Kleinmann's Tortoise
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Kleinmann's tortoise (''Testudo kleinmanni''), also called commonly the Egyptian tortoise, Leith's tortoise, and the Negev tortoise, is a critically endangered
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of cryptodire turtle in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Testudinidae Tortoises ( ) are reptiles of the family Testudinidae of the order Testudines (Latin for "tortoise"). Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like ot ...
. The species is native to
Libya Libya, officially the State of Libya, is a country in the Maghreb region of North Africa. It borders the Mediterranean Sea to the north, Egypt to Egypt–Libya border, the east, Sudan to Libya–Sudan border, the southeast, Chad to Chad–L ...
and possibly extinct in
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
. The species was once more widespread, but its numbers are now dwindling, and complete extinction in the wild is a looming threat unless more actions are taken to protect this species.


Etymology

The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, ''kleinmanni'', is in honor of , a French stockbroker who collected the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
in 1875. The specific name, ''werneri'' (of the synonym ''Testudo werneri)'', is in honor of Israeli herpetologist Yehudah L. Werner.


Description

Kleinmann's tortoise is the smallest tortoise in the Northern Hemisphere. Females are larger than males; males are more slender and have a longer tail. The
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
has a high dome, and ranges in color from ivory to pale gold to dark brown to pink or dull yellow. This allows the paler tortoise to stay in the desert heat for longer. It is also an effective
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
in the desert. The
plastron The turtle shell is a shield for the ventral and dorsal parts of turtles (the Order (biology), order Testudines), completely enclosing all the turtle's vital organs and in some cases even the head. It is constructed of modified bony elements such ...
is light yellow, often with two dark triangles on each abdominal
scute A scute () or scutum (Latin: ''scutum''; plural: ''scuta'' "Scutum (shield), shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of Bird anatomy#Scales, birds. The ter ...
. The plastral scutes have dark edgings that fade with age. The head and limbs are a very pale ivory-yellow to yellowish-brown colour.


Systematics

The proposed
subgenus In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
''Pseudotestudo'' is invalid, based on immature characters. It has been proposed to unite this species with the
marginated tortoise The marginated tortoise (''Testudo marginata'') is a species of tortoise in the family Testudinidae. The species is endemic to Greece, Italy, and the Balkans in Southern Europe. It is the largest European tortoise. The marginated tortoise is her ...
in the genus ''Chersus''. These clearly share a common ancestor with the common tortoise (Greek tortoise). The former two are somewhat more similar to each other than to the Greek tortoise regarding
DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...
data. Considering biogeography, however, this is either due to (rather unlikely) dispersal across the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
, or the supposed "
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
" is invalid and the similarity due to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
.


Negev tortoise

The
Negev The Negev ( ; ) or Naqab (), is a desert and semidesert region of southern Israel. The region's largest city and administrative capital is Beersheba (pop. ), in the north. At its southern end is the Gulf of Aqaba and the resort town, resort city ...
subpopulation In statistics, a population is a set of similar items or events which is of interest for some question or experiment. A statistical population can be a group of existing objects (e.g. the set of all stars within the Milky Way galaxy) or a hypo ...
had been separated as a distinct species, "Negev tortoise" (''Testudo werneri)'', as it did not appear to have distinct or strongly reduced
haplotype A haplotype (haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material (DNA) which is inherited from two parents. Normally these organisms have their DNA orga ...
diversity, consistent with the recent extinction of the Egyptian population and slow DNA sequence evolution rates in ''Testudo''. Baha el Din (2006) synonymized ''T. werneri'' due to lacking mtDNA differences between ''T. kleinmanni'' from western Libya compared to tortoises from the Negev Desert. In the 2017 checklist of turtles of the world, it remained a synonym. Image:Carapace T.werneri destra, T.kleinmanni sinistra.JPG, Characteristic specimens of the Egyptian tortoise (left) and Negev tortoise (right), dorsal view Image:Piastrone T. werneri destra , T.kleinmanni destra.JPG, Ventral view: Differences are slight and unreliable (see above)


