
becoming less abundant in many areas, the price of meat and low costs of hunting ensures that the
maintained enclosures.
bushmeat market stays profitable. The forests of this region have seen
tremendous increases in both commercial and subsistence hunting
with the arrival of logging camps. With these camps also comes
an expanding network of roads that fragment the forests and open
the areas to growing rural populations armed with non-traditional
hunting devices including shotguns and wire snares. Logging and other
extraction activities cause a large influx of workers, jobseekers, and their
families into previously remote areas. Foreign companies are often eager
to build a vast infrastructure capable of supporting these operations
and the masses of workers; however, many often fail to supply their
employees with adequate provisions of animal protein causing an
increased demand for bushmeat. In a study by Poulsen et al. (2009),
surveys of multiple logging concessions indicate alarming increases
in bushmeat consumption associated with the arrival of workers.
Immigrants were observed to harvest 72% of animals in markets and
consumed 66% of all bushmeat. The current rate of bushmeat harvest
in Africa has certainly reached unsustainable levels. Companies that
are large contributors to this trend can also play the largest role in
mitigating the effects of bushmeat hunting (Wilkie and Carpenter
1999). Guaranteeing adequate sources of protein for the workers and
their families while keeping prices competitive with bushmeat will
likely be good incentive for many locals to cease hunting. For workers
2
15
Matt Muir
who wish to continue hunting, companies should ensure
that hunting is conducted legally with proper licenses and
permits and impose penalties or fire workers who break the
law. Additionally, natural resource management should be
formalized with designated hunting territories and priority
should be given to local indigenous people. Furthermore,
a great deal of unsustainable bushmeat harvest may be
avoided if forest roads are restricted for company use only
and closed when not actively in use.
1. Hingeback tortoise in
a bushmeat market in
Equatorial Guinea. To
guarantee fresh meat for
consumption, tortoises are
kept alive until purchase
and are often prepared by
being dropped alive into
boiling water (BBC News
2012).
2. A fetish market in
Lomé, Togo displaying
a wide array of wildlife
including multiple
Kinixys species.
Hingeback tortoises are common in bushmeat and fetish
markets. Fetish markets include the sale of talismans,
objects that are believed to be infused with the power
of the divine and can heal and protect against various
1
ailments (Morton 2013). Traditionally indigenous people
have long utilized Kinixys for multiple purposes. Tortoise
shells are used for musical instruments, purses, cowbells,
etc. and meat is consumed as well as used for medicinal
purposes. The forest species K. homeana and K. erosa
are typically favored likely due to their larger size. Some
communities within the Niger Delta worship Hingeback
tortoises and do not consume them however many of these
tribes are losing their traditional cultures as immigration
has increased, making Kinixys more vulnerable (Luiselli
2003b). Historically the hunting of Kinixys was likely not
a significant threat to population sizes due to low human
population densities and the proportion of their habitat
that was not accessible to humans. In recent decades
considerable increases in human populations coupled with
extensive habitat loss has caused the collection of Kinixys
from the wild to reach unsustainable levels. Within various
fetish markets, dozens of shells of Kinixys and whole
animals kept in corrals and pens are available for purchase.
2
Locals will cut an L-shaped notch into the tortoises’ shell
for ease of tying string for transport from the field. Because
of the damage to the shell and reduction in aesthetic appeal
it is likely these individuals are not intended for the pet trade. In a study by Luiselli et al.
(2013), comparisons of bushmeat market data revealed a significant decline in the overall
number of Kinixys specimens likely indicating a decrease in the availability of tortoises in the
wild. Testimony from locals near the Kibale National Park in Uganda have reported increased
hardship finding resources such as wildlife in the area and many are concerned with shortages
in the near future (Hartter 2010).
The reptile pet trade has increased substantially in the past fifteen years with millions of reptiles
bought, sold, and traded annually around the world (Laidlaw 2005). Recent restrictions on
wild bird exportations have likely contributed to this increase as collectors have been forced
to shift to more exploitable species (Milledge 2000). While a majority of reptiles are collected
Burden Ltd ,Japan
16
Eric Fouchard
1
1. K. homeana for sale
at a Japanese pet show.
All Kinixys species are
available for sale in
Japan and the market
for these species in
Asia is growing.
2. K. homeana ready
for export in Togo to
the United States. K.
homeana is by far the
most heavily exported
Kinixys from Africa.
Many of these animals
will die in pet shows
or shortly after due
to improper care and
husbandry.
2
from Africa, Asia, and Latin America, the primary markets for
these animals are in North America, Europe, and Japan (Laidlaw
2005). Although this spectrum of the pet trade may be legal,
regulations in native range countries are often loose and animals
are known to be held and shipped in unsanitary and inhumane
conditions without food, water, or appropriate environmental
conditions. Recently at a Johannesburg airport in South Africa, a
large shipment of 1,600 amphibians and reptiles was discovered;
nearly 400 of these animals had already died due to dehydration
and improper shipping (BBC News 2014). This shipment of
animals was a legal export destined for the United States and
cases such as this although less frequent than historically, are
not uncommon. Though legal exportation of animals produces
large revenues across the globe, the illegal pet trade industry
generates substantially more. Interpol has recently estimated
illegal animal trade to be worth more than $10 billion annually
which is second only to the drug industry (Laidlaw 2005). A
large proportion of illegally obtained wildlife and their parts
(e.g., elephant tusk, rhinoceros horn) are brought into Asia to
feed an enormous demand for animal products used for food
and medicinal purposes. There has also been a historically high
demand for chelonian species and products within Asia and
it is very likely that Kinixys are highly valued (Laidlaw 2005).
Within these cultures, the rarer a species is, the higher its value.
This unfortunate belief leads collectors to aggressively search for
species that are rapidly declining in the wild. Corruption, weak
judicial systems, and light sentences are allowing dangerous
international crime networks to exploit natural resources
throughout Africa at alarming rates.
Historically Hingeback tortoises have been common in the pet trade industry. Prices vary greatly
for tortoises depending on the location, species, and rarity. Recently Kinixys have been priced at
over $150 each, with some species selling for up to $1000. K. homeana is the exception to this rule
with prices that have been low historically; however, due to increased difficulty in collection, the
prices have increased. In the United States K. nogueyi, which has been banned from importation
since 2001, sells for $300 to $800 per animal although exporters from Togo list these at $10
apiece. In Europe an adult K. erosa typically sells for $350-500 and pair of K. homeana can sell for
up to $500. As it is anticipated that the status of Kinixys species will be increased to a minimum of
Vulnerable with others elevated to Endangered, it is likely that the demand and therefore prices of
these tortoises will increase significantly.
17
Species Summaries
Jegler Herder - digitalnature.org
John Lewis
18
2. Introduction
Kinixys homeana
Home’s Hingeback Tortoise
Toby Hibbitts
Description
1
K. homeana reaches up to 220 mm in length. The overall coloration of
the shell is typically dark with shades of brown, red, and yellow. The
centers of the scutes remain lighter. The serrated carapace is flattened
dorsally and angled sharply downwards in the back, giving a squared
appearance from the side. The plastron is yellow with black markings
in the center of the scutes, notched in the front and projects slightly
beyond the carapace. Broadley (1989) notes the head and limbs are
typically pale yellow, though the head can range in color from dull
brown to orange.
Richard Bartlett
Geographic Range and Distribution
1-2. The color patterns
of K. homeana’s head and
limbs can vary greatly as
seen between these two
individuals.
This species is known from Liberia, Ivory Coast, Ghana, Togo,
Benin Republic, Nigeria, Cameroon, Equatorial Guinea, Gabon,
and Democratic Republic of the Congo. There is some uncertainty
regarding the easternmost records, particularly those from Democratic
Republic of the Congo.
2
Liberia: In the west portion of Liberia, the potential range of K.
homeana includes the eastern regions of Bomi, Gbarpolu, and Lofa
and is limited to more coastal states near the southeastern portion of
the country.
Ivory Coast: The range is limited to the southern portion of the country within the coastal regions
of Bas-Sassandra, Sud-Bandama, Lagunes, Agnebi, and Sud-Comoe. The species has been recorded
in all of the forested regions of this range, for instance around Aboisso (Iverson1992).
Ghana: Historically found in the southern region of this country (Iverson 1992) and more recent
19
observations include the Muni-Pomadze Lagoon near Krua (Raxworthy and Attuquayefio 2000,
Luiselli and Diagne 2013). Multiple specimens were observed by Luiselli et al. (2003) in markets
within the states of Greater Accra and Eastern with very few specimens found in the field, indicating
that occurrence in this country is becoming uncommon.
Togo: The potential range includes the hilly montane forests bordering Ghana in the Plateaux Region.
Localities have recently been recorded near Badou, Kpalimé, and Tomegbe (Luiselli and Diagne
2013). Historically this species may have occurred across much of southern Togo, though absence of
vouchers and significant land use changes limit our ability to accurately predict historic range.
Benin: This species potentially occurs in the southeastern tip of the country. Observations were
recently recorded in Oueme near the Nigerian border. Specimens are also often found in bushmeat
markets. Similarly to Togo, prior to land alteration and collection pressure this species may have
occurred over a larger area of the country.
Nigeria: Found only in southern regions of the country (Iverson 1992). Long-term studies by
Luiselli et al. (2000, 2003) and Luiselli (2003a, 2003b, 2003c, 2005) have revealed much about the
distribution within this area of its range with some of the results recorded below by state.
20
Kinixys homeana
Size
Male: 211mm
Female: 223mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Flattened dorsally, serrated scutes at
rear; dark in color, scutes may have
lighter centers
Plastron
Front lobe thick and notched, may
project slightly beyond carapace;
yellow or darker with yellow along the
margins
Hinge
Well developed, between 7th and 8th
marginal
Beak
Unicuspid and not serrated; moderately
to strongly hooked; slightly upturned
nose
21
Lagos: Assumed to be rare in this small state, specimens have
been found only in fragments of forest and bush near Maryland
and Lekki Lagoon.
Edo: Common in mature forest patches but likely extirpated
from recently deforested areas.
1
Delta: Severe habitat loss and active hunting has left the species
very endangered in this state. Once found along the Etiope River
in swamp forests in 1982, the population has since declined and
no tortoises have been observed recently (Lea et. al 2003).
Richard Bartlett
Rivers and Bayelsa: Observed in swamp forest patches
throughout both of these states (Powell 1993, Luiselli et al.
2000) and locally common around communities where it is
worshipped (Luiselli 2003b).
2
Anambra: Extremely rare in this state and found only in the
southernmost region within fragments of riverine forest. The
state is dominated by agriculture leaving little to no suitable
habitat for this species.
John Lewis
Akwa-Ibom: Populations are fragmented in this state and found
in a few riverine forests along the River Kwa-Ibo and in the
Stubbs Creek Forest Reserve. Many specimens are traded in the
Uyo market however it appears that a majority of them come
from neighboring Cross River State (Eniang unpublished data).
Habitat loss near the coast due to oil production expansion is
severely threatening populations in the state. The species is more
common in areas bordering Cross River State.
3
1. Typical forest habitat associated with K. homeana near
Kpalimé, Togo.
2. Captive specimen showing variability in shell colors.
3. Posterior view of K. homeana showing pronounced break
(hinge) and dorsally flattened, sharply angled carapace
characteristic of this species, giving a squared appearance
from the side.
Cross River: Common in lowland forests such as the Ikpan
Forest
Equatorial Guinea: The potential range of K. homeana extends
throughout the country. Specimens have been found on Bioko
Island and in Rio Muni (Iverson 1992).
Cameroon: Known occurrences exist throughout coastal regions
and also in the southern province (Iverson 1992). Distribution is
well documented in the southwest region of the country (Lawson
1993, 2000, 2001; Chirio and Lebreton 2007). Presence has
22
been recorded at multiple sites including, Abat, Mundemba,
Baro, Besingi, Ekundu Kundu, Pamol Palm Plantations,
Nfainchang, Bipindi, Dibongo, Jengwe, Kribi, Déhané,
Nkolembembe, Nko’olong, and Akom II (Lawson 1993, van
Dijk 1994, LeBreton 1999).
Tomas Diagne
Gabon: No locality data is available for this region. Potential
range likely occurs along the Equatorial Guinea border.
1
Democratic Republic of the Congo: Reported to only occur
in isolated populations in the country’s easternmost forests
that lie close to political borders with western Uganda and
Rwanda (Iverson 1992).
Tomas Diagne
Habitat and Ecology:
1. Plastron view of a pair
of K. homeana. Male (left)
with concave plastron
and long tail. Female
(right) with flat plastron
and comparatively
shorter tail (not visible).
2. Anterior view of K.
homeana highlighting the
elongated plastron and
carapace which provides
additional protection to
the tortoise.
The habitat preferences of K. homeana populations are
known largely from anecdotal data. This species prefers moist
forests that provide closed canopies and shady microhabitats
(Luiselli 2005); however it has also been associated with
streams and swampy areas (Branch 2008, Maran and Serpol
2006). Research in the Niger River Delta in southern Nigeria
has shown populations of K. homeana to be habitat specialists
which are associated with mature secondary dry forests
around mangroves and mature secondary swamp forests
(Luiselli et al. 2000). It is reported to inhabit forests that
are drier than those inhabited by the similar species K. erosa;
however in Cameroon and Nigeria the opposite is typically
2
observed (Ernst and Barbour 1989, Chirio and LeBreton
2007, Luiselli and Diagne 2013). Overlap in range between
the species may be a limiting factor in habitat occupancy due
to interspecific competition for local resources (Luiselli 2003a). Studies have shown that males
tend to be highly territorial, driving out other males from home ranges that may exceed 0.5 km2
(Lawson 2000). These results indicate that the density of K. homeana is likely to be low even in
areas with highly suitable habitat. Within the eastern region of the Niger Delta, Luiselli (2003b)
demonstrated the type of treatment K. homeana receives by indigenous people may play a large role
in habitat selection. In communities that regard K. homeana as a holy animal, individuals can be
found in diverse habitats such as dense brush, sparse brush, swamps, and plantations. In contrast,
in communities where K. homeana is actively hunted, observations are limited to dense, brushy
areas.
This species is primarily more active during the wet season and are known to use floods as a
way to disperse by floating rather than swimming. In areas where seasonal flooding occurs, this
type of movement is common (Luiselli et al. unpublished observations). These tortoises are
23
generally crepuscular regarding their daily activities, preferring
cooler times during dawn and dusk. In captivity this species
has also been reported as nocturnally active. K. homeana has an
omnivorous diet that includes plant matter, seeds, fungi, worms,
snails, slugs, and various arthropods (Luiselli 2006a). This diet
composition can vary slightly between habitat type and with
seasonal availability. K. homeana is also known to forage on
small amphibians and carrion (Branch 2008) and has a more
carnivorous diet when compared to other Testudinidae species
(Luiselli 2006c). These tortoises forage for food by following
olfactory paths with straight, darting jabs of their heads (Eglis
1962). Although there is little data on the predators of this
species, it is likely that hatchlings are vulnerable to a variety of
larger taxa and their heavily serrated marginal scutes are likely
an adaptation to discourage predation. Adults and juveniles are
vulnerable to animals such as mongooses, leopards, primates,
and birds of prey.
Tomas Diagne
1
1. Typical mixed use tropical forest with cultivated and
native vegetation. This secondary forest is habitat to K.
homeana.
2. Dense cover provides additional protection to these
cryptic tortoises.
2
In the wild mating has been reported from mid-March to end of
May in southern Nigeria (Luiselli and Akani unpublished data).
Females oviposit one to two times per year, typically between
December and January which falls in the dry season (Blackwell
1968, and George 1994). In captivity multiple clutching is
known for this species supporting observations in the field.
Wild collected specimens exported from Togo to the U.S. from
August through October 2013 were gravid upon arrival. The
highest proportion of gravid females was detected in October
shipments. These findings indicated that the potential nesting
season may extend from August through January and possibly
year-round, similar to K. erosa.
Eggs are brittle, oval to nearly spherical in shape, and measure
approximately 46 x 35 mm (Branch 2008). Nests contain 2-4
eggs and are typically made in forest litter and excavated burrows
(Blackwell 1968). Incubation time in the wild is unknown;
however, in captivity eggs will hatch 89-102 days after being
incubated at 28-30°C (Koehler 1997). Hatchlings generally
measure about 42-47 mm (Branch 2008) but are known to
reach up to 50 mm (Farkas and Satorhelyi 2006). Some level
of diapause in egg development is suspected for many Kinixys
including K. homeana. Developmental cues and triggers likely
are influenced by humidity and temperature.
24
1. K. homeana for sale
at wholesale facility in
Togo. This country is
the largest exporter of
K. homeana in the world.
2. Dead K. homeana for
sale in a fetish market in
Lomé, Togo. Tortoise
shells and heads are
believed to have medical
benefits as part of local
customs and beliefs.
Population Status and
Threats:
There is wide evidence to support
that this species is declining over its
entire range (Branch 2008, Chirio and
LeBreton 2007, Luiselli et al. 2006). In
southern Nigeria research has examined
the difference in tortoise density between
areas where the animals are worshipped
versus those where they are actively
1
hunted. Density ranged from 0.15 to
0.9 tortoises per ha in harvested areas
and from 1.65 to 2.85 tortoises per ha in areas where the species is worshipped (Luiselli 2003b).
Interviews of local hunters selling the tortoises in bushmeat markets have also found a decline
(Luiselli et al. 2013). Population sizes of K. homeana are likely reducing especially in areas where
these tortoises are actively hunted.
K. homeana populations have appreciably declined due in large part to habitat loss and collection.
Deforestation is occurring rapidly in West Africa for timber, mining, conversion to agriculture,
and industrial expansion. These destructive activities present a significant threat for the loss of high
quality forested habitat that this species relies on. Nigeria is the dominant oil-producing country
in Africa and as a result has experienced prolific deforestation with extensive losses occurring in the
Niger Delta where a large number of K. homeana historically occurred (Luiselli and Akani 2003).
Hunting by locals interested in selling or consuming the tortoises is also a factor in population
declines (Lawson 2000, Luiselli 2003c). Kinixys are sold in markets
throughout Africa and exported internationally for the pet trade,
consumption, and medicinal purposes (Luiselli 2003b). Review
of CITES export data for Ghana, Togo and Benin from 1975 to
2012 showed a reported export of nearly 80,000 K. homeana most
of which occurred since the mid 1990’s (CITES Trade Database
2012). Numerous surveys of bushmeat markets in the Niger Delta
and Togo conducted by Luiselli et al. (2003b) over multiple years
revealed significant decline in the number of K. homeana and other
hingeback tortoise specimens for sale, indicating that the numbers
of wild individuals are declining. K. homeana can be found in more
diverse, open habitats in regions where the tortoise is worshipped
compared to areas of active hunting where they occupy habitat
dominated by thick brushy vegetation where presumably detection
by humans is less. It is likely that the species has been extirpated
from areas where they are easily captured.
2
25
Will Ahrens
Kurt Edwards
Regulation and Management:
1
2
Beyond standard habitat management in protected forests,
there are no known specific conservation actions currently being
taken to protect this species. In Togo, the species is protected
under Article 62, Section 2, Chapter II of law N° 2008-005,
regulating environment protection and wildlife conservation
and in Nigeria exports are currently not allowed. Observations
of trade data trends have shown when one country closes
export or reduces quotas the exportation shifts to an adjacent
country. This is most notable in Ghana and Benin, both major
exporting countries for Kinixys. Application of regulation
on the ground requires enforcement and illegal hunting of
tortoises even in protected areas is widespread through West
Africa (Lawson 2000). Even if not directly hunted on protected
lands, this species faces unexpected challenges of habitat
loss. Near Kpalimé at a protected Nature Reserve, an area of
previously intact tropical forest in 2005 was by 2013 converted
into agricultural lands and inhabited by multiple families. The
landscape no longer supported conditions suitable to support
K. homeana.
K. homeana is currently listed on CITES appendix II, which
restricts the legal exporting of specimens, presumably limiting
the harvest to the point that a wild population should be
sustainable, though within-country collection and use may
be unregulated. Until recently, IUCN Red List described this
species as Vulnerable. In August 2013, a Red List assessment
focused on sub-Saharan African chelonians resulted in the
elevation of K. homeana to the draft designation of Critically
Endangered. At the national and subnational level, it is
necessary to include this species among the protected fauna in
all the countries of occurrence. At present, it appears that there
is virtually no country that can reliably preserve this species.
Further legislation should also be developed to protect this
species at the international level.
1. Hundreds of K. homeana are imported into the U.S.
annually. Many die from improper care and stress though
some make their way into the hands of competent keepers.
3
2. Some keepers and institutions in the U.S. and Europe have
been successful in keeping and breeding this species.
3. Conversion of habitat is a major threat to K. homeana.
Terry E. Kilgore
26
2. Introduction
Kinixys erosa
Forest Hingeback Tortoise, Serrated Hingeback Tortoise, Eroded Hinged Tortoise, Schweigger’s Tortoise
David Korte
Description
1
K. erosa is the largest of the Hingeback tortoises reaching lengths of up
to 400mm. The marginal scutes are noticeably serrated, a feature that is
unique to this species. The carapace slopes forward dorsally and ranges
from orange-yellow to gray-brown with costal scutes that occasionally
have contrasting markings. The plastron projects slightly beyond the
carapace and is black with yellow on the outer margins and along
sutures. Broadley (1989) remarks that the head and limbs area typically
yellow though they can vary from brown to orange depending on the
individual.
Tomas Diagne
Geographic Range and Distribution
1. K. erosa displaying its
adaptive color pattern on
the floor of a west Africa
tropical forest.
2. Large K. erosa rescued
from a village where it
was likely heading to the
bushmeat market.
This species is known or presumed to occur from Senegal, Gambia,
Guinea-Bissau, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana,
Togo, Benin, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo,
Central African Republic, Democratic Republic of the Congo, Angola,
and Uganda.
2
Gambia: Observations are reported by Andersson (1937), Hakansson
(1981) and Iverson (1992) and this species may occur throughout the
country.
Senegal: Observations are not reported for this region and distribution is unknown. Suitable
habitat for this species is limited to the extreme southern coastal region.
Guinea-Bissau: There is no locality information reported from this region. The species may
potentially occur throughout the country.
27
Guinea: Little locality information is known for this country and
the range of the species is likely limited to the western and coastal
regions of the country between Guinea-Bissau and Sierra Leone.
Sierra Leone: K. erosa potentially occurs throughout the country;
however due to a large amount of deforestation in this area, it is
likely that remaining habitat is limited. Observations have been
reported in the southeast region.
1
1. Deforestation and
conversion of tropical
forest is a significant
stressor to K. erosa.
Incompatible land use
practices will continue
to negatively impact this
species.
Liberia: There are individuals found throughout the country and
Iverson (1992) lists reports from both the Northwest and Southeast
provinces. Unpublished data also reports finding individuals in
bushmeat markets in Monrovia. Buchanan, and Gbarnga in 2000.
Ivory Coast: Here the range is limited to the southern portion of the country within the coastal
regions of Moyen-Cavally, Bas-Sassandra, Sud-Bandama, Lagunes, Agnebi, and Sud-Comoe.
Observations have been reported from the forests around Abidjan in Lagunes from Iverson (1992)
and Maran (2004).
28
Kinixys erosa
Size
Male: 323mm
Female: 260mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Slopes dorsally, scutes flat, serrated
marginals, scarecely to moderately
notched nuchal region; dark in color
with lighter scute centers
Plastron
Front lobe thick and notched, projects
beyond carapace, black with yellow on
outer margins and along sutures
Hinge
marginal
Beak
Unicuspid and not serrated
Matt Muir
29
Ghana: Iverson (1992) has reported K. erosa to have a wide
distribution through southern and central territories, though the
current potential range is restricted to the southern region of the
country. Individuals have been found by Raxworthy and Attuquayefio
(2000) in two sites (Onyadze and Ony-Winn) of Muni-Pomadze
Lagoon in the southern region. Additional specimens were observed
in the south around Cape Coast and researchers have reported that
many tortoises are traded for food in local markets especially near
Accra and Koforidua.
1
Togo: Found only in Southwestern Togo in remnant forests around
the towns of Kpalime and Badou (Harris 2001). It is rare to find
wild specimens in the few forest habitats where the tortoise occurs;
however, many specimens can be found in bushmeat and fetish
markets (Luiselli and Segniagbeto unpublished data, Segniagbeto
et al. 2013).
Bruce G. Marcot
Benin: The species has likely been extirpated from much of this
country, however they are known from a restricted range along the
southern region of the country near the Nigerian border (Trape et
al. 2012).
Olivier Pauwels
2
1. A dorsal view of K.
erosa showing its plastron
protruding past the
carapace.
2. Several K. erosa for sale
at a bushmeat market in
Equatorial Guinea.
3. Newborn K. erosa,
showing the egg tooth,
in Gamba, OgooueMaritime Province,
southwestern Gabon.
Nigeria: Distribution is well documented in this country because of
numerous studies conducted in the region with some of the results
recorded below by state (Luiselli et al. 2000, 2003. Luiselli 2003a,
2003b, 2003c, 2005). Current range is restricted to the southern
region of the country.
Lagos: Presumed to be rare, few individuals were found in fragments
of forest and bush around the coastal lagoons.
Oyo: Considered very rare in this area, it was found in fragmented
forests around Ibadan where it is actively hunted. Its presence and
3
exploitation has also been recorded by Butler and Shitu (1985).
Edo: Presumed to be extirpated from recently deforested areas.
Potential range includes the southwestern region. Studies from 1999 to 2005 have verified its
presence in only 3 of 19 potential sites of occurrence.
Delta: The species occurs in this area however severe habitat loss and subsistence hunting has left it
very endangered here. The potential range is currently limited to the southern half of the state.
30
Jean Francoi Trape
1
1.Potential habitat of
K. erosa in the Plateaux
Region of Togo.
2. Young K. erosa from
Bopolu, Liberia with
heavily serrated marginal
scutes--a likely defense
against predation.
Bayelsa and Rivers: Found in swamp forests throughout
both states. In some areas where local communities worship
the tortoise, observations are common. Active hunting for
food and medicine coupled with habitat loss is likely causing
the species to decline in these regions (Luiselli 2003b)
Anambra: believed to be very rare in this state and found only
in a few riverine forest patches in the southernmost region
surrounding Oguta. Much of the state is used for agriculture
leaving very little suitable habitat for the tortoise.
Akwa-Ibom: The species is found in a few riverine forests
that are severely fragmented and in the Stubbs Creek Forest
Reserve. Many specimens have been found for trade in Uyo
markets; however, a large portion of these are likely to be
from the adjacent state, Cross River. Habitat loss due to oil
production expansion near the coast threatens the tortoise
in this region. The species is most common at the border of
Cross River where some sacred forests are protected.
Cross River: Presumed rare in lowland forests where the
similar K. homeana is more abundant, K. erosa is considered
by some to be common and even abundant in some hilly
and mountainous areas of these regions. Within the Cross
River National Park, K. erosa is the more common hingeback
tortoise in the Akamkpa and Okwangwo divisions of the
park (Eniang, Ebin, Luiselli unpublished data).
Cameroon: Found throughout the coastal, southern, and
southwestern regions of the country (Chirio and LeBreton
2
2007, Iverson 1992, Lawson 1993, 2000, 2001). Lebrenton
(1999) reports observing specimens at the following sites:
Bipindi, mouth of the Doumé River, Campo, Efulen, Lolodorf, Metet, Sakpabayémé, Sangmelima,
Yaoundé, Kribi, Ngam, Foulassi, Nkoétyé, Djomedio, Dja River Faunal Reserve, and additionally,
Takamanda Forest, where it is actively hunted (LeBreton et al. 2003). Lawson (2000) has reported
very strong hunting pressure in western and south western regions.
Equatorial Guinea: The potential range of K. erosa extends throughout the country. Iverson (1992)
reported observations in Rio Muni. This tortoise is often found in bush-meat markets here (Fa et
al. 2000, Fa and Garcia Yuste 2001).
31
Gabon: Maran (2006) reported it as the most commonly observed chelonian
species in the country. Observations were made in 86 locations throughout the
entire country including all the dense forest types.
Congo: Potential range currently extends throughout the country with the
exception of Brazzaville, Pool, and southern portions of Plateaux.
2
1. K. erosa are known to be
semiaquatic, spending considerable
amounts of time in water
foraging, traveling and even
mating. Maintaining clean, clear,
watercourses with canopy cover is
essential in providing the critical
habitat feature for this declining
species.
2. Mushrooms are an important
food for Kinixys. Seasonally they
make up a significant portion of a
tortoise’s diet.
Democratic Republic of the Congo: A majority of the species’ potential range
here is located within the northern and central regions of the country, with a
small band extending to the southern border through Kasai Occidental. Most
of the locality data has been reported in the northern region (Iverson 1992).
Angola: Here the range is limited to extreme northern regions of the country.
Observations were reported from Posto de Lóvua, Perto de Rio Camualua (Dundo),
North Lunda, and Cabinda. A photographic record of an individual traded in a market
in Cambulo (Luiselli 2005) indicates that the species is hunted in this area.
Central African Republic: K. erosa likely occurs in the extreme southwest portion of
the country with the majority in Sangha-Mbrare and the extreme south western tip of
Mambere-Kadei.
Uganda: Found through the central portion of the country with a limited range in the
state Eastern. Absence of observation from well-preserved forested habitats suggests
that this species is uncommon here (Vonesh 2001) or perhaps significantly collected.
1
32
Habitat and Ecology:
Cris Hagen
2. Though typically
crepuscular or even
nocturnal, K. erosa can
with time become
more active during
the day in captivity.
Russ Mittermier
1. K. erosa are known
to use rivers and
streams as corridors
and will float or swim
between habitats. As
logging, agriculture,
and mining
operations continue,
these habitats
become degraded
and unsuitable for
supporting this
unique tortoise
species.
The habitat preferences of K. erosa
include lowland evergreen forests,
marshy areas, and gallery forest
along rivers and streams (Lawson
1993). Research in the Niger
River Delta in southern Nigeria
have shown populations of K.
erosa to be habitat specialists and
associated with mature or secondary swamp forests (Luiselli et al.
2000). Although this species
1
typically inhabits dense forests,
individuals can be found crossing adjacent habitats such as savannah to reach nearby forested zones (Maran 2006). Reports of K.
erosa using watercourses as travel corridors have also been noted (Lawson 1993). Within the eastern
region of the Niger Delta where communities treat K. erosa as a holy animal, the tortoise can be
found in more diverse habitats such as dense brush, sparse brush, swamps, and plantations. In
contrast, in communities where K. erosa is actively hunted, observations are limited to dense brushy
areas. This use of secondary or cultivated lands may indicate that this species, when not threatened
by hunting can be somewhat adaptive and use more disturbed habitats. Home range and site fidelity of K. erosa are not well understood.
The range of K. erosa is sympatric with K. homeana increasing the potential for interspecific
competition due to similar diet composition, habitat, and geographic range (Lawson 2006, Luiselli
2008). The density of K. erosa is likely low in even in areas with highly suitable habitat due to the
territorial nature of these tortoises. Studies have shown a high degree of competition between males
who will force others out from their home ranges that may exceed 0.5 km2 (Lawson 2000, 2006).
Research in Cameroon indicates that home ranges between all
specimens have considerable overlap; however, analysis reveals
little overlap occurs in the core areas that tortoises occupy the
most (Lawson 2006). On a local scale, competition between
the tortoises results in one species that is dominant over the
other (Luiselli 2006c, 2008). K. erosa tend to inhabit forests
that are wetter than those inhabited by K. homeana (Ernst
and Barbour 1989) except in Nigeria where the opposite is
generally observed (Luiselli and Diagne 2013). Analyses of
the diets of these two species suggests that at least within
parts of their range, K. homeana is more of a generalist than
K. erosa (Luiselli 2003a, 2003b). Within heavily altered
mature forests, the diet composition of the two species
becomes more similar.
2
Olivier Pauwels
Gabriel Segniagbeto
33
1 Habitat loss displaces
Kinixys increasing their
exposure to collection.
2. Live adult K. erosa sold
as food along the road
in Kougouleu, Estuaire
Province, western
Gabon.
1
K. erosa is active primarily in the wet season with limited activity
during the dry months. It is crepuscular with most activity occuring
during cooler times of the day, such as early morning or late
afternoon. During periods of prolonged rain the tortoise may be
active at any time of day. Throughout their inactive period during
the dry season, individuals will often spend several consecutive
days without moving from their refugia, which are beneath tree
roots or under leaf litter. This tortoise can be cryptic and elusive,
preferring shaded microhabitats, though it will occasionally bask
(Luisellie 2003a, Maran 2006, Naulleau, 1988). Observations
have been reported in both the U.S. and Europe of this species
showing activity in temperatures as low as 50° F. K. erosa has an
omnivorous diet and will eat earthworms, snails, arthropods, fish,
carrion, plants, seeds, fruits, and fungi. Mushrooms are generally
the preferred type of food (Luiselli 2006a). These individuals are
hosts to many parasites in the wild (Bouamer and Morand 2007)
and leeches are often found on them. Little is known about the
natural predators of this species. It is likely that hatchling tortoises
are vulnerable to a host of species, though sharp, serrated marginal
scutes deter many predators from swallowing them whole. Juveniles
and adults are likely vulnerable to mongooses, leopards, primates
and birds of prey.
Data on the reproductive biology of this species in the wild and
captivity is limited. In Gabon, nesting is observed throughout
the year and females with eggs have been observed in August,
November, December, February, and March with a peak in
March, indicating potential for multiple clutches, which is further
supported by observations of captive animals. The eggs are hard
shelled, slightly flat, white, and measure 38-45 mm in length, 3439 mm in width, 31-34 mm in height, and 27-30 g in weight
(Maran, 2006). Clutches with up to 10 eggs have been reported
though in captivity typical clutches are between 6 and 8 eggs.
2
Population Status and Threats:
Currently, no extensive data is available for population size of
this species, but it believed that populations are declining throughout most of its range based on
collection pressure and habitat loss. In some portions of its range, such as Liberia and Gabon, this
species appears to be more stable, though increase in mining and logging, and shifting demand
for tortoises in food markets and the pet trade may impact these populations as well. In southern
Nigeria research has examined the difference in tortoise density between areas where the animals
are worshipped versus those that where they are actively hunted. Density ranged from 0 to 0.35
tortoises per ha in harvested areas and from 0.40 to 1.10 tortoises per ha in areas where the species
34
is worshipped Luiselli (2003). In Cameroon the estimated mean annual harvest rate is 0.7 to 0.8
animals per km2 in the sites with high habitat quality and low human densities (Lawson 2000).
Populations of K. erosa are likely declining in areas where the tortoises are under pressure due to
hunting.
2. Multiple K. erosa
shells at a fetish market
Lomé, Togo.
Hunting by locals living in desperately poor
conditions is a factor in population declines (Lawson
2000, Luiselli 2003c). Although this species is known
to be valued as a holy animal in some communities
1
of the Niger Delta, its use as a source of food and
income in much of this area is increasing. Tortoises
are sold in markets throughout Africa and imported internationally for the pet trade, consumption,
and medicinal purposes (Ceballos and Fitzgerald 2004,). According to the CITES trade database,
reported exports from range countries of K. erosa from 1976 to 2012 was over 9,100 animals. Of
that total, over 2,400 individuals were exported between 2000 and 2012 (CITES Trade Database
2012). Though these numbers are lower in comparison to
other species in this genus, these also do not account for
the significant number of animals that end up in bushmeat
markets. Surveys of bushmeat markets in the Niger Delta
and Togo conducted by Luiselli et al. (2008) over an
extended period of time revealed significant decline in the
number of K. erosa and other hingeback tortoise specimens
for sale, indicating numbers of wild individuals are rapidly
declining. Research has shown that K. erosa can be found
in more diverse, open habitats in regions where the tortoise
is not actively hunted compared to areas of active hunting
where they are found in dense brushy areas that are hard
for humans to reach. It is likely that the species has been
extirpated from areas where they are easily captured.
2
This reveals that in areas where K. erosa is threatened by
Tomas Diagne
1. Tortoises and other
wild meats can serve
as a significant source
of protein for local
people. At one time
their consumption
may have had minimal
impact on the overall
populations. With
increased habitat loss
and growing human
densities this practice
now has a greater effect
on population stability.
K. erosa populations are declining throughout much
of its range, due to multiple factors including habitat
loss and persecution by local people. Deforestation is
occurring rapidly in West Africa for timber, mining,
conversion to agriculture and industrial expansion.
These destructive activities present a significant threat
for the loss of high quality forested habitat that this
species relies on. K. erosa is considered to be a habitat
specialist because it prefers mature or secondary
swamp forests. These habitat types within the Delta
states have dramatically dropped from 2% of state area
in 1976/78 to 0.2% in 1993/5 (Geomatics 1996).
35
interaction with humans, it occupies only a small portion of suitable
habitat that may be available (Luiselli 2003b).
Kelly Hull
Beyond limited protection of this species critical habitat, there are no
specific conservation actions that are currently being taken to protect
K. erosa. The species is currently listed on CITES appendix II, which
restricts the legal exporting of specimens, presumably limiting the
harvest to the point that a wild population should be sustainable,
though within-country collection and use may be unregulated.
Further legislation should be developed to protect this species on
both international and national levels. Until recently, IUCN Red
List described this species as Data Deficient. In August 2013, a Red
List assessment focused on sub-Saharan African chelonians resulted
in the elevation of K. erosa to the draft designation of Endangered. In
order to protect it on a national level, this species should be included
1
among the protected fauna in each country of occurrence. In Togo,
the species is protected under Article 62, Section 2, Chapter II of
the law N° 2008-005, regulating the environment protection and
wildlife conservation and in Nigeria exports are also not allowed. However, at this time there is no
country that can protect the species dependably and researchers have witnessed specimens being
traded for food or medicine even within protected areas (e.g., Upper Orashi Forest Reserve in Rivers
State, southeastern Nigeria). Another problem at the national level is the excessive exportation rate of
individuals within Africa to markets in western countries such as Ghana, Benin, and Togo (Lawson
2000). This trade needs to be controlled to ensure that it is not being supplemented with free-ranging
individuals. It is usually unclear whether these specimens were captured from the wild because tortoise
farms are generally underdeveloped in these areas (Harris 2001).
1. Captive breeding
via assurance colonies
both in situ and ex situ
will be an important
tool in conserving
Kinixys.
2. An increasingly
rare site to see in west
Africa: a young K. erosa
found in Tai National
Park, Ivory Coast
Alex Krohn
2
1
2. Introduction
Kinixys belliana
Bell’s Hingeback Tortoise
1. K. belliana from
Yabello, Ethiopia
traveling. This species
has the largest, most
diverse range, occuring in
a wide variety of habitats.
2. Ornately patterned
K. belliana from Uganda.
Though the most
commonly known, K.
belliana is one of the least
studied Kinixys. Their
status throughout Africa
is unknown.
Description:
This species can reach up to 230mm in length. The carapace has convex scutes, giving the shell
a domed appearance in contrast with the square shape of K. homeana and K. erosa. The carapace
has a radiated pattern with color ranging from beige to brown with surrounding darker circles
and seams that are cream or whitish. The plastron can vary from uniformly light in color to
darker with black markings that radiate outwards. Coloration and patterns can vary greatly
between individuals (Broadley 1989).