Habitat and ecology

Kleinmann's tortoise lives in
desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...
s and semiarid
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
s, usually with compact sand and gravel plains, scattered rocks, shallow, sandy
wadi Wadi ( ; ) is a river valley or a wet (ephemerality, ephemeral) Stream bed, riverbed that contains water only when heavy rain occurs. Wadis are located on gently sloping, nearly flat parts of deserts; commonly they begin on the distal portion ...
s, dry woodlands, shrubby areas, and coastal
salt marsh A salt marsh, saltmarsh or salting, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastal intertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. I ...
habitats. In captivity, it eats grasses, fruits, and vegetables, but the diet of ''T. kleinmanni'' in the wild is unknown. Kleinmann's tortoise tend to live in habitats that receive less than 50 mm of rain every year. These habitats tend to be countries with expansive desert areas such as Egypt and Libya. It is least active when the weather is very cold or very hot. During the colder months, it is out most during midday. During the warm season, it is active in the mornings and evenings. The rest of the day is spent under bushes or in
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
burrow file:Chipmunk-burrow (exits).jpg, An eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of Animal lo ...
s.


Reproduction

Kleinmann's tortoise becomes sexually mature when about 5 years old. In the wild, mating has only been observed in March, but in captivity, it mates in April and August to November. During courtship, the male will ram the female, sometimes chasing after her. Unlike any other Mediterranean tortoise, ''T. kleinmanni'' may make a
mating call A mating call is the auditory signal used by animals to attract mates. It can occur in males or females, but literature is abundantly favored toward researching mating calls in females. In addition, mating calls are often the subject of mate choi ...
similar to the call of the
mourning dove The mourning dove (''Zenaida macroura'') is a member of the dove Family (biology), family, Columbidae. The bird is also known as the American mourning dove, the rain dove, the chueybird, colloquially as the turtle dove, and it was once known a ...
.
Eggs An egg is an organic vessel in which an embryo begins to develop. Egg, EGG or eggs may also refer to: Biology * Egg cell, the female reproductive cell (gamete) in oogamous organisms Food * Eggs as food Places * Egg, Austria * Egg, Switzerland ...
are laid in shallow bowls beneath bushes, or in vacant burrows. Each
clutch A clutch is a mechanical device that allows an output shaft to be disconnected from a rotating input shaft. The clutch's input shaft is typically attached to a motor, while the clutch's output shaft is connected to the mechanism that does th ...
contains one to five eggs, which hatch in the summer or early autumn.