Geographic Range and Distribution:
This species is known to occur in Cameroon, Central African Republic, South Sudan, Sudan,
Eritrea, Ethiopia, Somalia, Kenya, Uganda, Rwanda, Democratic Republic of the Congo, and
Angola and may occur but it is not confirmed in Burundi, Djibouti, and Tanzania. Predicting the
range and distribution of K. belliana is problematic due to a lack of verified locality information.
The range map below was created based on available data. A potential alternate or additional
corridor where this species may occur extends from Central African Republic to Angola were
animals may travel down the Congo River system (D. Broadley, personal communication, April 7,
2014).
Cameroon: Locality information is not available for this
country. Species may potentially occur in a small region
extending through northern portions of East and Central
states.
Mark Stanton
Central African Republic: The range is restricted to the
southern provinces of the country. Observations have been
reported in Ouaka and Bangui.
2
South Sudan: The potential range of this species spans the
entire country with the exception of the northern region of
Western Bahr el Ghazal. Specimens have been reported in
Western Equatoria, and Central Equatoria.
Trevor and Margaret Hardaker
36
37
Sudan: The species range in this country is limited to the southeastern
states of the country. Observations have been reported in South
Kordofan and Khartoum.
Vladimír Trailin
Eritrea: The species potential range is limited to the southern regions of
the country.
1. K. belliana from Centrální
Etiopie, Langano, Rift
Valley, Ethiopia. This
species has considerable
color variability and is often
confused with other species
of Kinixys.
2
Ethiopia: Range includes western and central regions of the country
with a large gap where the Ethiopian Highlands occur. Observations
have been reported in Oromia near the Eastern Rift Valley, and in the
Southern Nations, Nationalities, and People’s Region division.
Somalia: The potential range is limited to areas in the northwest and
southwest regions of the country.
Kenya: Within this country, a large portion of the species’ potential range occurs along the
western border extending as far south as Nairobi and east to Meru. The range potentially
extends from the southern tip of Somalia into the states of Garissa, Tana River, and
Kilifi. It is not likely that the species is found in central portions of the country. Specimen
observations have been recorded around the Winam Gulf near Lake Victoria and Tana River
near the southern coast (Spawls et al. 2002).
?
38
Kinixys belliana
Size
190-220mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Convex, scutes raised; radiated pattern
on carapace, areolae brown with light
ring surrounding
Plastron
Truncated and usually protrudes beyond
carapace; black splashes radiating
outward or uniform light color
Hinge
marginal
Beak
Unicuspid and not serrated; rarely
hooked
39
Uganda: The species is likely to occur throughout the entire
country. Observations have been recorded from Eastern,
Northern, and Western Regions (Spawls et al. 2002).
Jaime Traynor
V.Socha
Rwanda: This species is predicted to occur throughout the
country. Previously a record has been reported from Akagera
(Spawls et al. 2002).
1. K. belliana from
Welkite, Gurage zone,
in Western Ethiopia.
Historically (and as
recently as the 1990s),
upwards of 20% of this
region was covered in
forested habitat, though
much of it has since
been developed.
2. Grassland habitat
in Uganda. These
grasslands, although
ideal for K. belliana, can
serve as a population
sink when these areas are
aggressively burned.
1
Democratic Republic of the Congo: A majority of
the locality records within this country have occurred in
Orientale and a limited number are from within Bandundu
and Equator. The potential range of K. belliana within this
country includes a small area located in the northern tip
of Equator, which expands into the northern and eastern
borders of Orientale. Its range continues along the eastern
borders of North and South Kivu where a small corridor
begins to span across the country beginning in southern
Maniema and extending through the southern regions of
Kasai Oriental and Kasai Occidental. Distribution continues
through the southern and central portions of Bandundu
where it extends slightly into Equator near Lac Tumba.
Angola: The range occurs through the northern region
of the country and ends near the southwestern coast.
Observations have been reported in the states of Namibe,
Huila, and Bie.
2
Djibouti: Potential range of the species occurs throughout
the country; however no specific locality data is currently
available.
Tanzania: Although this species has not been recorded in this country, it is likely to occur in the
northernmost region surrounding Lake Victoria.
Burundi: No locality data is currently available for this country; however it is likely that this
species occurs in the northern tips of the country, north of Lake Tanganyika.
Habitat and Ecology Information:
Within its range K. belliana prefers to inhabit areas that are sandy with savannah, dry bush, or
grassland (Bonin et al. 2006). This tortoise is crepuscular and most active in the early morning
or late afternoon during the coolest times of the day. Activity peaks for this species during the
wet season while they are known to estivate during the dry season. During periods of heavy rain,
the tortoise is known to use waterways to disperse by floating. Unlike most tortoise species, this
species is omnivorous and prefers to eat a variable diet of plant matter, fungi, invertebrates, and
is also known to feed on small animal carcasses when encountered (Broadley 1989).
Nesting is known to occur from November to April and females typically lay 2-7 eggs in a single
clutch though more than one clutch may be laid per year. Data on reproduction of K. Belliana
Tomas Diagne
40
1
is limited. This species likely exhibits similar breeding and nesting
behaviours to other dry grassland species. Presumably, incubation
may take up to a year for hatchlings to emerge based on the
ecology of southern African species (Boycott and Bourquin 2000)
. Populations located further north in the continent may exhibit
incubation periods more similar to species such as K. nogueyi. There
is little information regarding known predators of this species. It
is likely that hatchlings are prey to a variety of taxa including birds,
snakes, and rodents while the less vulnerable juveniles and adults
are likely to be killed by larger animals such as mongooses, leopards,
primates, and ground hornbills.
Though longevity data is limited for Kinixys, it is presumed that this
species may live a minimum of 40 to 50 years. Numerous natural
predators and pressure from collection by humans likely limits the
potential longevity of many animals, though in captivity, adults are
known to live for over 30 years (Branch 2008, Harris 2001).
Vladimír Trailin
Patti Murray
1. K. belliana habitat.
2. K. belliana can be
distinguished from other
Kinixys by its highly
domed shell.
3. A young adult
specimen of K. belliana
from Addis Ababa,
Ethiopia.
At this time there is no quantitative data on the size of K. belliana
populations. This species’ current range is uncertain and abundance
of the tortoise within its known range is not well understood. Available
2
data does support that K. belliana populations are declining throughout
much of its range, likely due to multiple factors including habitat loss
and collection by local people for food, use as traditional cowbells,
sale in fetish markets and the pet trade. Deforestation and clearing of
grassland habitat is occurring rapidly in many parts of Africa for timber
harvest for charcoal and lumber, agriculture, mining and industry.
Increase in conversion of grassland to agriculture and the common
use of fire to prep fields may have a significant negative impact on
K. belliana populations and population viability. These destructive
activities present a significant threat for the loss of high quality habitat.
Increase in roads and road traffic also likely has a deleterious effect
on tortoise populations and impact habitat connectivity. Additionally,
electric fences are commonly used throughout Africa and are known
3
to cause direct mortality to tortoises (Beck 2010). Kinixys are sold in
markets throughout Africa and exported internationally for the pet
trade, consumption, and medicinal purposes (Ceballos and Fitzgerald
2004). Exact reported trade data on K. belliana is difficult to interpret because until recently multiple
species were lumped into the K. belliana complex and CITES trade data did not differentiate which
subspecies were exported. In addition, Kinixys have been sold internationally since at least the 1960’s
in large quantities with shipments of hundreds to thousands of animals. Like other Kinxys, the rate
of loss from various anthropogenic sources is presumed to be negatively impacting the stability
of K. belliana populations and their ability for recovery, adaption, and long-term viability. Habitat
loss and landscape management especially the extent to which fire is used are likely the largest
direct threats. It is probable that a majority of collection has historically been localized for cultural
and opportunistic consumption. Increasing presence by foreign nationals, particularly from China
is driving the demand for chelonians including Kinixys for food and creating a black market for
41
collection of these and other declining species
(BBC News 2012). This has been reported for
multiple countries where Kinixys is found and the
trend is likely to increase without intervention.
Trevor and Margaret Hardaker
Within the range of K. belliana, critical habitat
protection is limited, and there are no specific
conservation actions that are currently
being taken to protect K. belliana outside of
restrictions on export. Throughout a majority
of its range, this species is likely in decline
and immediate conservation actions are
needed to prevent further population decline
or eventual collapse. Seasonal use of fire as
a management strategy likely kills numerous
K. belliana annually. Evaluating management
to time burning to avoid tortoises or manage
through alternative means is recommended.
Enacting stronger regulations to protect
1
Kinixys within countries and providing
subsequent enforcement of regulations will
be important in helping reduce collection
pressure. Establishing protected areas especially in regions
were multiple species overlap is critical in providing longterm conservation of species.
1. Tortoise shell cowbells
up for auction. The
specimen on the right
is a Leopard Tortoise,
while on the left is K.
belliana. These cowbells
are still used today by
local people, especially in
Ethiopia.
2. K. belliana from
Yabello, Ethiopia.
K. belliana is currently listed on CITES appendix II,
which restricts the legal exporting of specimens, presumably
limiting the harvest to the point that a wild population
should be sustainable, though within-country collection
and use may be unregulated. Since 2001 this species and
associated subspecies have been banned from importation
to the United States based on the potential concern of the
tortoise being a vector for heart water disease, a tick-born
2
bacterial disease which affects cattle (Kaplan 2014, U.S.
Department of Agriculture 2000). The status of this species
was recently assessed at the IUCN Red List assessment
focused on sub-Saharan African chelonians in August of 2013 which resulted in its elevation
to the draft designation of Vulnerable. Further legislation should be developed to protect this
species on both international and national levels. Future research should focus on the range
and distribution of this species due to the current lack of locality data with effort to also
gather more information regarding population size and age structure. In order to protect it
on a national level, this species should be included among the protected fauna in each country
of occurrence. Currently, to our knowledge as the time of publishing, no research is being
conducted on K. belliana.
42
Kinixys nogueyi
2. Introduction
Western Hingeback Tortoise
1. K. nogueyi found near
Kpalimé Togo. Note
the four toes visible, a
diagnostic trait of this
species.
2. A large adult from
Sabodala, Senegal.
Description:
This species is unique among Kinixys tortoises in having only four claws on each front limb. Its
domed shell is similar in appearance to K. belliana. The carapace varies between patterns with
areolae that are uniformly brown or yellow with dark borders. The plastron is uniformly colored
or sometimes has dark blotches (Broadley 1989).
It is known to occur in Senegal, Gambia, Guinea, Guinea-Bissau, Sierra Leone, Ivory Coast, Mali,
Burkina Faso, Ghana, Togo, Benin, Nigeria, Cameroon, and Central African Republic.
Russ Mittermier
Senegal: Here the range is limited to the south with a majority of the
provinces Louga and Matam excluded.
Gambia: No locality information is currently available for this country
however it is likely that the range of this species extends throughout the
country.
1
Guinea-Bissau: Potential range of the species occurs throughout the
country; however no locality data is currently available.
Jean Francoi Trape
Guinea: Found throughout the country with the exception of the
southern most regions past the Guinea Highlands.
Sierra Leone: Potential range extends the width of the country but
probably not into the southern region.
2
Mali: Here the range extends through the southwestern region of the
country and ends near the southern border of the Mopti province.
43
Burkina Faso: The species is believed to occur throughout the country
with the exception of the Sahel Province.
Ghana: Found throughout the country with a small area of absence in
the southwest portion of the Western province. Observations have been
reported in Central, Eastern, and Northern Provinces.
1
1. Grasslands make
up a major portion
of K. nogueyi habitat,
though they can also be
found in forests living
sympatrically with K.
erosa and K. homeana.
Togo: Here the range is believed to extend throughout the country.
Benin: Potential range of the species extends throughout the country;
however no locality data is currently available.
Nigeria: Range extends throughout the central regions of the country and is believed to be
absent from the northernmost provinces and coastal region of the Niger Delta.
Cameroon: Range extends through the entire country with the exception of the Extreme North,
the southern half of East, and South regions.
44
Size
Kinixys nogueyi
200-300mm
Digits
Forelimb: 4
Hindlimb: 4
Carapace
Domed carapace which is uniformly
brown or with yellow areolae with a
dark border
Plastron
Irregular or uniform dark blotches
Hinge
marginal
Beak
Unicuspid and not serrated; Upper
jawline is indented in front of the eye
45
Central African Republic: Here the range extends through
the western half of the country with the exception of SanghaMbare and southern Lobaye.
Tomas Diagne
Tomas Diagne
1. K. nogueyi in Benin.
2. Various K. nogueyi
showing incredible color
variation.
1
Within its range K. nogueyi prefers to inhabit areas that are
open with savannah, dry bush, or grassland (Ernst and Barbour
1989, Luiselli 2003a, 2003c). It is sympatric with K. homeana
and K. erosa within much of its range; however it is the least
observed of the three species likely due to habitat preference
(Luiselli 2003a). Additionally, K. nogueyi is sympatric with K.
belliana in a small range in Cameroon and Central African
Republic where there is the potential for the two species to
intergrade. This tortoise is crepuscular and most active in the
early morning or late afternoon during the coolest times of
the day. Activity peaks for this species during the wet season
while it is known to estivate during the dry season. Like other
members of Kinixys this species is omnivorous and prefer to
eat a variable diet of plant matter, fungi, invertebrates, and is
also known to feed on carrion when encountered (Bonin et
al. 2006).
Information on this tortoise in the wild is limited and gathering
data can be difficult due to its small size, and cryptic coloration
and behaviour. Captive populations have given some insight
into the life history of the tortoise. Males, especially during the
mating season, are very aggressive and territorial with other
males. Observations of captive males showed intense and
prolonged battles resulting in multiple wounds to the limbs
2
and head. This dominance has been documented in other
Kinixys both in captivity and the wild (Lawson 2000, 2006,
Mifsud unpublished data). In captivity, the majority of mating behaviour is observed between
June and November but it is also known to occur periodically at other times of the year (Morris
1994). A captive male that had no contact with other Kinixys for approximately 20 years initiated
courtship almost immediately after being introduced to a group, and mating was observed year
round (Mifsud unpublished data). Captive females typically nest from November to January and
will display restless behaviour in weeks leading up to nesting season (Morris 1994). In captivity,
this species produces clutches of up to 6 eggs and can lay multiple clutches per season. Like other
Kinixys this species does have a developmental diapause lasting roughly 30 days in captivity. Work
has not been conducted to evaluate the natural history, reproduction, and development of this
species in the wild.
46
2. Young adult male
K. nogueyi in a village
near Kpalimé, Togo.
Locals collect these
animals to supply the
pet trade.
3. Kinixys in west
Africa are often
collected when locals
are out looking for
snails.
4. Roads serve as
a form of habitat
fragmentation as
well as a source
for collection and
loss due to vehicle
encounters.
Currently there is no comprehensive data on the
population size of K. nogueyi. Abundance of the
species within its range is not well understood
as it has been only locally studied as a secondary
component to other work. Within limited
portions of the K. nogueyi range, this species
exhibits very low population densities of 0.1 to
1.4 individuals per ha (Luiselli 2003c, 2006a,
2008). Effort should be made to better assess
population size and health throughout the species
range.
1
A combination of habitat loss and collection
by local people is likely the leading cause of K.
nogueyi population declines. Deforestation and
clearing of grassland habitat is occurring rapidly
in many parts of this species range for timber
harvest and charcoal, mining, and industry.
Increase in conversion of grassland to agriculture
and the common use of fire to prep fields may
have a significant negative impact on K. nogueyi
populations and population viability. These
destructive activities present a significant threat
2
for the loss of high quality habitat. Tortoises
are sold in markets throughout West Africa and
collected for the pet trade, consumption, and
cultural/medicinal purposes (Luiselli 2003a).
Historically thousands were imported into the
United States, Europe, and Japan and today many
are still collected and sold for the international pet
trade. Based on trade data for reporting countries,
over 85,000 K. nogueyi were reported exported
between 1975 to 2012. The major exporting
countries of K.
nogueyi include
3
Togo, Benin,
and Ghana
representing greater than 95% of all exportation of this species
(CITES Trade Database 2012). Since 2001, this species (formally
a subspecies of K. belliana) has been banned from importation
into the United States due to potential threats to U.S. cattle
industry associated with heart water disease (Kaplan 2014, U.S.
Department of Agriculture 2000). Based on trade data this species
is still imported into Europe and demand is growing in Asia,
particularly Japan and China.
4
Eric Fourchard
1. K.. nogueyi for sale
at a Lomé exporter.
Note the incredible
color variability in
this species, similar
to the variability of
patterns found in the
North American Box
Turtle, Terrepene.
47
1. Conversion of
grassland to agriculture
and aggressive use of
fire negatively impact K.
nogueyi populations.
2. Captive bred K.
nogueyi displaying color
variability even as very
young animals.
3. This young female
K. nogueyi was collected
in a small village near
Kpalimé Togo and
offered for sale.
2
Beyond limited habitat protection of this species, there are no specific conservation actions that
are currently being taken to protect K. nogueyi. Throughout a majority of its range, this species
is likely in decline and immediate conservation actions are needed to prevent further population
decline or eventual collapse.
K. nogueyi is currently listed on CITES appendix II, which restricts the legal exporting
of specimens, presumably limiting the harvest to the point that a wild population should be
sustainable, though within-country collection and use may be unregulated. Further legislation
should be developed to protect this species on both international and national levels. The species
was recently elevated from data deficient to the draft designation of Vulnerable in August 2013
following an IUCN Red List assessment targeting sub-Saharan chelonians. Effort needs to be
made to get a better census on population size, age structure, and distribution locally by country
and rangewide. In order to protect it on a national level, this species should be included among
the protected fauna in each country of occurrence.
Currently, very limited effort to conduct research on K.
nogueyi is under way. Future research should be focused on
assessing populations, monitoring habitat loss and how these
changes in the environment affect local populations. More
ecological studies in the wild are encouraged to gather a
better understanding of the species’ behavior in the natural
environment.
3
Will Ahrens
1
48
Kinixys spekii
2. Introduction
Speke’s Hingeback Tortoise
1.Large female K.
spekii from Limpopo,
South Africa, showing
its characteristic flat
carapace.
Description
Richard Hay
The potential range of this species includes Angola, Namibia, Botswana,
Zambia, Democratic Republic of the Congo, Zimbabwe, South Africa,
Swaziland, Malawi, Mozambique, Tanzania, Burundi, and Kenya.
2
Kenya: Here the potential range extends through the southeastern
provinces of the country. It is believed to occur from Narok to
Laikipia and south to the coast from Kwale to Kilifi. Specimens have
been recorded in the southern tip of the country and isolated records
have been reported from the Kerio Valley.
Victor Loehr
This Hingeback tortoise is a medium-sized
species reported to reach up to 200mm in
length. The marginal scutes are not serrated
and the carapacial hinge is more developed
when compared to other species. The overall
2. K. spekii in veld near
coloration is typically brown or yellow
Nelspruit, South Africa.
becoming darker and often redder with
Note that color can
age. The shell lacks distinguishing markings
vary even among adult
and individuals can vary greatly within the
females, from heavily
1
patterned to solid brown. species (Broadley 1989). Females of this
species typically grow larger than males
(Coulson and Hailey 2001). A captive female
specimen in the U.S. measures 202mm and has shown new growth
indicating the maximum size may yet to be determined (Mifsud
unpublished data).
49
Ryan van Huyssteen
Tanzania: Range extends through the central provinces of the
country, south of Lake Victoria and north of Morogoro. Most of
the observations have been reported from the northern region within
Kigoma, Rukwa, Shinyanga, Arusha, and Manyara.
Burundi: The potential range here extends along the southern portion
of the country and the species is believed be absent north of Lake
Tanganyika.
1
1. Juvenile K. spekii, Masebe Nature Reserve, South
Africa.
Democratic Republic of the Congo: Range is believed to be
limited to the southeastern regions of the country within Katanga and
portions of South Kivu, Maniema, and Kasai Oriental. Observations
have been reported in Katanga and South Kivu.
Zambia: Range extends throughout the country, with observations
concentrated along the eastern and southern borders.
50
Kinixys spekii
Size
Male: 181mm
Female: 200mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Flattened, scutes relatively flat with
well-defined annuli; Male: uniform
color, Female: uniform to short
radiations
Plastron
Front lobe thick and projects well
beyond carapace, concave in males
Hinge
marginal
Beak
Unicuspid and not serrated; not or
weakly hooked
51
Bernard Dupont
Malawi: Found throughout the country with the exception of the
southeastern border near Lake Chilwa.
Botswana: Here the range is potentially limited to the
northeastern region of the country where a majority of
observations have been from North West and Central Provinces.
1
Namibia: The potential range in this country is limited to the
small segment of land that lies between Angola, Botswana, and
Zambia including Caprivi and portions of Kavango.
Curtis Hart
Zimbabwe: Found throughout most of the country, many of the
observations are from the Sengwa Wildlife Research Area (SWRA)
in the central Gokwe District. Its range is also known to extend
into Manicaland near Odzi.
2
Mozambique: Limited to the western portion of the country
between Malawi and Zimbabwe and a small region bordering
South Africa in Gaza Province. Observations have been reported
from the Tete Province.
Adriane Ohanesian
South Africa: Potential range extends into the northeast of the
country entering Limpopo and Mpumalanga
1. Roads are likely a
significant threat to K.
spekii. As road density
and traffic increases
this may impact tortoise
density.
2. K.spekii emerging after
rain in Kruger National
Park, South Africa.
Swaziland: Here the range is believed to cover the northern tip of
the country, primarily spanning the Hhohho province along with
the northernmost portions of Lebombo and Manzini Provinces.
In eastern Africa, K. spekii prefers to inhabit savannahs and coastal
thickets, often in rocky areas (Spawls et al. 2002). In the southern
region of the continent, it utilizes moist savannah woodlands and
drier woodlands farther north in its range. At the Sengwa Wildlfie Research Area (SWRA) in
Zimbabwe, Hailey and Coulson (1995) recorded K. spekii using 13 of the 16 vegetation types
observed in the park. The greatest number of specimens was found within various types of
mixed woodland, grassland, and wooded grassland. The range of K. spekii overlaps with all
members of the genus except K. nogueyi, K. homeana, and K. erosa (sensu Kindler et al. 2012).
3
K. spekii is a crepuscular species that uses the early morning and late afternoon hours to forage
while seeking refuge from the heat during the middle of the day (Spawls et al. 2002). This
species often finds shelter in rock crevices (facilitated by the flattened carapace) and horizontal
3. Typical savanna habitat tree trunks (D. Broadley, personal communication, April 7, 2014) During the dry season, the
for K. spekii.
species spends most of its time estivating under cover. Hailey and Loveridge (1997) notes that K.
52
spekii may be dormant for up to 8 months during the winter
and the following dry season. Activity peaks during the wet
season in which tortoises will move from dense woodlands
into woodland/grassland areas that provide more food
(Scoones 1986). Studies by Hailey and Coulson (1995) at the
SWRA, Zimbabwe showed the home range of this species
to be 3.1 ha. Data from the same study has shown that the
tortoise prefers to use mammal burrows as a form of cover
but are also known to use felled trees and dense shrub/
thickets. These refuges serve as protection from predation as
well as the environment.
K. spekii has an omnivorous diet with vascular plants making
up less than half of its food intake, part of which is fruit
rather than foliage. Fungi and invertebrates make up the rest
of the variable diet (Hailey et al. 1997). Scoones (1986) notes
1
that observations of the tortoises are often associated with
1.Mushrooms make up
the presence of mushrooms which are the preferred food.
a seasonally significant Natural predators of this tortoise include a variety of mammals and birds. A study by Coulson and
portion of this species’
Hailey (2001) investigated the mortality of K. spekii due to predation within undisturbed habitat.
diet. Alteration to
landscapes or changes They demonstrated that at least 77% of specimens found dead within the study area were killed
by predators. A significant source of this predation is caused by large birds such as the ground
in precipitation may
affect food availability. hornbill.
2. Moist savanna
woodlands are an
important habitat for
K. spekii.
2
Ryan van Huyssteen
3
Liana May
3. K. spekii resting in
the shade of a shrub
in Masebe Nature
Reserve, South Africa.
This species displays sexual dimorphism with
females generally growing larger than the
males. Average age at sexual maturity is around
9 years for females and 7 years in males
(Coulson and Hailey 2001). In the southern
part of the range, K. spekii has been observed
nesting from November to April. 2 to 3 eggs
are typically laid at once, however clutches with
as many as 6 eggs are known to occur. The
tortoise can lay more than one clutch within
a year and larger females generally produce
clutches with more eggs (Hailey and Coulson
1997, Boycott and Bourquin 2000). Nesting
was observed by Hailey and Coulson (1997) in
Zimbabwe where all successful nests were built
within 30 m2 of bushes and/or small trees, all
under tree canopies. Nests were built during
late afternoon and evening. In captivity nesting
has been observed from November through
April. Incubation duration of eggs varies, but
53
ranges between 7 and 9 months, possibly up to a year. Hatchling
growth is fast and within a month can grow from 30 to 50 mm
to approximately 75 mm within 3 months. Though this may in
part be due to diet, it is likely based on the shorter active seasons
that K. spekii may grow quickly.
David Beadle
1
Currently there is no extensive data on the population size
of the species. Similar to other Kinixys species, gathering
information on this tortoise can be difficult due to its small size,
and cryptic coloration and behaviour. Because it is seasonally
inactive, it can be hard to find in the wild which may help this
species remain more common even in densely populated areas
(Hailey and Coulsen 1999). Lambiris et al. (1989) and Coulson
and Hailey (2001) performed mark-recapture studies in separate
research areas in Zimbabwe (the Boulton Atlantica Research
Station and the Sengwa Wildlife Research Area) and calculated
the density of local populations to be 2.25 per ha and 0.16 per
ha, respectively. The large contrast in these figures may be due in
part to the difference in abundance of predators between these
two sites, with the SWRA having more natural predators present.
Both of these sites are located in reserves where wildlife is
protected thus it can be assumed that outside of these locations,
densities of K. spekii populations are lower due to increased
threat of habitat loss and persecution by humans.
2. Dead K. spekii from
Selati, South Africa. This
species has numerous
natural predators,
athough this animal
appears to have been hit
by a vehicle.
This tortoise is collected from the wild by humans for consumption and trade. Locally, the
tortoise is traded for food and traditional medicines where the meat is consumed and the shell is
often used for tools. There are reports of this species in turtle markets in Hong Kong however it
is only sold in the pet trade whereas other Kinixys species are included in the food trade which is
ten times larger (Cheung and Dudgeon 2006). This species is sought after and has also appeared
on multiple Japanese and U.S. websites for sale as a pet selling for hundreds of U.S. dollars per
Ryan van Huyssteen
1. A hinged carapace
is unique to the genus
Kinixys.
K. spekii populations are likely declining throughout its range, in
part due to extensive habitat loss that is occurring. Deforestation
has dramatically increased in Africa recently due to agriculture
and industrial expansion. Climate change causes habitat
modification which could affect the presence of this species in
parts of the native range. K. spekii was identified by Erasmus et
al. (2002) as susceptible to a range shift and contraction in South
2
Africa after global warming creates an environment with mean
2°C temperature increase and increasing aridity. However, South
Africa is at the edge of the species range, thus the extent of potential change in this area is not
likely to represent that of the entire range.
Brian Horne
54
1. K. spekii has been
imported by the
thousands into the U.S.
Trade data only dates
back to 2006, though
this species has been
consistently imported
since at least 2003. Most
die from improper
care. Establishment
of assurance colonies
will be one tool to help
prevent the loss of this
species and increase our
understanding of its
needs.
2-3. Beautifully patterned
K. spekii.
animal. Trade numbers are challenging due to the relatively
recent recognition of K. spekii as a full species. Based on
data from 2007 through 2012 over 1,500 K. spekii have
been exported from Africa with nearly 80% exported from
Mozambique (CITES Trade Database 2012). Another
hazard that it faces is mortality due to man-made structures
such as electric game fences. Literature indicated that
Kinixys species are known to be killed on occasion by these
fences which also act as barriers for traveling animals (Beck
2010).
1
Currently this species is protected in many game reserves
and national parks throughout its range (Caro et al. 2011)
and is listed as present in 16 major reserves throughout
southern Africa (Branch et al. 1995). Beyond this standard management in protected areas,
there are no specific conservation actions that are being taken to further protect the species and
poaching likely still happens even within protected areas.
K. spekii is currently listed on CITES appendix II, which restricts the legal exporting of
specimens, presumably limiting the harvest to the point that a wild population should be
sustainable, though within-country collection and use may be unregulated. The species was
recently elevated from Data Deficient to the draft designation of Vulnerable in August 2013
following an IUCN Red List assessment targeting sub-Saharan chelonians. Further legislation
should be developed to protect this species on both international and national levels. Effort
needs to be made to get a better census on population size, age structure, and distribution locally
by country and range-wide. In order to protect it on a national level, this species should be
included among the protected fauna in each country of occurrence.
The ecology of K. spekii has been studied in detail in Zimbabwe and to a lesser extent in other
countries. Great effort must be made to expand research into other countries within the species
range. At this time there is no known effort to conduct new research with this tortoise.
3
Ozan oanguen
2
55
Kinixys zombensis
2. Introduction
Southeastern or Radiate Hingeback Tortoise, Madagascan Hingeback Tortoise
Jo-anne Hounsom
Description:
K. zombensis zombensis
K. z. zombensis reaches up to 220mm in length. The carapace is
domed with sloping sides and flattened dorsally with broad black
radial patterns. The color patterns can vary between individuals from
brown to yellow. The plastron is very dark in juveniles and lightens
in color with age. (Broadley 1989).
1
K. zombensis domerguei
The subspecies K. z. domerguei is extremely similar in appearance
to K. z. zombensis and can only be distinguished from the mainland
relative by a fragmented frontal scale.
Thomas Leuteritz
2
1. K. z. zombensis within the Imfolozi Game Reserve,
Zululand Nata. As this species ages, it tends to develop a
lumpy or pyramided appearance.
2. K. z. domerguei on Nosy Faly island. This endemic
form of Kinixys on Madagascar closely resembles K. z.
zombensis. The origins of the Malagasy population are
uncertain, but it has been a part of Madagascar’s biota
for thousands of years.
The mainland subspecies, K. z. zombensis is known to occur in Kenya,
Tanzania, Malawi, Mozambique, Zimbabwe, and South Africa while
the Madagascan Hingeback Tortoise (K. z. domerguei) occurs on Nosy
Faly, a small island off the northwest coast of mainland Madagascar
and the adjacent peninsula.
Kenya: Range is limited to the southeastern corner of the country
within Kilifi, Mombasa, and Kwale. Isolated records are reported
from the Taita Hills (Spawls et al. 2002).
Tanzania: Range extends from the far northeast of the country
in Tanga region and continues south expanding as far west as the
Dodoma, Iringa, and Ruvuma regions. Spawls et al. (2002) reports
observations are in Dar es Salaam, the eastern Udzungwa Mountains
and isolated records are reported from the Rondo Plateau.
56
Mozambique: The species range occurs throughout the country with
the exception of the Tete Province and the western portion of the Gaza
Province.
Craig Gibbon
Malawi: Here the species is known to be widespread in the southeastern
region of the country (D. Broadley, Personal Communication, April 7,
2014).
Zimbabwe: The potential range covers a small portion of the eastern
region of the country including Manicaland, Mashonaland East, and
Masvingo.
1
1. K. zombensis at False Bay in Kwazulu Natal near
the town of Hluhluwe, South Africa
South Africa: Range extends into the far northeastern region of the
KwaZulu-Natal Province and specimens have been reported to occur in
St. Lucia Park.
Madagascar: The subspecies K. z. domerguei is endemic to Madagascar.
Currently the range is presumed to be limited to Nosy Faly, a small island
off the northwest coast of mainland Madagascar, as well as the adjacent
peninsula. The largest density is found in the central and western regions
of Nosy Faly (Pedrano 2008).
57
Thomas Leuteritz
Data on the ecology of the K. zombensis complex is lacking likely due in
part to its recent taxonomic elevation. As a result, much of the general
information on this tortoise is from research focusing on the K. belliana
complex which included K. z. zombensis and K. z. domerguei as subspecies.
1
1. As K. z. domerguei age their colors tend to
fade, as shown in this individual. This tortoise is
culturally significant to the people of the region as
it is considered a guardian of spirits.
K. z. zombensis prefer to inhabit moist savannah woodlands, coastal
grasslands, and dune forest edge (Branch 2008, Broadley 1989, Pedrano
2008). K. z. domerguei have been observed utilizing secondary forest as
well as coconut plantations, crops fields, and villages where people do not
consume them (Pedrano 2008). These tortoises tend to be crepuscular and
are most active in the early morning or late afternoon during the coolest
times of the day (Spawls et al. 2002). Activity peaks for these species during
the wet season while they are known to aestivate during the dry season
using animal burrows, dead logs, and leaf litter (Branch 2008, Bonin et al.
2006).
58
Kinixys z. zombensis
Size
Male: 206mm
Female: 217mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Domed carapace with a broad black
radial pattern
Plastron
Front lobe truncated and extends to
or projects beyond carapace; uniform
light color or black splashes radiating
outward
Hinge
marginal
Beak
Unicuspid and not serrated
59
Kinixys z. domerguei
Size
Male: 194mm
Female: 196mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Flat dorsal surface, with unflared
marginal; variable in color from yellow
to brown, darker radiated pattern in
adults
Plastron
Front lobe truncated and projects
beyond carapace; uniform light color or
black splashes radiating outward
Hinge
marginal
Beak
Unicuspid; frontal scale is fragmented
and the width of the frontals is less than
the width of the prefrontals
60
The breeding patterns of these species are poorly
known. Mating is observed from November to
April. Multiple clutches are laid with 2 to 10 eggs
in each clutch. Incubation times for individuals
within mainland Africa are typically 90-110
days (Branch 2008). The period of incubation
for individuals in Madagascar is unknown but
3
believed to be around 200 days (Pedrano 2008).
Tortoises become sexually mature between 8 and
11 years old. Like other Kinixys longevity is unknown but speculated based
on captive animals to be over 50 years.
Andy Dasypeltis
Data on the population of this species is lacking in part due to the recent
change in taxonomic status. Based on studies and observations of other
grassland Kinixys, it is presumed that density of individuals is rather low.
Work should be conducted to better assess population structure and
density to better assess overall population health.
4
Although data regarding the current number of K.z.zombensis is limited,
the population is likely declining throughout much of its range due to
multiple factors including habitat loss and collection by local people.
Deforestation and clearing of grassland habitat is occurring rapidly
in many parts of Africa for timber harvest, mining and industry.
Increase in conversion of grassland to agriculture and the common
Margaret Hardaker
4. Older and lightly
patterned K. z. zombensis
on a road in Hluhluwe
Umfolozi Park, South
Africa.
2
Thomas Leuteritz
particularly vulnerable
to road-related mortality
after rain events as they
drink water from road
puddles.
1
Thomas Leuteritz
Like other species of the genus, members of the K.
zombensis complex are omnivorous and prefer to eat
a variable diet of plant matter, fungi, invertebrates,
and are also known to feed on small animal
carcasses when encountered (Broadley 1989,
Bonin et al. 2006, Branch 2008, Pedrano 2008).
Limited information is available regarding natural
predators of this species. Common predators of
other Kinixys species include a variety of mammals
2. K. z. domerguei hiding
and birds. Large birds such as the ground hornbill
in vegetation. To date, no
comprehensive study has are common predators of many tortoise species
and are known predators of Kinixys (Bonin et al.
looked at this endemic
2006, Coulson and Hailey 2001). On Nosy Faly,
Malagasy Kinixys.
the main predator of the tortoise observed was
3. Female K. z. zombensis
the African Bush Pig (Potamochoerus larvatus),
near a pool of water
however the tortoise population does not appear
along a road in Mkuze
to suffer significantly from this predation (Pedrano
Game Reserve, South
2008). Feral dogs also are known to predate young
Africa. Kinixys are
tortoises (Pedrano 2008).
1. Typical habitat where
K. z. domerguei can be
observed. Clearing of
habitat for agriculture
and seasonal burning
negatively impacts this
species across its small
geographical range.
Megan English
61
1. Roads serve as
physical breaks in the
landscape, increasing
exposure to predators
and vehicle-related
mortality. Opportunistic
collection and targeted
poaching can often occur
in conjunction with other
illegal wildlife trade.
2. Conversion of habitat
into agricultural land has
a significant impact on K.
zombensis. Depending on
the crop, tortoises may
be killed as pests as they
may feed on the plants,
flowers, or fruits.
1
more each wholesale .
use of fire to prep fields may have a significant negative impact
on K. zombensis populations and population viability. These
destructive activities present a significant threat for the loss of
high quality habitat (Broadley 1989). Kinixys are sold in markets
throughout Africa and exported internationally for the pet trade,
consumption, and medicinal purposes. Large numbers of K.z.
zombensis are exported annually for the international pet trade,
though these numbers likely represent a small percentage of
the total number consumed for food. Most of the exports of
this species come from Mozambique. Over 10,000 grassland
Kinixys were exported between 1990 and 2012 with Kenya,
Tanzania and Mozambique being the highest percent of exports
(CITES Trade Database 2012). This data presumably represents
K zombensis based on the range of exporters though some
individuals may represent K. spekii prior to CITES recognition
of K. spekii as a full species and its trade data separation in 1993.
Because of bans on importation into the U.S. limited numbers
of individuals are currently known and when found they will
sell for $800 to $1,000US for a female. In Europe where
importation is currently allowed, they will sell for $150US or
The subspecies K. z. domerguei is especially at risk due to the extremely small range it inhabits.
Within this range, bush fires are a common practice used for agriculture and land clearing.
This method is known to be a direct cause of mortality and areas that are frequently burned
currently support very few or no individuals. Within the restricted range, as little as 500 hectares
of suitable habitat remains (Pedrano 2008). Fortunately for the tortoise, in the Ambanja region
of Nosy Faly locals believe them to guard royal spirits and do not kill them directly. However,
in other regions of the island with
different culture the tortoises are not
worshipped and often are collected for
consumption (Pedrano 2008).
2
Megan English
It is estimated that on Nosy Faly less
than 5,000 K. z. domerguei individuals
remain giving this area the largest
density of the these tortoises in the
world (Pedrano 2008). Based on
Cities Trade data nearly 350 were
exported over a two year period with
20 animals heading to the United
States in 2000 and 328 destined for
Japan and China in 2001 (CITES
Trade Database 2012). This coincides
with the mass collection and export
of Pyxis tortoises from Madagascar.
62
When found on a Japanese price list this Malagasy endemic form
was being offered for $1,400US. The status of those imported to
the U.S. is unknown.
Will Ahrens
Jacob Mueti Ngwava
Regulations and Management:
1. Noticeable break
(hinge) of K. z. zombensis
on this specimen from
Kenya.
2. Though once
imported in large
numbers, K. z. zombensis
is rarely found in U.S.
private or institutional
collections due to
concerns of potential
disease transmission
to cattle via ticks. It is
imperative to establish
assurance colonies by
consolidating remaining
captive animals before
they succumb to old
age. This species is also
relatively uncommon in
European collections.
1
Within the range of K. zombensis, critical habitat protection is
limited, and there are no specific conservation actions that are
currently being taken to protect it outside of restrictions on export.
Throughout a majority of its range, this species is likely in decline
and immediate conservation actions are needed to prevent further
population decline or eventual collapse. Seasonal use of fire as
a management technique likely kills numerous K. z. zombensis
and K. z.domerguei annually. Evaluating management to time
burning to avoid tortoises or manage through alternative means is
recommended.
Kinixys tortoises are currently listed on CITES appendix II,
limiting the harvest to the point that a wild population should
be sustainable, though within-country collection and use may be
unregulated. Until recently, IUCN Red List described these species
as Data Deficient. In August 2013, a Redlist assessment focused
on sub-Saharan African chelonians resulted in the elevation of
2
the K. zombensis complex to the draft designation of Vulnerable.
Further legislation should be developed to protect this species on
both international and national levels. Effort needs to be made to
get a better census on population size, age structure, and distribution locally by country and
rangewide. In order to protect it on a national level, this species should be included among the
protected fauna in each country of occurrence.
Currently, no known studies on K. zombensis are under way. Work will begin in fall of 2014
to assess status, threats and distribution of K. zombensis both in mainland Africa as well as
Madasgascar. Future research of the K. zombensis complex should focus on the endemic
populations of K. z. domerguei due to a current lack of information on this vulnerable subspecies.
Victor JT Loehr
63
2. Introduction
Kinixys natalensis
Natal Hingeback Tortoise
Victor JT Loehr
Description:
1
K. natalensis is the smallest of the hingeback tortoises reaching
lengths of up to 160 mm with females being slightly larger than
males. The strongly tricuspid beak is unique to this species with all
other tortoises in the genus having unicuspid beaks. The species can
also be distinguished from other Kinixys members because it has the
least-developed hinge of the group. The carapace is primarily brown
with yellow or orange central scute markings surrounded by black.
The plastron is yellow with roughly symmetrical black markings.
With age, the color and patterns of the shell typically dull (Broadley
1989).
Richard Boycott
This species is known to occur in South Africa, Swaziland, and
Mozambique.
1. The diagnostic
tricuspid beak of a K.
natalensis from KwazuluNatal, South Africa.
2. This is the smallest of
all the Kinixys species.
South Africa: Here the species is limited to small band along the far
eastern region of the country. The range begins in the north east of
Mpumalanga and extends into Kwazulu-Natal.
2
Swaziland: Currently the range is believed to span the entirety of this
small country.
Mozambique: The range is limited to the extreme southwestern tip of the country.
This tortoise species is restricted to the inland region of south eastern Africa. Within the
KwaZulu-Natal Province they are generally absent from coastal regions preferring to inhabit
mountainous landscapes to the west and north. It occurs in rocky grasslands, savannah, and
John van der Woude
64
bushveld habitat but avoids thick forested areas and areas
of deep sand (Bourquin 2004). It is found primarily in
the Lebombo Mountain range where it has been recorded
between 50 m and 1100 m above sea level (Boycott and
Bourquin 2000). K. natalensis is sympatric with K.z.
zombensis in northern KwaZulu-Natal Province within hilly,
montane habitats (Broadley 1981, 1993). There is sympatry
between K. natalensis and K. spekii in southern Mozambique
through the Manyeleti Game Reserve (Boycott and Jacobsen
1988, Broadley 1993). This species is the only member of
the genus that has a tricuspid rather than unicuspid beak
(Broadley 1993). The tortoise is active when weather
1
conditions are overcast and humid or during and after
1. Typical habitat for K. natalensis includes bushveld communities. rain. During inactive times, it can be found seeking shelter
This species tend to occupy drier habitats than other Kinixys
under rocks and in the foothills of mountains (Boycott
species. Fires, both natural and artificially set, can have significant 2001). There is little data on the biology of K. natalensis in
impacts on populations.
the wild; however Boycott and Bourquin (2000) provide
65
Kinixys natalensis
Size
Male: 125mm
Female: 155mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Moderately convex, posterior forms
“gutter” w/slight serration, well-marked
annuli; concentric pattern on each scute
with brown areolae
Plastron
Front lobe may project slightly beyond
carapace, male asa distinct concavity;
yellow centers and margins, 2 black
rings on abdominals
Hinge
Poorly developed, between 7th and 8th
marginal
Beak
Tricuspid
Andreas Karsten
Darglec Cnservancy
66
1. Fire as a form of
management and
conversion of habitat
to agriculture both have
a negative impact on
K. natalensis and other
Kinixys species.
2. Electric fences have
been shown to result in
mortality to K. lobatsiana
and are suspected to
impact other Kinixys
including K. natalensis in
southern Africa. These
fences also serve as a
physical barrier.
information gathered from captive individuals. Mating is observed
in February and clutches of 2 eggs are laid in April. After 5 to 6
months of incubation eggs typically hatch in September. Hatchlings
have carapace length of approximately 35 mm and a mass of 8 to
10 g. The species has an omnivorous diet consisting of plants, and
invertebrates and likely fungi which are a favorite of most Kinixys.
1
Population density information regarding K. natalensis is lacking.
It is listed as near threatened in Swaziland (Monadjem et al. 2003).
The species was initially listed in the South African Red Data BookReptiles and Amphibians (Branch 1988) as rare; however over the
past two decades observations in new localities suggest that this
may not be the case across its range. Nonetheless, in Swaziland the
species is still rare. No formal data is available regarding this species
in Mozambique.
The main threats that K. natalensis faces are habitat loss,
fragmentation, and degradation. This species occupies a small
range, thus any loss in high quality habitat can potentially have a
2
large effect on nearby populations. For example, a wild population
of several hundred individuals in the southern edge of the South
African range are being threatened by a future dam that could be built in the Tugela basin
(Boycott and Bourquin 2000). Structures like this can severely limit a tortoise’s ability to disperse
between suitable habitats. Increase in conversion of land to agriculture and the common use
of fire to prep fields can have a devastating impact on populations and viability. The current
populations are somewhat fragmented and increased conversion of land to agriculture, roads,
mining will only increase the distance between populations
The harvesting of wild specimens poses a high threat to several members of the Kinixys group and
although reports of harvested K. natalensis is limited, this species is involved in the pet and likely
food trade and as demand grows this species will be more heavily collected. Multiple reports of this
species in the United States and Japanese pet trade are due in part to misidentified K. zombensis or
K. spekii juveniles. Based on the CITES trade database, between 1994 and 2008, 28 were reported
as exported from South Africa and Zambia (thought not known to be found here) and over 130 in
total have been exported or re-exported with most going to Japan (CITES Trade Database 2012).
Currently this species is protected in a number of game reserves and national parks throughout
its range including, Kruger National Park, Manyeleti Game Reserve, Mbuluzi Nature Reserve,
Mlawula Nature Reserve, Ndzindza Nature Reserve, Itala Game Reserve, Pongola Nature Reserve
and Weenen Nature Reserve. Beyond this standard management in protected areas, there are no
specific conservation actions that are being taken to further protect the species. Less than a fifth
67
Megan English
of the entire potential range of this species is protected.
1. Conversion of natural
landscapes for grazing
livestock and agriculture
reduces the amount of
available K. natalensis
habitat and reduces
population connectivity.
1
K. natalensis is currently listed on CITES appendix II,
limiting the harvest to the point that a wild population
should be sustainable, though within-country collection
and use may be unregulated. Until recently, IUCN Red
List described this species as Near Threatened. In August
2013, a Red List assessment focused on sub-Saharan African
chelonians resulted in the elevation of K. natalensis to the
draft designation of Vulnerable. Further legislation should be
developed to protect this species on both international and
national levels. Effort needs to be made to get a better census
on population size, age structure, and distribution locally by
country and range-wide. In order to protect it on a national
level, this species should be included among the protected
fauna in each country of occurrence.
At this time there is no known effort to conduct new research on K. natalensis. Monadjem et
al. (2003) had proposed to continue distribution surveys in Swaziland to determine population
density and localities. Little is known on the captive husbandry of K. natalensis and work with
captive tortoises can be beneficial for better understanding behavior of the animals while also
establishing valuable assurance colonies abroad. There are no known assurance colonies in the
2. Protecting large
United States and the number of captive animals in European collections is uncertain though
contiguous tracts of
likely small.
habitat is essential for the
long-term conservation
of this species. Adaptive
management is
important when
conserving these
and other vulnerable
Kinixys species.
Lisa Call
2
68
Kinixys lobatsiana
2. Introduction
Lobatse Hingeback Tortoise
Victor Loehr
Victor Loehr
Description:
1
2
This species is smaller than most Hingeback Tortoises but is larger
than K. natalensis and can reach lengths of 170mm. The slightly
convex carapace has sloping sides and is buff to dull yellow with
brown areolae. Irregular radiations are sometimes seen in the species
although there is a lot of variation between individuals. The plastron
extends to or beyond the carapace and is buff with a brown border
and sometimes black rays are present (Broadley 1989). Broadley
(1993) commented that South African K.lobatsiana have more distinct
markings than tortoises from south-eastern Botswana. Until 1993
this species was not recognized as a separate species from K. belliana
(Boycott and Bourquin 2000). The overall appearance of this species
is very similar to that of K. belliana with the exception of a much
flatter overall appearance (Boycott and Bourquin 2000). K. lobatsiana
is sexually dimorphic with the females reaching sizes of around 200
mm and 1.5 kg while males will only reach 170 mm in length and 820 g
(Boycott and Bourquin 2000).
.
K. lobatsiana is known to occur in Botswana and South Africa.
1. Plastral view of K.
lobatsiana showing the
serrated marginal scutes,
a diagnostic characteristic
for this grassland species.
Botswana: Here the species range is restricted to South-East District from Lobatse to Gaborone
(D. Broadley, personal communication, April 7, 2014).
2. K. lobatsiana from
Limpopo, South Africa.
South Africa: The range covers the north of North-West Province, the southwest of Limpopo,
Guateng, and the northwest of Mpumalanga. Observations have been reported from multiple
nature reserves within Ohrigstad, Loskop, and Nylsvley (Bonin et al. 2006).
69
Megan English
1. K. lobatsiana prefers drier, more rocky
terrain as shown here in this thornveld
habitat.
Within its range K. lobatsiana is found in rocky terrain that is made up
of savannah, bushveld, and thornveld habitats (Boycott and Bourquin
2000). This species appears to be absent from highveld grassland and
subtropical lowveld. K. lobatsiana occurs across a range of vegetation from
dense, short bushveld to open tree savanna though it prefers areas with
rocky hillsides and rocky ridges (Boycott & Bourquin 2000). Compared
to other members of the genus, K. lobatsiana is the most adapted to arid
regions (Branch 2008). The northern populations are sympatric with that
of K. spekii and eastern populations likely occur with K. natalensis (Boycott
and Bourquin 2000).
1
This species is most active in the summer particularly during or after
rain. During the cold season K. lobatsiana will hibernate from May to
September using old burrows of other animals as refuge (Boycott and Bourquin
2000). When the rainy season returns the tortoises will become active again. Mating
usual occurs during November to April with much of the nesting occurring in April
70
Kinixys lobatsiana
Size
Male: 162mm
Female: 167mm
Digits
Forelimb: 5
Hindlimb: 4
Carapace
Moderately convex, well-marked
annuli, buff to dull yellow with
brown areolae, sometimes irregular
radiations; posterior marginal
shields are upturned at edges
Plastron
Front lobe truncated and extends
to or beyond carapace; buff with
brown border, sometimes black
rays
Hinge
Well developed, between 7th and
8th marginal
Beak
Unicuspid and not serrated; weakly
to moderately hooked
71
1.Savannah bushveld
habitat of K. lobatsiana.
Currently there is no extensive data on the
population size of the species. Though range
data are relatively available, abundance of the
species within its range is not well understood.
Gathering information on this tortoise can
be difficult due to its small size, and cryptic
coloration and behaviour. Because it is
seasonally inactive, it can be hard to find in the
wild which may help this species remain more
common even in densely populated areas.
2
This species is threatened by impacts of agricultural conversion and use
of fires as a management tool. It is collected for the pet trade and likely
locally collected for food and medicinal/cultural purposes. Based on
reported trade data between 2008 and 2012, 51 K. lobatsiana were exported
from South Africa (CITES Trade Database 2012). Though numbers are
low, they will sell for $800 to $1,200 each when available in the U.S. and
are often presold upon arrival. The export from 2008 to 2012 has risen by
700% and with increased demand will likely continue to grow. Currently
Botswana does not allow export, however this could change in the future.
Because of its small range local events can have very large impacts on
the population. Human activities such as overgrazing and events such as
wildfires can negatively affect this tortoise (Bonin et al. 2006).
Paul Rabiega
3
1
K. lobatsiana is currently listed on CITES appendix II, which restricts
the legal exporting of specimens, presumably limiting the harvest to
the point that a wild population should be sustainable, though withincountry collection and use may be unregulated. Until recently, IUCN
Megan English
Paul Rabiega
2-3. Carapacial and
plastral view of a
juvenile K. lobatsiana
found outside of
Johannesburg, South
Africa.
(Boycott and Bourquin 2000). Nests can include
up to 6 eggs and hatching is known to occur
after 313 days. This extended incubation period
is likely due to a winter diapause (Branch 2008).
The omnivorous tortoise will feed primarily
on herbaceous plants but has been known
to eat fungi, as well as several different types
of invertebrates including snails, millipedes,
and beetles (Bonin et al. 2006, Boycott and
Bourquin 2000).
72
Will Ahrens
Red List described this species as Data Deficient. In August
2013, a Red List assessment focused on sub-Saharan African
chelonians resulted in the elevation of K. lobatsiana to the
draft designation of Vulnerable. Further legislation should be
developed to protect this species on both international and
national levels. Effort is needed to better assess population
size, age structure, and distribution locally by country and
range-wide. In order to protect it on a national level, this
species should be included among the protected fauna in each
country of occurrence.
Within the potential species range, most of the protected areas
are located within South Africa. Only one protected area is
1
located in Botswana. In total less than 25% of this species’
fragmented range includes protected lands. Viable suitable
habitat within the species’ range has been reduced over the last 20 years and continues to decline.
Little formal research has been done on K. lobatsiana though an assessment of southern Africa
including the range of K. lobatisana will be conducted in fall of 2014 to evaluate habitat use,
threats, and species range. Future research should be focused on assessing populations, monitoring
habitat loss and how these changes in the environment affect local populations.
1. Though relatively
uncommon in captivity,
this species is known
to be offered for
sale in Japan and the
U.S. Exportation has
increased in recent years
and these tortoises are
often listed for more
than $1,000 U.S.
2
SA Reptiles Forum
2. Though locally
uncommon, K. lobatsiana
has been found in
highveld grassland on
vertical rock crevices, at
elevations of 1510m.
73
Conservation, Management, and Recovery
1. Conservation of
intact forest habitat is
critical for long-term
population stability
of forest Kinixys.
Alterations in habitat
patch size can also
affect local weather
patterns impacting
remaining habitat.
Habitat Protection
Habitat destruction, degradation, and fragmentation are among the most serious causes of
current and future population declines among Kinixys and other chelonians ( Böhm, Collen et al.
2012, Klemens 2000, Swingland and Klemens 1989). Based on the degree and rate of habitat
conversion and loss, the fragmentation of African landscapes and wildlife populations is occurring
at an alarming rate. Fragmentation can result in increased mortality, reduced genetic diversity,
increased predation pressure, increased edge habitat, reduced habitat quality, alteration to local
weather and humidity, and invasive species colonization. Habitat protection is the single most
important action that can be taken to assure Kinixys have a contiguous high quality landscape to
maintain viable populations in Africa. Though some species have relatively large theoretical ranges,
available suitable habitat within these has been reduced over the last 30 years and this decline is
expected to continue and worsen.
Connectivity is therefore essential
for long-term viability of Kinixys
populations on a landscape level.
In an effort to maintain population
viability, increasing the number
of protected lands within the
Kinixys range would be beneficial
in preserving connectivity of high
quality habitat. Protected lands
are generally established with
the consideration that exclusion
of land users is necessary to
conserve habitat and biodiversity
while appropriate uses include
tourism and scientific research.
Although these conditions are
ideal, different approaches are also
necessary to ensure that protected
1
74
1. Conversion of habitat
(including on protected
lands) to agriculture
degrades the overall
landscape, fragments
remaining habitat, and
displaces Kinixys and
other wildlife.
Liana May
2. Fire, though
a natural part of
African ecosystems,
is manipulated and
frequently set by
humans. Overuse
impacts habitat integrity
and exposes Kinixys to
greater risk of firerelated trauma and
death.
lands are effective at conserving
biodiversity among the landscape.
Protected areas have long been
the dominant management
strategy for preserving African
wildlife; however, increasing rates
of poaching and other illegal
activities within these areas has
demonstrated the need for new
management techniques. An
emerging strategy that can be
effective is the implementation
of an Integrated Conservation
and Development Project
(ICDP) which attempts to link
1
the conservation of biodiversity
within a protected area to social
and economic development outside that area (Newmark and Hough 2000). Local people that
lose their traditional right to use wildlife populations without receiving any compensation
typically see little incentive to conserve these resources which leads to poaching within protected
lands (Johannesen 2007). Increasing the economic benefits from wildlife for neighboring
populations is a broadly recognized means to elevate local support of wildlife (Weber et
al. 2001). By implementing ICDPs, community members will be engaged through various
incentives including shared decision-making authority, employment, revenue sharing, limited
harvest of plant and animal species, and provision of community facilities. In communities
that are located outside of protected natural areas, because the encroachment on protected
lands and harvesting resources is restricted,
locals often turn to neighboring unprotected
areas and deplete resources just outside of the
protected landscapes (Hartter et al. 2011). To
avoid this situation from occurring, consideration
should be given to linking existing protected
areas with corridors of well-protected habitat
sufficiently wide enough to allow movement
of a range species (McCullough et al. 2005). In
areas where locals worship Kinixys species the
threat of collection is less severe and creating
habitat corridors among these regions would be
beneficial (Luiselli 2003b). Priority should also be
given to zones of sympatry that support multiple
species to provide the greatest value for resources
spent.
2
75
Tomas Diagne
Trade
1.Kinixys can sell for
hundreds of dollars in
the international pet
trade. This demand
makes it worthwhile to
continue to collect.
2. Numerous K. nogueyi
being held in poor
condition awaiting
pick up for sale in the
international pet trade.
Trade in Kinixys has been ongoing since at least the 1960’s with
tens of thousands of animals exported out of Africa since that
time. Since 1975 nearly 200,000 Kinixys were reported as being
exported from Africa. During this time period, over 81,500 K.
homeana were exported with the three major exporting countries
being Togo (43,324), Ghana (20,572), and Benin (16,041) (CITES
Trade Database 2012). Most of this trade has occurred since the
1990’s and in general has increased in volume over time. In recent
years some species have declined in export and it is speculated by
multiple sources this may be the result of a decrease in availability
1
of animals. For some species such as K. natalensis and K. lobatsiana,
the number of animals exported has increased in recent years.
This may be attributed to the relative uniqueness and small ranges
of these species. This may also be the case for K. z. domerguei
which has had some limited export from Madagascar. In total,
nearly 350 individuals were legally exported between 2000 and
2001, which coincides with mass collection and exportation of
Pyxis from Madagascar. Though the volume of exported animals
is relatively small when compared to other Kinixys, this species
occupies an extremely small range. Population estimates for K.
z. domerguei indicate approximately 5,000 individuals remain with
total available habitat of less than 500 hectares (Pedrano 2008).
This amounts to a potential 7% loss of the population in two
years’ time through legal trade. It is known that illegal trade of
tortoises occurs throughout Madagascar despite efforts to stop
2
it (Razafison 2014). There is an increasing variety and volume of
chelonian species being sold as pets on the internet and Kinixys
species have recently been found for sale on websites in Japan. Retail prices for Kinixys range
from $80 to over $1,400US depending on the species and country the animals are for sale in.
It is likely that even relatively inexpensive species such as K. homeana will rise in price due to
elevated global status and increased difficulty in obtaining animals.
Based on the life history traits and our understanding of Kinixys threats, it appears that
there are no means for sustainable harvest of this genus that would not result in decline and
eventual localized and regional population collapse. Elimination of quotas for wild collected or
“ranched” animals for K. homeana and K. erosa are strongly recommended. Significant reduction
in quotas for all other Kinixys species is strongly recommended. Due to the demand and interest
in these species, migration toward greater emphasis on captive bred animals and facilities and
elimination of wild collected Kinixys from trade is encouraged. This shift can reduce collection
pressures and still provide revenue for locals within range countries.
Currently all species of Kinixys are listed under appendix II of CITES. Review of elevating
target Kinixys species to CITES I should be considered with species that experience significant
exportation or those with small ranges reviewed as a priority. Care must be taken as this stricter
designation often results in increased valuation and black market sales of rare species. This
designation may also increase difficulty in international collaboration and moving animals back
into range countries if reintroduction is ever proposed. Furthermore, limiting the trade of these
species will only work as part of an overall conservation strategy that also includes additional
habitat protection and improved management.
76
Tomas Diagne
Within range countries, regulations should be enacted to
protect live Kinixys from sale in food markets or animals parts
sold in fetish markets. Bushmeat trade is likely the largest
contributor to declines in K. erosa. Increasing demand from
foreign nationals in eastern and southern Africa warrants
greater local protection for all Kinixys.
1
1. One of the strongest
conservation tools we
have is engaging the
local community in the
role and importance
of biodiversity and
conservation. This
has been shown to
be effective, but
local partnership and
engagement is critical .
2. Multiple K. erosa.
offered for sale in a
bushmeat market in
Equatorial Guinea
Education and Outreach
Community based conservation will be critical to help combat
the collection of Kinixys and reduce the loss of habitat. Engaging locals in research opportunities
and conservation initiatives may help minimize the pressures these species are facing through
locally initiated efforts. This system has been shown to be effective in helping preserve species
(Wolmer et al. 2004). This approach requires strong collaboration and communication between
project partners within range countries and abroad. Due to recent outbreaks of Ebola in West
Africa and the potential link to bushmeat consumption (Fombu 2014), encouraging locals to
reduce use of bushmeat may improve community health while also protecting wildlife. For local
conservation to be successfully implemented, adequate resources as well as support and guidance
are necessary. Conservation efforts at the local level are warranted and necessary to engage
community members in helping protect and
value these tortoises. This can include wildlife
protection, assisting with research, and,
when appropriate, developing and managing
assurance colonies.
Adaptive Management
When dealing with species in a crisis
conservation mode we must also consider the
needs for adaptive management and flexibility
in design to allow and account for new and
emerging threats. In some cases action and
implementation is required using the best
available data.
3
Vin Crosbie
Adaptive Management is a structured
approach to resource management. Through
this iterative process, conservation managers
and on-the-ground biologists work together
to improve conservation and management
of Kinixys over time by learning from
2
Matt Muir
3. Barriers, including
electric fences, fragment
the landscape and can
result in direct mortality.
Effort to allow
passage of tortoises
would minimize this
unintended impact.
Restoring connectivity
and identifying priority
corridors for protection
is an essential part
of long-term species
management.
It is also important that the international conservation
community recognize that the illegal black market trade in
wildlife extends beyond our charismatic mega-fauna and
includes tortoises and other herpetofauna. Conservation
groups operating within Africa should consider wildlife
protection on a bottom up approach and develop conservation
strategies that incorporate these declining important sentinel
species in recovery and management plans.
77
management outcomes. Adaptive Management entails a multistep process:
1. Considering various actions to meet management objectives;
2. Predicting the outcomes of these management actions
based on what is currently known;
Dwight Lawson
3. Implementing management actions;
4. Monitoring to observe the results of those actions; and
1
5. Using the results to update knowledge and adjust future
management actions accordingly.
By repeating this cycle and increasing the body of knowledge
about the system in question, managers are able to refine their
prescriptions to more closely meet the original objectives (U.S.
Geological Survey 2013).
Tomas Diagne
Assurance Colonies
1. Adaptive
management requires
the ability to adjust to
various situations. Here
founder stock for a
K. homeana assurance
colony is set up to be
cared for by zoo and
veterinary staff while
working with various
partners to accept
animals.
2. Range country
assurance colonies are
an essential part of
genus conservation
as they also
provide educational
opportunities for locals
to help engage them in
species protection.
Though the conservation priority is to protect habitat and
species in situ, effort should also be taken to provide additional
safeguards in the event that short-term efforts resulting from
social, political, financial or environmental factors do not result
in sufficient protection to assure species survival.
Both in-situ and ex-situ assurance colony programs should
be developed to optimize and increase redundancies and
learning opportunities. Programs within a species’ native
range have the benefit of natural climatic conditions as well as
opportunities for local community engagement and increased awareness of species protection.
In-situ programs also result in reduced logistical issues regarding animal release although ex-situ
programs have also been effective in helping species recover. These programs often include
dedicated individuals and resources including trained veterinary teams who can help optimize
animal health condition and species reproduction.
2
To date, no formal assurance colony program has been established within range countries though
effort is underway at an existing facility in Senegal to establish multiple West African Kinixys
species. Ex-situ conservation programs on various Kinixys species including the establishment
of assurance colonies have recently been initiated in the U.S. to better understand these unique
tortoise’s behaviour and husbandry and to help protect wild populations. The first Kinixys
assurance colony established was for K. homeana in 2013 following the IUCN Redlist workshop and
recommendations. In total approximately 100 founder animals have been incorporated into the
assurance colony. Significant work still needs to be done for other Kinixys species though additional
assurance colonies are being developed for K. erosa and K. spekii. Recently studbook coordinators
78
1. Proper husbandry
and record keeping
are key to successful
assurance colonies of
Kinixys. Little is known
about these species in
the wild as well as in
captivity. Collaboration
and communication are
vital components to
successfully managing
and expanding assurance
colonies.
2. Increased interest
in the conservation
of Kinixys and the
development of
better communication
mechanisms for those
working with the species
in various ways are
important in developing
stronger partnerships
and tangible success.
for both K. homeana and K. erosa were selected
and work on these will be ongoing. This is
a collaborative effort between multiple zoos,
conservation organizations, and private
individuals.
Husbandry and Captive
Breeding
As part of the development of an assurance
colony program, establishing guidelines
for basic husbandry and captive breeding
guidelines is essential. Captive collections
can be beneficial in acquiring much needed
information about these species and how
1
to help protect them. Kinixys species that
are being successfully bred outside of range
countries include K. erosa, K. homeana, K. nogueyi, in both the U.S. and Europe, and K. spekki in the
U.S. It has also been reported that captive colonies of K. lobatsiana and K. z. zombensis are being
successfully bred within range countries. In an effort to continue expanding the understanding
of this little understood genus and the success of captive collections, a questionnaire has been
developed and will be sent to various partners and individuals working with Kinixys to expand
the knowledge and individual experiences on how to care for and breed these tortoises. Kinixys
have a reputation for being difficult to maintain in captivity, so guidelines for their general care
have been provided in Appendix D as well as a chart describing time frames of their activity in
Appendix F. In an effort to remain adaptive and current with our knowledge about these poorly
understood animals, a questionnaire was also developed (Appendix E) as a means to consolidate
available knowledge on the genus and successful ways they are maintained and bred. This ongoing
effort and knowledge sharing will be critical in successfully establishing assurance colonies or
developing repatriation or reintroduction program.
Jim Harding
Reintroduction Programs
2
Though certainly in decline, with some species critically so, it is not
recommended that any reintroduction program be initiated until
there is greater understanding of species natural history and overall threats, demonstrated protection of habitat, and efforts to stop
the collection of wild animals for consumption and the pet trade.
Though a viable conservation tool when properly implemented,
this strategy should not be implemented in the near future. Application of on the ground research, proactive habitat conservation,
and species protection will be the most effective approach to the
prevention of continued declines and potential extinction. Mitigation of threats and education of the local people is needed to
ensure the survival of reintroduced animals.
79
Sean Zera
Genetic Management
1.Success with Kinixys
conservation can be
measured in various
ways. Understanding
and breaking
developmental diapause
is a critical step in
successfully breeding
Kinixys in captivity
1
Evaluation of genetics and maintaining sufficient gene
diversity in captive colonies will be an important part of
establishing viable long term assurance colonies. Because
many Kinixys occur over a large range, evaluation of
genetic clades within species will help best determine
which specimens should be bred together. A management
strategy and breeding transfer plan should be developed
in coordination with taxon managers and Kinixys advisory
groups. Initial work should focus on K. homeana and K.
erosa as these populations are likely experiencing the most
significant declines. Additional effort should be placed
on K. nogueyi and K. spekii as both grassland species are
successfully being produced in small numbers within
the U.S. and Europe and greater understanding of their
genetics will be necessary as future offspring are produced.
Research Needs
There is a considerable lack of available data for this genus
and effort should be put into conducting research to better understand the species’ distribution
and ecology Building on the coarse scale analyses conducted as part of this document,
conservation groups and regional stakeholders will benefit from the development of more
refined habitat suitability models including amounts of protected lands in addition to conducting
population viability analysis as a means of risk assessment. Spatial analysis considering landscape
factors that reduce Kinixys presence should be evaluated to develop more accurate maps of
the true potential available habitat, degree of fragmentation, and potential for restoration of
connectivity Once the true extent of available habitat is better understood and where in this
range species are least vulnerable to exploitation, habitat corridors that create linkages between
these communities can be identified. Identification of field localities is of the utmost importance
in ranges where locality data is limited or species are known only from the markets. During
future research, emphasis should be placed on K. belliana due to the considerable lack of locality
information for this species in parts of its range. Additionally, habitat analysis focused on the
relationship between humans and tortoises among different communities may be beneficial to
identifying critical habitat corridors where animals are worshipped and do not face as significant
threat of collection.
To gain a better understanding of Kinixys ecology, studies are encouraged to focus on
reproductive behavior, nesting, incubation and associated diapause, juvenile diet, and habitat
selection. Work should also assess population structure including size, age class, and distribution
while monitoring the extent of suitable habitat and how changes in availability are affecting local
populations. Future genetic work is warranted to assess population-level and range-wide genetic
information. Collection of genetic samples during research activities will help better verify the
number of species within the genus while defining geographic distribution and relatedness
among species. This establishment of assurance colonies will also benefit from the collection of
genetic data. Genetics can be used to determine the influences of inbreeding and outbreeding
depression in colonies with limited numbers of founder individuals. This information will also
establish reference blood ranges for future veterinary and health analyses as well as gaining a
better understanding of disease pathology.
80
Appendices
Appendix A:
Species Table
Forelimbs/
hindlimbs
Carapace
Plastron
Hinge
Beak
Range
5/4 claws
(Swingland and Convex, scutes raised;
M+F: 190Klemens 1989) radiated pattern on
220mm
No tubercles
carapace, areolae
(Boycott and
on buttocks
brown with light ring
Bourquin
(Boycott and surrounding (Boycott
2000)
Bourquin 2000) and Bourquin 2000)
Kinixys homeana
M: /323mm,
F: /260mm
(Swingland
and Klemens
1989)
Gambia to Dem. Rep. of Congo
and Uganda, S to Cabinda, N
Carapace slopes
Front lobe thick and
Angola, and N shore of Lake
dorsally, scutes flat,
notched, projects
Well developed,
Tanganyika (Swingland and
serrated marginals,
beyond carapace;
5/4 claws
btwn 7th and 8th Unicuspid and not Klemens 1989); Guinea, Sierra
scarcely to mod notched Black with yellow
(Swingland and
marginal (Boycott serrated (Swingland Leone, Liberia, Cote Divoire,
nuchal region; Dark in on outer margins
Klemens 1989)
and Bourquin
and Klemens 1989) Nigeria, Cameroon, Equatorial
color with lighter scute and along sutures
2000)
Guinea, Gabon, Congo, Central
centers (Swingland and (Swingland and
African Republic, Dem. Rep.
Klemens 1989)
Klemens 1989)
of Congo, Angola, and Uganda
(Vetter 2011)
NE Dem. Rep. of Congo through
S Sudan and Ethiopia to NW
Truncated and
Unicuspid and not Somalia and S to Uganda and W
usually protrudes
Well developed, serrated (Swingland Kenya (Swingland and Klemens
beyond carapace;
btwn 7th and 8th and Klemens 1989)
1989)
black splashes
marginal (Boycott
Rarely hooked
Cameroon, Central African
radiating outward
Republic, Dem. Rep. of Congo,
and Bourquin
(Boycott and
or uniform light
Rwanda,
2000)
Bourquin 2000)
color (Boycott and
Sudan, Ethiopia, Uganda,
Bourquin 2000)
Tanzania, Kenya, Eritrea, and
Somalia (Vetter 2011)
81
M: /211mm,
F: /223mm
(Swingland
and Klemens
1989)
Front lobe thick
and notched, may
Unicuspid and not
Flattened dorsally,
Dem. Rep. of Congo W to
project slightly
Well developed, serrated; mod to
serrated scutes at rear;
Liberia (Swingland and Klemens
5/4 claws
beyond carapace; btwn 7th and 8th strongly hooked
dark in color, scutes
1989); Liberia, Cote Divoire,
(Swingland and
yellow or darker marginal (Boycott (Swingland and
may have lighter centers
Ghana, Togo, Benin, Nigeria,
Klemens 1989)
with yellow along
and Bourquin
Klemens 1989)
(Swingland and Klemens
Congo, and Dem. Rep. of Congo
the margins
2000)
Slightly upturned
1989)
(Vetter 2011)
(Swingland and
nose
Klemens 1989)
Kinixys erosa
Min/Max
Length
Kinixys belliana
Species
Min/Max
Length
Forelimbs/
hindlimbs
Carapace
Plastron
Hinge
Beak
Range
W Africa from Senegal to Cameroon
Unicuspid and not
Domed carapace
and the Central African Republic
Well developed, serrated (Swingland
4/4 claws
which is uniformly Irregular or uniform
(Ernst and Barbour 1989); Senegal,
btwn 7th and 8th and Klemens 1989);
Adult: 200 - (Swingland brown or with yellow
dark blotches
Gambia, Guinea Bissau, Guinea,
marginal (Boycott Upper jawline is
300mm
and Klemens areolae with a dark
(Boycott and
Sierra Leone, Mali, Burkina Faso,
and Bourquin
indented in front
1989)
border (Ernst and
Bourquin 2000)
Cote Divoire, Ghana, Togo, Benin,
2000)
of the eye (Boycott
Barbour 1989)
Cameroon, Central African Republic
and Bourquin 2000)
and Nigeria (Vetter 2011)
Kinixys spekii
Kinixys z. zombensis
Central plateau south of the
equatorial forests; Kenya to N South
Africa (Transvaal), Swaziland and
Front lobe thick and Well developed, Unicuspid and not S Mozambique, W to S Dem. Rep.
projects well beyond btwn 7th and 8th serrated, not to
of Congo, Angola and N Botswana
carapace, concave marginal (Boycott weakly hooked
(Swingland and Klemens 1989);
in males (Swingland and Bourquin
(Swingland and
Kenya, Rwanda, Burundi, Dem.
and Klemens 1989)
2000)
Klemens 1989)
Rep. of Congo, Malawi, Zambia,
Mozambique, Angola, Namibia,
Botswana, Zimbabwe, Swaziland, and
South Africa (Vetter 2011)
5/4 claws
(Swingland
M: /181mm, and Klemens
F: /200mm
1989)
(Boycott and No tubercles
Bourquin
on buttocks
2000)
(Boycott and
Bourquin
2000)
Front lobe may be
M: /206mm,
slightly notched and Well developed,
5/4 claws Domed carapace with
F: /217mm
projects beyond btwn 7th and 8th Unicuspid and not
Kenya, Tanzania, Malawi,
(Swingland a broad black radial
(Swingland
carapace; uniform marginal (Boycott serrated (Swingland Mozambique, Zimbabwe, and South
and Klemens pattern (Swingland
and Klemens
light color or black
and Bourquin and Klemens 1989)
Africa (Vetter 2011)
1989)
and Klemens 1989)
1989)
splashes radiating
2000)
outward
Kinixys z. domerguei
82
Kinixys nogueyi
Species
M: 19.4cm, F:
19.6cm
(Pedrono
2008)
5/4 claws
Flattened, scutes
relatively flat with
well-defined annuli;
M: Uniform color,
F: Uniform to short
radiations (Swingland
and Klemens 1989)
Flat dorsal
Front lobe may be
Unicuspid; frontal
surface,with unflared
Northeastern Madagascar (Vetter
slightly notched and Well developed, scale is fragmented
marginal; Variable
2011)
projects beyond btwn 7th and 8th and the width of
in color from yellow
Nosy Faly (satellite island) (F.
carapace; uniform marginal (Boycott the frontals is less
to brown, darker
Andreone and Raxworthy 2003),
light color or black
and Bourquin
than the width of
radiated pattern in
Ambato peninsula (Pedrono 2008)
splashes radiating
2000
the prefrontals.
adults. (Pedrono
outward
(Pedrono 2008)
2008)
Kinixys natalensis
5/4 claws
Front lobe, may projMod convex, Poste(Swingland
ect slightly beyond
rior forms “gutter”
E Transvaal, South Africa and MozamM: /125mm, and Klemens
carapace, male has Poorly developed,
w/ slight serration,
bique, S through Swaziland to Natal
F: /155mm
1989)
distinct concavity; btwn 7th and 8th Tricuspid (Swingwell-marked annuli;
midlands at Pietermaritzburg (Swing(Swingland No tubercles
yellow centers and marginal (Boycott land and Klemens
concentric pattern on
land and Klemens 1989); Mozamand Klemens on buttocks
margins, 2 black
and Bourquin
1989)
each scute with brown
bique, Swaziland, and South Africa
1989)
(Boycott and
rings on abdominals
2000)
areolae (Swingland
(Vetter 2011)
Bourquin
(Swingland and Kleand Klemens 1989)
2000)
mens 1989)
Kinixys lobatsiana
5/4 claws
(Swingland
M: /162mm, and Klemens
F: /167mm
1989)
(Swingland No tubercles
and Klemens on buttocks
1989)
(Boycott and
Bourquin
2000)
Mod convex, wellSE Botswana, E to central Transvaal
marked annuli, buff
middleveld (Swingland and Klemens
to dull yellow with Front lobe truncated
1989)
brown areolae,
and extends to or be- Well developed, Unicuspid and not
South Africa and
sometimes irregular yond carapace; Buff btwn 7th and 8th serrated; weakly to
Botswana, found in three reserves in
radiations (Swingland with brown border, marginal (Boycott moderately hooked
South Africa (Boycott and Bourquin
and Klemens 1989);
sometimes black
and Bourquin
(Swingland and
2000)
Posterior marginal rays (Swingland and
2000)
Klemens 1989)
South Africa and Botswana (Vetter
shields are upturned
Klemens 1989)
2011)
at edges. (Boycott and
Bourquin 2000)
83
84
Appendix B: Maps
1. Sub-Saharan Africa land cover classification types.
2. Sub-Saharan Africa elevation displaying in meters above sea-level
3. Locations of Sub-Saharan Africa mining sites and type of method practiced including: unknown mining methods,
open-pit or surface mines, surface mines with underground component, underground mines with surface operations, and
underground mines.
4. Locations of Sub-Saharan roads, mapped in black, and population density centers (people/square km) with 2.5 km
spatial resolution. The combined range of all Kinixys species is included in pink.
5. K. homeana range with available potential habitat defined by the presence of suitable land cover and community types
associated with this species. Range area and available habitat given in hectares, and percentage of available habitat are
provided as well.
6. K. homeana range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014. The
colors represent a count of the number of fires observed within a 1,000-square-kilometer area. White pixels show as many
as 100 fires in a 1,000-square-kilometer area per day. Yellow pixels show as many as 10 fires, orange shows as many as 5
fires, and red areas as few as 1 fire in a 1,000-square-kilometer area per day.
7. K. erosa range with available potential habitat defined by the presence of suitable land cover and community types
provided as well.
8. K. erosa range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014. The
9. K. belliana range with available potential habitat defined by the presence of suitable land cover and community types
provided as well.
10. K. belliana range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014.
The colors represent a count of the number of fires observed within a 1,000-square-kilometer area. White pixels show as
many as 100 fires in a 1,000-square-kilometer area per day. Yellow pixels show as many as 10 fires, orange shows as many
as 5 fires, and red areas as few as 1 fire in a 1,000-square-kilometer area per day.
11. K. nogueyi range with available potential habitat defined by the presence of suitable land cover and community types
provided as well.
12. K. nogueyi range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014. The
85
13. K. spekii range with available potential habitat defined by the presence of suitable land cover and community types
provided as well.
14. K. spekii range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014. The
15. K. zombensis range with available potential habitat defined by the presence of suitable land cover and community types
provided as well.
16. K. zombensis range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014.
17. K. natalensis range with available potential habitat defined by the presence of suitable land cover and community types
provided as well.
18. K. natalensis range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014.
19. K. lobatsiana range with available potential habitat defined by the presence of suitable land cover and community
types associated with this species. Range area and available habitat given in hectares, and percentage of available habitat are
provided as well.
20. K. lobatsiana range with fire density and frequency. Fire intensity data was recorded from May 2013 - February 2014.
Map 1
86
Source: GLOBCOVER by ESA/UNEP/FAO/JRC/IGBP/GOFC-GOLD
Map 2
87
88
Map 3
Map 4
89
90
Map 5
Range Area Habitat within % Available
in Hectares Range in Hectares Habitat
46,332,430
13,613,700
29%
Map 6
91
92
Map 7
285,723,701
88,919,937
31%
Map 8
93
94
Map 9
in Hectares Range in Hectares
Habitat
389,611,672
228,919,938
59%
Map 10
95
96
Map 11
284,200,067
98,075,500
38%
Map 12
97
98
Map 13
353,103,172
209664054
59.38%
Map 14
99
100
Map 15
116,825,768
41,967,396
36%
Map 16
101
102
Map 17
13,004,682
5,119,592
39%
Map 18
103
104
Map 19
13,252,785
10,560,175
79.68%
Map 20
105
106 Kinixys Conservation Blueprint
Appendix C: Data Tables
Kinixys Exported by Top Six African Countries for
Exportation Between 1975-2012
120,000
108,128
100,000
Kinixys exported
Total Kinixys African Exports Between 1975-2012
#
% of total Kinixys
Country
Exports
Exports
Equatorial Guinea
1
0%
Angola
2
0%
Guinea-Bissau
2
0%
Sierra Leone
2
0%
Somalia
2
0%
Lesotho
3
0%
Malawi
3
0%
Central African Rep.
10
0%
Gabon
16
0%
Rwanda
20
0%
Liberia
29
0%
Nigeria
36
0%
Niger
52
0%
Burundi
53
0%
Rep. of the Congo
66
0%
Burkina Faso
200
0%
Uganda
203
0%
Zimbabwe
204
0%
D. Rep. of the Congo 216
0%
Guinea
269
0%
Madagascar
349
0%
South Africa
358
0%
Cameroon
734
0%
Mali
740
0%
Chad
820
0%
Ivory Coast
856
0%
Zambia
1,253
1%
Tanzania
2,194
1%
Kenya
3,311
2%
Mozambique
6,317
3%
Benin
33,161
17%
Ghana
34,474
18%
Togo
108,128
56%
TOTAL = 194,084
80,000
60,000
33,161 34,474
40,000
20,000
2,194
3,311
6,317
0
% of Total Kinixys Exported Between 1975-2012
Tanzania Kenya
1%
2%
Mozambique
other
3%
3%
Benin
17%
Togo
56%
Ghana
18%
The table to the left contains the total count of Kinixys exported
from African countries. This count includes only primary
exportation and excludes re-exports between countries to produce
a more accurate number of tortoises leaving the country. The top
bar graph comparatively illustrates the top six exporters of Kinixys,
while the pie chart illustrates the countries responsible for the
largest amount of Kinixys exported.
107
Kinixys Exports 1975-1989
Country
# Exports
Lesotho
1
Sierra Leone
1
Guinea
2
Nigeria
2
Somalia
2
Burundi
16
Dem. Rep. of Congo
16
Cameroon
16
Mozambique
20
Rwanda
20
Zimbabwe
23
Liberia
29
South Africa
34
Tanzania
1,912
Kenya
2,961
Ghana
3,832
Togo
23,352
Total = 32,239
Country
# Exports
Sierra Leone
1
Angola
2
Guinea-Bissau
2
Lesotho
2
Ivory Coast
4
Gabon
9
Cen. African Rep.
10
Dem. Rep. of Congo
28
Nigeria
28
Burundi
37
Uganda
53
Mali
110
South Africa
119
Madagascar
148
Zimbabwe
197
Burkina Faso
200
Kenya
350
Cameroon
566
Zambia
765
Tanzania
1,059
Mozambique
3,258
Ghana
13,073
Benin
18,427
Togo
49,191
Total = 87,639
Country
# Exports
Equatorial Guinea
1
Malawi
3
Nigeria
6
Gabon
7
Niger
52
Rep. of Congo
66
Uganda
150
Cameroon
152
Tanzania
152
Dem. Rep. of Congo
172
Madagascar
201
South Africa
222
Guinea
267
Zimbabwe
488
Mali
630
Chad
820
Ivory Coast
852
Mozambique
3,039
Benin
14,734
Ghana
17,569
Tanzania
152
Togo
39,181
Total = 78,916
The above tables detail the total exports per country over
approximate decade time constraints. Since 1975 there
has been a steady increase in both exports and countries
exporting however it peaks between 1990-2000, with 20012012 onward experiencing a slow decline in later years.
49191
Top Kinixys Exporter Growth
50000
39181
45000
40000
35000
23352
30000
13073
17569
18427
14734
25000
20000
15000
Tanzania
Mozambique
Kenya
0
3832
2961
350
0
3258
3039
20
1912
1059
152
0
Ghana
Benin
1990-2000
2001-2012
10000
5000
1975-1989
Togo
The table to the left illustrates the number of exports per
decade of the top six countries with the highest number
of Kinixys exported. Tanzania and Kenya have experienced
steadily decreasing exports since 1975, however Ghana’s
Kinixys trade has continued to grow. Togo and Benin both
spiked to high numbers initially then experienced a small
decline at the latter end of the last decade.
Total Kinixys Exported
14000
12000
10000
8000
6000
4000
2000
0
2010
2005
2000
1995
1990
1985
1980
1975
Total Kinixys African Exports
Between 1975-2012
Species
# Exports
K. belliana complex
97,790
K. homeana
81,559
K. erosa
9,104
K. spekii
1,516
K. natalensis
28
K. lobatsiana
51
The line graph above demonstrates the total number of Kinixys
exported from all African countries per year. Trade has been on a
steady increase until 2002 where it peaks and has declined to a lesser,
but steady amount.
The table to the left is a species breakdown of the total exports
composition. Note that during the years export data was recorded, a
handful of individual Kinixys species were not recognized as separate
from K. belliana. K. spekii was recognized after 2006 in trade data, K.
natalensis in 1980, and K. lobatsiana in 2006.
K. belliana complex is composed of the following species:
K. belliana, K. spekii, K. zombensis, K. nogueyi, K. natalensis and K.
lobatsiana either through old nomenclature or misidentified tortoises.
109
8000
7000
6000
5000
Togo
4000
Ghana
Benin
3000
2000
1000
0
2011
2006
2001
1996
1991
1986
1981
1976
The above graph details total Kinixys exports for the top three ranking countries in Africa for Kinixys export. Togo has
been exporting Kinixys since 1977 and has in general maintained a high count with its export numbers peaking in the
year 2000 with 7,066 exports. Ghana has been exporting Kinixys since 1978 and peaked in the year 1989 with 3,360
exports. Ghana’s trade numbers seesaw with Benin’s, which started exporting in 1991 and hit its peak in 1997 with 4,612
exports while Ghana hit record lows. Similarly, Ghana’s trade increased between 1999 and 2001 while Benin hit a low.
The record low for Benin also corresponds with the record high for Togo in the year 2000, implying the trade from Benin
shifted largely to Togo. Another rise in Ghana’s trade numbers between 2004 and 2008 corresponds with a period of low
exports for Benin, implying that periods of trade alternate between these two counties.
K. belliana complex
1975-2012
Country
# Exports % of total
Angola
2
0%
D. Rep. of Congo
2
0%
Liberia
2
0%
Lesotho
2
0%
Somalia
2
0%
Malawi
3
0%
Rwanda
8
0%
Burundi
42
0%
Rep. of the Congo
66
0%
Uganda
150
0%
South Africa
151
0%
Burkina Faso
200
0%
Zimbabwe
202
0%
Cameroon
281
0%
Madagascar
348
0%
Chad
360
0%
Ivory Coast
402
0%
Zambia
949
1%
Tanzania
2,179
2%
Kenya
3,311
3%
Mozambique
4,044
4%
Ghana
8,929
9%
Benin
16,422
17%
Togo
59,733
61%
Total = 97,790
% of Total K. belliana Complex Exported
1975-2012
Tanzania Kenya
4%
2%
Mozambique
4%
Zambia
1%
Ghana
9%
Benin
17%
Togo
63%
Note: K. belliana should be regarded as K. belliana complex as some species of
Kinixys weren’t recognized as separate during the years of data collection. Likely
contains: K. belliana, K. spekii, K. zombensis, K. nogueyi, K. natalensis and K.
lobatsiana either through old nomenclature or misidentified tortoises.
The table to the left and the corresponding bar graph below detail the total
numbers of K. belliana complex exports from African countries between
1975-2012. Judging by known locality: the majority of K. belliana trade from
Mozambique is most likely K. spekii and K. z. zombensis. Exports from South
Africa likely contain K. natalensis and K. lobatsiana as well as K. belliana. Exports
from Togo, Benin, and Ghana likely represent K. nogueyi and exports from
Madagascar are K. z. domerguei. The above pie chart illustrates the percentage of
countries making up the bulk of K. belliana complex exports between the top 7
exporters.
59733
K. belliana Complex Exports
1975-2012
60000
50000
4044
3311
2179
949
402
360
348
281
202
200
151
150
66
8
3
2
2
2
2
0
2
10000
42
20000
8929
30000
16422
40000
111
K. erosa 1975-2012
Country
Gabon
1
0%
Equatorial Guinea
1
0%
Nigeria
1
0%
Zambia
1
0%
Sierra Leone
2
0%
Zimbabwe
2
0%
Central African Rep.
10
0%
Burundi
11
0%
Rwanda
12
0%
Liberia
27
0%
Mali
50
1%
Uganda
53
1%
Cameroon
134
1%
Guinea
159
2%
D. Rep. of Congo
214
2%
Benin
698
8%
Ghana
3,471
38%
Togo
4,257
47%
Total = 9,104
% of Total K. erosa Exported
1975-2012
Uganda
1%
Cameroon
1%
Guinea
2%
D. Rep. of the
Congo
2%
Benin
8%
Togo
47%
Ghana
39%
The table to the left and the corresponding bar graph below detail the
total numbers of K. erosa exports from African countries between 19752012. The above pie chart illustrates the percentage of countries making
up the bulk of K. erosa exports between the top 7 exporters.
K. erosa Exports
1975-2012
4500
4000
3500
3000
2500
2000
1500
1000
500
0
4257
3471
698
1
1
1
1
2
2
10
11
12
27
50
53
134
159
214
K. homeana 1975-2012
Country
Guinea-Bissau
2
0%
Nigeria
3
0%
Gabon
12
0%
Niger
52
0%
Zambia
55
0%
Guinea
110
0%
Mali
200
0%
Cameroon
277
0%
Ivory Coast
451
1%
Chad
460
1%
Benin
16,041
20%
Ghana
20,572
25%
Togo
43,324
53%
Total = 81,559
% of Total K. homeana Exported
1975-2012
Ivory Coast
0%
Chad
1%
Benin
20%
Togo
54%
Ghana
25%
The table above and the corresponding bar graph below detail the total numbers of K. homeana exports from African
countries between 1975-2012. The above pie chart illustrates the percentage of countries making up the bulk of K.
homeana exports between the top 5 exporters.
K. homeana Exports
1975-2012
45000
40000
35000
30000
25000
20000
15000
10000
5000
0
43324
16041
2
3
12
52
55
110
200
277
451
460
20572
113
K. lobatsiana 1988-2012
Country
South Africa
51
100%
Total =
51
% of Total K. spekii Exported
1993-2012
South Africa
1%
K. natalensis 1994-2012
Country
South Africa
14
50%
Zimbabwe
14
50%
Total =
28
K. spekii 1993-2012
Country
South Africa
Zimbabwe
Mozambique
Zimbabwe
22%
Mozambique
77%
10
1%
335
22%
1,171
77%
Total = 1,516
The tables above detail the total numbers of recorded K. lobatsiana, K. natalensis, and K. spekii exports from African
countries between 1975-2012. K. lobatsiana wasn’t accounted for in this data until 2008, K. natalensis in 1980,
and K. spekii in 2006 as the species became differentiated in trade data. As such, the numbers do not represent an
accurate picture of the trade over the entire data set and the majority of them were included under K. belliana. The
majority of K. belliana trade from Mozambique was most likely K. spekii judging by known locality. The above pie
graph illustrates the countries making up the highest percentage of K. spekii trade. K. natalensis are exported from
South Africa and Zimbabwe, while South Africa is responsible for all recorded K. lobatsiana exports.
K. spekii Exports
1975-2012
1400
1171
1200
1000
800
600
335
400
200
0
10
South Africa
Zimbabwe
Mozambique
Togo
species composition of 108128 total exports 1975-2012
Ghana
1%
4%
4%
10%
26%
40%
55%
K. belliana
K. belliana
K. homeana
K. homeana
K. erosa
K. erosa
unknown spp.
unknown spp.
60%
Benin
Mozambique
2%
17%
50%
48%
K. belliana
K. belliana
K. homeana
K. spekii
19%
unknown spp.
K. erosa
64%
The above pie charts illustrate the majority species composition of the total Kinixys exports of four African countries.
Togo’s exports are largely K. belliana complex, likely K. nogueyi (55%), Ghana’s are mostly K. homeana (60%), Benin
strikes a close 50:50 between K. belliana complex (likely K. nogueyi) and K. erosa, and Mozambique’s exports are largely K.
belliana (which is likely to be K. spekii as well).
115
Benin 1997-2001, Top Ten Species in Trade
The bar graphs to the left
illustrate the underlined
Kinixys species regularly
placing in the top ten
species in trade for
Benin.
Ghana 1997-2001, Top Ten Species in Trade
species in trade for Togo.
117
Togo 1997-2001, Top Ten Species in Trade
species in trade for Togo.
Appendix D: Assurance Colony Husbandry Recommendations
1. The establishment
of assurance colonies
is an important
component to the
overall conservation
of this genus. Through
collaborative efforts,
healthy colonies of
tortoises are being
established in the U.S.
and Europe.
As Kinixys become increasingly rare in the wild and pressures continue to threaten their existence,
measures both in-situ and ex-situ measures should be taken to preserve these species and reduce
the risk of species collapse or extinction. Development of assurance colonies is one component
to an overall conservation approach to safeguard these imperiled species. These guidelines are
general in nature and do not necessarily apply for each situation or tortoise. Observations of
your colony, location, and their responses to various husbandry techniques will help best direct
how to approach their care. As the different species occur over an expansive range encompassing
a majority of Africa and a variety of climatic zones and ecosystems, care should be tailored to
their specific needs. Though this husbandry summary covers all Kinixys (both tropical forest and
grassland), specific needs for a species are noted when warranted .
Housing
Indoor Enclosures
Depending on the geographic location of an assurance colony, Kinixys can
be maintained in either indoor or outdoor environments. Kinixys colonies
in most parts of the U.S. and Europe will require indoor facilities for at
least part of the year due to cold winters.
1
Enclosures should be spacious with at least three square feet per inch of
shell. If kept in groups, sufficient area must be provided. Cypress mulch
or hardwood mulch are generally readily available products that are
good bases for substrate. In addition peat or coconut fiber are beneficial
as additives that provide more natural substrate texture and moisture
retention when mixed with other base substrates. A mixture containing
large proportions of peat can become too saturated or soggy and it is
recommended that it be supplemented with sand to aid in maintaining
aeration. These mixtures provide quality substrates that allow tortoises to
express natural burrowing behaviors and provide optimal indoor nesting
substrate. Sphagnum moss is another substrate that is effective at retaining
moisture; however, care should be taken to use material that has been
119
Ruth Basham
1
1. Enclosures should
contain ample hiding
areas which can be
created using various
materials including
plastic plant pots.
2. This K. nogueyi
hatchling is soaking
itself in a water dish. To
ensure proper hydration,
tortoises should
always have access to a
source of water and be
maintained in a humid
environment.
2
sustainably harvested. The depth of substrate within
an enclosure should be maintained along a gradient
with portions containing at least 10 to 12 inches of
material to allow for successful nesting. Leaf litter
for moisture preservation and providing refugia is
a worthwhile addition to Kinixys enclosures. This
additional structure can result in increased natural
behavior of individuals and an improved sense of
security. Do not collect leaves or other outdoor
materials where pesticides or domestic animal feces
are present. Hides are encouraged and can be made
from a variety of things including wood, slate, or even
plastic plant pots. Ease of access for maintenance
and inspection of tortoises is vital. Preference is
given to plastic pots as these can be easily cut in half,
come in a variety of sizes, and are easy to clean and
decontaminate. Cork bark is another material that
is useful as it can easily be sterilized by heating in an
oven at 325° F for roughly 10 minutes. Alternatively
wood structures can be used as they are durable, but
should be maintained with a pet-friendly paint that
prevents moisture buildup and potential rotting.
Painting these structures also allows for easier clean
up and decontamination. In addition to housing,
plastic or silk plants can create a more naturalistic
enclosure and provide an added security that will help
reduce stress and facilitate a more comfortable Kinixys
resident. Artificial plants have the added benefits of
being easier to maintain indoors and of not being
eaten by the tortoises.
Hatchlings do well in a mixture of coconut fiber (coir) and play sand, which is easily moisture
regulated. Hides need to be provided as these young tortoises prefer a place to find shelter
and can include materials used for adult enclosures scaled down in size. Plastic flower pots cut
into halves or terracotta pots buried halfway on their sides provide simple and ample hides.
Additionally, an overturned plastic basket with an entryway cut into it or a hide constructed of
wood can provide suitable shelter.
Kinixys are most active and healthiest when maintained at 60-90% humidity though grassland
species can be maintained on the lower end of this range. This can be achieved by misting
several times daily or use of a fogging system. In addition partially covering the enclosure with a
plastic tarp and/or incorporating nontoxic live plants can help maintain higher humidity levels.
As a general rule, temperatures should be kept between 70°- 85°F for day temperatures and
60°-70 °F at night. Though Kinixys will bask, ambient temperatures seem to be sufficient for
1
1. To achieve optimal
daytime temperatures of
70°- 85°F, multiple heat
sources can be used
including incadescent
lights and ceramic heat
bulbs.
2. In regions where
Kinixys can be
maintained outside,
enclosures with high
walls should be utilized
as these tortoises are
capable of climbing.
most individuals to achieve their desired temperature for activity. It is still
encouraged that a heat source be maintained by overhead incandescent lights
though heat can also be can be supplemented with radiant heat emitters,
ceramic bubs, or under-the-tank heating pads. If using a heating-pad, care
should be taken to avoid overheating of the pad and surrounding material. A
gradient of temperature should be provided with the basking spot on one end
of the enclosure to allow the tortoises to move around and thermoregulate.
The environment should be kept at low-light to simulate their natural
conditions of dappled light on a forest floor or shaded from vegetation to
help reduce stress. A full spectrum compact florescent light should be used
to provide as natural of light as possible. Fresh water must be provided and
can be presented in a dish or small animal automatic waterer. The latter,
commonly used for raising poultry, reduces contamination of the water by
urine and feces and is easy to clean and maintain. Tortoises will quickly learn
to use a waterer and the water reservoir on top helps prevents the water dish
drying out.
Seasonal or Permanent Outdoor Enclosures
If climate permits, outdoor enclosures can be utilized. Kinixys are excellent
climbers and can burrow so the enclosure must be designed with this in
mind. Outdoor enclosures should be landscaped with plants safe for grazing,
such as dandelions, chickweed, and clover. Multiple hides and an adequate
water source must be provided. These tortoises enjoy soaking themselves and
will benefit from a place to soak including a shallow pool. Ample shaded areas
must be provided, especially for the rainforest species such as K. homeana
and K. erosa who often avoid direct sunlight. For the winter months in colder
climates, indoor housing will be necessary as these tortoises do not hibernate.
In some areas of the southwestern U.S. including Burbank, CA, Phoenix,
AZ, and El Paso, TX, K. spekii have been maintained outside year round. It is
presumed that K. lobatsiana and K. natalensis are able to remain outside year
2
round in this region although there are currently no known colonies of these
species. These species do become inactive in parts of their range during winter
and appear to overwinter in some areas. Due to the relatively low numbers of
these animals in assurance colonies this activity should be monitored closely and only attempted
with surplus animals.
Diet
Kinixys are omnivorous and should be provided with a varied diet including high calcium
green leafy plant matter, fungi, vegetables, high fiber fruits, and animal protein. Kinixys relish
earthworms, are known to consume snails, and most will also aggressively take thawed feeder
mice. Young can also be given a variety of invertebrates such as crickets, mealworms, and
waxworms for protein and enrichment. Leafy weeds and greens like kale, chicory, clover, endive,
dandelion, escarole, mustard greens and red leaf lettuces are preferred. Note that rainforest
species of Kinxys, K. homeana and K. erosa, are more reluctant to feed on greens in general and
Christine Light
121
Terry E. Kilgore
1
1-3. Hingeback tortoises
are omnivorous and
should be fed a variable
diet that can include
snails and other live
food, mushrooms, and
plant matter. Variety
is important and
observing animals eating
can help detect early
signs of health issues.
incorporating them into mixes has been more effective. Plant matter
like squashes, bell peppers, tomatoes, and zucchini can be offered along
with vegetables like carrots and sweet potatoes. Plant matter that is
high in oxalic acid such as kale, spinach and turnip greens are safe to
feed as part of a varied diet. If fed in large quantities over a prolonged
period of time, oxalic acid can be harmful as it binds with calcium and
prevents absorption of it. To avoid calcium depletion when feeding out
these items, calcium supplement can be a beneficial addition to their
diet. Hatchlings in particular should have a diet low in oxalic acid in
order to ensure they are absorbing all the calcium they need. It has been
observed that young Kinixys raised on a higher percent protein diet
have healthier growth rates and tend to be more active. In addition to
earthworms and other invertebrates, another excellent source for protein
is mushrooms which Kinixys seem to ubiquitously love.
Studies of wild and captive Kinixys have shown mushrooms to be an
important part of their diet as fungi are high in protein and preferred by
this omnivorous tortoise genus. We have observed the greatest growth
rate during periods where mushrooms make up a larger percent of their
diet. For our collection this has been timed to the increase in misting
to simulate the rainy season and the period when mushrooms would
be most readily available. Mushrooms are best offered quartered unless
enrichment is an objective and they can also be scattered whole in an
enclosure to promote natural grazing behaviors.
Animal proteins can be best provided through mice, pinkie mice, snails,
or live earthworms. Live invertebrates form an important part of the
2
wild Kinixys diet and they are known predators of millipedes, however
such a treat is difficult to provide in captivity. Cooked liver or poultry
can be offered sparingly, but raw beef should be avoided.
Cris Hagen
Fruits in general are best fed sparingly or as an occasional treat, as many are high in sugar and
low in fiber, leading to impaired gut function and increased lactic acid levels that promote
parasite activity. A high fruit diet is composed of sugars that ferment quickly in the gut and
compromise the natural gut flora in the process,
potentially leading to complications such as liver
abscesses. High sugar and low fiber fruits such
as berries, ripe bananas, and melons should be
avoided or used sparingly especially with wild
collected animals. However, since Kinixys are
attracted to brightly colored foods; if a tortoise is
a picky or reluctant eater, brightly colored food
items can be an enticing treat to tempt them into
eating. Kinixys can be introduced to a balanced
3
diet by slowly mixing treats that they will eat into
more nutritious and varied foods. Commercial tortoise chow,
such as Mazuri, can also be offered and are eagerly consumed
even by recently imported animals. The traditional Mazuri diet
lacks the desired fiber content and this can be augmented by use
of fresh alfalfa or similar items that can be blended into the food.
Newer products designed for tortoises have a higher fiber content
though it has been observed that they do not as aggressively
consume this more balanced diet. Commercial turtle chow,
while high in protein and not normally recommended for other
tortoise genera, can be mixed into Kinixys meals on occasion as
these tortoises maintain a higher protein diet.
1
Will Ahrens
Nutritional supplements can be a worthwhile addition to the
Kinixys diet. A fiber source, such as dried alfalfa helps to increase
fiber content especially during feedings that may include lower
fiber fruits. Commercial supplements like Oxbow’s FIBRevive
can also be added to meals and provides numerous benefits as
well as a pleasant and appetizing smell to attract Kinixys to food.
A calcium/vitamin D3 supplement powder can also be added
during periods when animals are maintained indoors though
used sparingly. The diet of hatchling and juvenile tortoises can
be supplemented more frequently with up to 3 times per week
depending on diet though over supplementing can result in
issues for both adults and young. Cuttle bone should always
be available and is quickly consumed in the breeding season.
2
Providing this material as a source of calcium in enclosures for
young tortoises helps prevent development of overgrown beaks
and provides a source for enrichment.
1. It is important to
monitor the feeding
of captive colonies so
that any behavioral or
physical issues can be
addressed immediately.
2. To avoid
complications from
impaction of ingested
mulch or sand, it is
suggested that captive
animals be fed on paper
plates or trays.
Care should be taken when feeding to avoid direct contact with substrate to avoid potential
impaction especially when sand is incorporated. Paper plates can be used as a simple and quick
feeding surface, though on occasion Kinixys may eat part of the plate if juices or color from food
items saturates it. Disposal is also an issue and we recommend composting these when possible.
Plastic trays are an effective alternative though great care must be taken to properly clean and
disinfect them.
Health
Kinixys maintain a reputation as being incredibly difficult animals to establish and maintain in
captivity. If dealing with recent imports, the quicker that individual can be set up and stabilized
the better. Hydration is key to setting up newly arrived tortoises. These tortoises are often
severely dehydrated after the long journey that typically includes multiple trips from a collector,
to an exporter, to an importer, to a pet shop, and finally an assurance colony. Immediate and
frequent soakings are essential in helping to stabilize Kinixys. We suggest a somewhat hands off
approach and to handle animals only when necessary. Recent acquisitions should be maintained
in low-light enclosures with plenty of hides and cover to help reduce stress. Kinixys also display
123
1. Thorough health
inspections and
substantial quarantine
periods are critical
when working with
newly imported
tortoises. Periodic
evaluations will help
track health and
animal condition.
Alain Soula
2. Inspections of new
animals for parasites
is vital to maintaining
healthy colonies.
Ticks are common
among Kinixys as
seen on this K. spekii
from the Masai Mara
Manyatta Camp in
Kenya.
certain natural physiological behaviors
that can be perceived as a health issues
including watery eyes and wheezing or
whistling sounds. Though both can be
signs of stress or illness, healthy animals
will have somewhat naturally watery
eyes and when excited or anxious (such
as when food is placed in front of them)
they will breathe heavily making a sort
of wheezing sound. Continued vigilance
and observation of Kinixys is important to
detect signs of distress including lethargy,
loss of appetite, discharge from nose or
mouth, swelling of eyes, or feces containing
parasites or blood.
1
Health conditions are often treatable and can be prevented by proper husbandry and early
detection. It is imperative that all new tortoise arrivals be quarantined from existing populations to
prevent cross contamination. Different species should also never be housed together. Wild caught
Kinixys are particularly prone to heavy parasite loads of worms and protozoa, although a proper
diet can maintain healthy gut flora and decrease the severity of parasites. It is important to note
that small numbers of worms and protozoa may be normal gut residents potentially providing
some benefits to the animal. In stressed or immune compromised animals
this balance can become disrupted. Though both are problematic and can
result in the decline or death of an animal, protozoans have likely historically
been the major killer of imported Kinixys. Diet can be key in minimizing and
controlling these organisms. Evaluation of the need for treatment for parasites
should be done on a case-by-case basis and under the supervision of a qualified
veterinarian with demonstrated experience working with tortoises. Excessive
or unnecessary treatment of Kinixys for parasites may damage gut health and
efficiency. Coccidia can be particularly problematic and lethal with this genus
and detection of it prior to any treatment for other parasites is important.
Examining the stool is a good indication of how balanced the tortoise’s diet
is. Scat should be firm and contain little undigested food. Diarrhea and loose
stool indicates a diet that is possibly too low in fiber, too high in fruit or water
content, overfeeding, or illness.
2
A general indication of healthy weight is that a tortoise should feel solid
when lifted. If it feels light and hollow, its weight will need attention. Any
unusual or unexpected changes in weight should be taken seriously and the
animal closely monitored for normal eating behaviors. The shell of a Kinixys
is another important health indicator. If a shell is pyramiding--extra keratin
building up on the scutes resulting in a lumpy or pyramided effect--it could
be an indication of a number of complications. Shell pyramiding can be a sign
1. Animals should be
monitored for any changes in
health including swollen or
irritated eyes.
2. Captive males can become
territorial in the presence
of females and care should
be taken to separate males
when signs of aggression are
shown.
3. Nests can be quite shallow
and enclosures should be
monitored for recently laid
eggs as species, including K.
spekii, have been observed
eating their own eggs and
those of other females.
of too much protein in a diet, a possible
lack of calcium or vitamin D3 (naturally
obtained through sunlight), or even an
indication of being overfed or kept at
too cool of a temperature. Insufficient
humidity and substrate moisture can
also impact shell growth and condition
1
resulting in the appearance of lumpiness
in some species or a sunken appearance of
scutes for others. Excessive moisture and
soggy substrate can lead to potential fungal infections and shell rot. Inspection of the shell
growth and appearance is part of the necessary regular inspections for these tortoises.
The eyes of Kinixys should be bright, clear and wide. Swelling of the eyes or any discharge
could be an indication of infection, a massive parasite load, or of a foreign body lodged
around the eyes or nares(nostrils). As previously noted, Kinixys are naturally prone to
weepy eyes. This has also been observed to occur particularly with gravid tortoises and this
shouldn’t be seen as a health concern. Watery eyes in general could also be indicative of
being kept in too dry of an environment.
Aggression can and often will occur especially among males due to territorial
disputes or over females. Sometimes severe trauma, especially to the front
limbs and face, can result. These acts of dominance may also result in
behavioral changes in subordinate Kinixys and could affect overall health.
Flicker User - Devil Fang
Rodney Lewis
Reproduction
2
3
Maintaining multiple male Kinixys in an enclosure can be challenging
depending on the space available, amount of cover, access to females, and
time of year. Male Kinixys tend to be territorial and will battle with other
individuals, especially other males during breeding season. It is reported that
K. erosa males are the most aggressive of the genus, though K. nogueyi have
also demonstrated severe aggression and inflicted extensive wounds. This
tendency may make it difficult to keep more than one male Kinixys in an
enclosure. This aggressive behavior during the breeding season also warrants
observation of females for increased stress from continuous advances.
Courtship in Kinixys includes the male nudging and shoving the female
for a short period of time while attempting to mount. Males will also on
occasion nip at females. The shape of the carapace presents some difficulties
to successful mating, especially in K. homeana, where the carapace comes to
a sharp point and then drops off at a right angle down to the tortoise’s tail.
When successful, a male will climb on to a female and lower his back until
touching the ground. With their front feet dangling or resting on a female’s
carapace, males will move their tail under the carapace of the female. Once
successfully mounted, mating between Kinixys can last for an hour or
125
Will Ahrens
Mallory Clark
more. During this time, males will stretch their necks and with a wide
gaped mouth, emit a number of high pitched grunts. Females have been
observed to pseudocopulate, presumably to assert dominance as well
as to potentially attract the attention of nearby males. With K. spekii,
this behavior along with aggressive pseudocourtship activities has been
observed in gravid females especially those looking for nesting sites.
1.Radiographs are a
beneficial tool when
assessing the health
and reproductive state
of captive and newly
imported tortoises.
2. Kinixys hatchlings
can grow at incredible
rates. These K. homeana
hatchlings are roughly a
month apart in age.
3. It is believed that
all Kinixys species
have developmental
diapause. Future work
with assurance colonies
and field work with
Kinixys will help better
determine the extent
of this developmental
characteristic and how
to break it.
1
2
Following successful mating it can take two to three months for a female
to fully develop the eggs and nest. Nesting can occur year round for some
species, though it tends to occur from November through May, which
coincides with the rainy seasons in their respective ranges. Nests can be
rather shallow for species like K. homeana, or at depths of 10 or more
inches as in K. spekii. Clutch sizes vary from 3 to 8 eggs for most species,
though K. erosa can lay up to 10 eggs per clutch. Kinixys can lay multiple
clutches under optimal conditions.
Kinixys, being omnivorous and opportunistic, will eat their own eggs
and those of other females. Both K. spekii and K. homeana have been
observed consuming eggs. On at least one occasion a captive female
K. spekii was observed watching another female K. spekii nesting and
attempted to consume her eggs.
Incubation time is variable as it is believed all Kinixys have at least
some level of diapause. The lowest observed was one month for K. homeana and K. nogueyi
(both tropical species) and the longest in captivity was five months for K. spekii. This time is
in addition to the 160 or so days for incubation once diapause is broken. In Swaziland, K.
spekii in captivity took nearly a year to hatch in an outdoor enclosure (Boycott and Bourquin
2000). Incubation substrate varies on personal preference and experimentation. Known
substrates have included: vermiculite, perlite, mulch, and hydroponic clay pellets. Humidity
is important, but too much and the eggs will crack as with many tortoise species. Incubation
temperatures have ranged from 80 to 90°F and
humidity levels average 80%. It is presumed
that Kinixys display Temperature-dependent Sex
Determination (TSD), but studies are lacking.
Incubation at higher ends of the range can result
in developmental problems including extra scutes
and in rare situations eyeless tortoises.
Hatchlings can take up to four days to fully
emerge from eggs and absorb their yolk sac. It
is important to soak these tortoises as they can
drink up to 30% or more of their body weight
in water during their first soak. Some individuals
have been observed to eat within a day or so of
emerging. Care should be taken when housing
3
1. Enclosures with
multiple hatchlings
should be monitored
for signs of territorial
disputes as smaller
individuals can
become stressed and
stop feeding.
2.-3. When kept in
proper conditions
and monitored
closely, it is possible
to maintain healthy
and robust captive
groups of Kinixys.
The establishment of
both ex situ and in situ
assurance colonies
will likely be a critical
component in the
conservation of this
genus in the future.
1
multiple young animals as territorial disputes can arise and smaller animals may stop feeding and
become highly stressed. Housing of juveniles of various sizes is discouraged unless ample space,
hides, and feeding areas are available. Growth can be fairly rapid and hatchlings which are about
1.5 inches can double in as little as three months for K. spekii and 5 months for K. nogueyi.
Maintenance
The mulch of enclosures should be turned by hand on an approximate monthly basis with priority
being placed on enclosures of nesting tortoises. Doing so aids in distribution of moisture and
aeration of the substrate which often facilitates the nesting process and encourages gravid females
to nest by providing quality substrate. Soaking tortoises during this task is an
effective way to reduce the stress of disturbance and providing supplemental
hydration and inspection. Spot cleaning the enclosures daily will help prevent
contamination between animals, which includes removing feces and old,
uneaten food. Colonies of pillbugs and springtails may naturally colonize
enclosures (especially if leaf litter is incorporated) and may help remove waste
and food scraps that are not collected to provide a more naturally healthy
indoor system. Water sources should be flushed and refilled every other day or
as needed to prevent contamination as these tortoises tend to soil their water
with substrate or feces. Take care to completely disinfect water dishes weekly, a
2
10% bleach solution is recommended for this.
The general appearance of enclosures should be closely maintained. Any
hides that are flipped or false plants that are uprooted should be resituated in
order to ensure the quality of the habitat and maintain low-stress levels. Spot
clean any plants or hides that may have gotten feces on them. Ensure that all
lights are properly situated and replace any bulbs that have burned out. It is
recommended that UV bulbs be changed every 6 months as the UV output
declines over time.
3
127
Appendix E: Kinixys Questionnaire
Goals: Quantify who is working with Kinixys in private and public
collections; Determine what species they currently or have historically
maintained and quantify total number and sex ratios; Assess success
with keeping and breeding Kinixys; Determine level of interest in
working more with Kinixys; Inquire about husbandry, breeding, and
incubation techniques; Engage questionnaire participant in becoming
active participants in Kinixys Conservation Task force and assurance
colonies; Develop comprehensive database of Kinixys keepers,
husbandry and breeding data.
Owner/Keeper Information: Name __________________ City/
State __________________ Email ________________
General species information
1) What types of hingebacks (Kinixys) do you have and what are their
genders? If you are unsure, would you be willing to send us pictures so
we can identify species and sex?
2) How long have you had your hingebacks?
3) Did you get them as adults? __ yes __ no
4) Have you had other hingebacks in the past or that are no longer
alive? __yes __ no
If yes, could you please indicate their type, sex, how long you had them,
and why they died (if you know) or if you got rid of them:
5) Do you keep other tortoises? __ yes __ no
If yes, what species do you keep:
6) Were your animals wild caught or captive bred? If both please
explain.
7) In an effort to catalogue variability in patterns by species would you
be willing to share photos of your animals?
8) Are you willing to include your animals in a studbook and
participate in conservation program?
5) What range of humidity levels do you maintain?
6) What temperature range do you maintain?
7) Do you maintain males and females together year round or separate
seasonally?
General health information
1) Has your hingeback(s) ever experienced medical issues? __yes __no
If yes, what were the issues and were they resolved?
2) Is your hingeback(s) seen by a veterinarian?
__ yes, routinely __ yes, but only when medical attention is required __
no
Diet
1) How often do you feed your hingeback(s)?
2) What types of food do you feed your hingeback(s)?
3) How have you been successful with addressing picky eaters?
4) Do you add supplements to your hingeback(s) food?
Breeding
With the following questions, if more than one female was involved,
please give the number of females and if possible how many each
female produced.
1) Have your hingeback(s) breed in the past or are they breeding now?
__yes __no
If yes:
1) Were any eggs produced? __ yes __ no If yes, how many eggs were produced? 2) Were any hatchlings produced? __ yes __ no
If yes, how many hatchlings were produced?
3) What temperatures did you incubate?
9) Are you willing to participate in exchanging animals for genetic
diversity?
4) Did you incorporate a diapause (delayed development) into
incubation?
General enclosure information
1) How do you house your hingeback(s)? __ Inside only __ Outside
only __Both inside and outside (please explain)
5) What type of incubator/incubation technique did you use?
2) What size is the enclosure(s)?
Summary Information
1) Would you like more information on hingebacks? __yes __ no
3) What type of substrate or bedding do you use?
2) Can we contact you in the future for more information about your
hingebacks? __yes __ no
4) What light cycles do you use? Do you use UVB lighting on them?
__yes __no
3) Would you be interested in becoming more active in hingeback
husbandry? __ yes __ no
Appendix F: Kinixys Activity Chart
129
130
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