Status and conservation

Once found in Egypt,
Libya Libya, officially the State of Libya, is a country in the Maghreb region of North Africa. It borders the Mediterranean Sea to the north, Egypt to Egypt–Libya border, the east, Sudan to Libya–Sudan border, the southeast, Chad to Chad–L ...
and
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
/
Palestine Palestine, officially the State of Palestine, is a country in West Asia. Recognized by International recognition of Palestine, 147 of the UN's 193 member states, it encompasses the Israeli-occupied West Bank, including East Jerusalem, and th ...
, the
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
of ''T. kleinmanni'' in Egypt has been all but destroyed, and the Egyptian tortoise is close to complete extinction there; in 2006, only 10 wild individuals were known from the vicinity of
Lake Bardawil Lake Bardawil ( or ), is a large, very saline lagoon nearby the protected area of Zaranik (also known for diversities of insects and waterbirds) in Egypt on the north coast of the Sinai Peninsula. Lake Bardawil is about long, and wide (at its ...
. Two populations can still be found in Libya, but much of the coastline habitat has been destroyed because of human activity. Habitat loss and the illegal pet trade are huge issues facing the species; it is also hunted locally for use in
folk medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
. The population is still on the decline, and the risk of extinction in the wild is very real if habitat degradation and illegal trade continue at their present rate. In
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
, ''T. kleinmanni'' (previously known as ''T. werneri'') populations are known from northwestern
Negev The Negev ( ; ) or Naqab (), is a desert and semidesert region of southern Israel. The region's largest city and administrative capital is Beersheba (pop. ), in the north. At its southern end is the Gulf of Aqaba and the resort town, resort city ...
and the Yemin Plain, with lone individuals occasionally found near
Yeruham Yeruham () is a local council (Israel), town in the Southern District (Israel), Southern District of Israel, in the Negev desert. It covers , and had a population of in . It is named after the Hebrew Bible, Biblical Jeroham. Until early 2011 th ...
. It is protected by law and several conservation programs, such as the establishment of new
protected area Protected areas or conservation areas are locations which receive protection because of their recognized natural or cultural values. Protected areas are those areas in which human presence or the exploitation of natural resources (e.g. firewood ...
s, but is still highly threatened by the conversion of habitat to agricultural fields, extensive animal herding, and the approval of new developments on the
sand dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
of the western Negev. In 2010, the population in
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
/
Palestine Palestine, officially the State of Palestine, is a country in West Asia. Recognized by International recognition of Palestine, 147 of the UN's 193 member states, it encompasses the Israeli-occupied West Bank, including East Jerusalem, and th ...
was estimated at less than 2,000 individuals. On the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological ...
, Kleinmann's tortoise is classified as
critically endangered An IUCN Red List critically endangered (CR or sometimes CE) species is one that has been categorized by the International Union for Conservation of Nature as facing an extremely high risk of extinction in the wild. As of December 2023, of t ...
(CR A2abcd+3d). Less than three ''Testudo'' generations ago, an estimated 55-56,000 adult Kleinmann's tortoises existed. Today, about 7,500 remain, and decline due to
habitat destruction Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
and unsustainable exploitation appears to be ongoing. While the former threat is believed to be alleviated, illegal pet trade is feared to have reduced the population to maybe 1,000 adult tortoises in the two or three remaining subpopulations. Given that ''T. kleinmanni'' is a slow-maturing, long-lived species with few offspring (see
K-strategist In ecology, selection theory relates to the selection of combinations of traits in an organism that trade off between quantity and quality of offspring. The focus on either an increased quantity of offspring at the expense of reduced individual ...
), this is well possibly less than the
minimum viable population Minimum viable population (MVP) is a lower bound on the population of a species, such that it can survive in the wild. This term is commonly used in the fields of biology, ecology, and conservation biology. MVP refers to the smallest possible si ...
size, eventually dooming the species to extinction in the wild. There are plans for augmenting the Israeli sub-population by releasing individuals from local controlled breeding centers in Israel, but these projects have yet to be approved by authorities. Mixing individuals from the subpopulations for
reintroduction Species reintroduction is the deliberate release of a species into the wild, from captivity or other areas where the organism is capable of survival. The goal of species reintroduction is to establish a healthy, genetically diverse, self-sustain ...
purposes should be avoided, as the ecological parameters of habitat differ.
DNA fingerprinting DNA profiling (also called DNA fingerprinting and genetic fingerprinting) is the process of determining an individual's deoxyribonucleic acid (DNA) characteristics. DNA analysis intended to identify a species, rather than an individual, is cal ...
of individuals to help maintain
heterozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
in captive and reintroduction populations can be performed during routine stool analyses. In this context, notably, there may be a
haplotype A haplotype (haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material (DNA) which is inherited from two parents. Normally these organisms have their DNA orga ...
uniquely found in the Egyptian tortoise. Captive breeding requires more care than in other ''Testudo'' species, as the species is more delicate and the clutch is very small, but is not highly difficult for experienced ''Testudo'' breeders. ''T. kleinmanni'' is not generally available for hobbyists and even if legal to keep, should be avoided without valid documentation. Smuggling continues to be a problem, but confiscated individuals are used to establish a captive safeguard population. On May 21, 2007, Rome's main zoo, Bioparco, reported it has successfully bred the species from parents rescued from a smuggler's suitcase in 2005. It is also kept and bred at several other zoos in Europe and North America. Image:Testudo kleinmanni&werneri1.jpg, Intercepted shipment of Kleinmann's tortoise from Libya for the illegal pet market, port of
Genoa Genoa ( ; ; ) is a city in and the capital of the Italian region of Liguria, and the sixth-largest city in Italy. As of 2025, 563,947 people live within the city's administrative limits. While its metropolitan city has 818,651 inhabitan ...
, 2005. Image:Testudo kleinmanni&werneri8.jpg, ''T. kleinmanni'' now in Bioparco: Smuggling conditions are typically worse. Image:Testudo kleinmanni&werneri3.jpg, ''T. kleinmanni'' from Libya. Image:Testudo werneri.jpg, Kleinmann's tortoise,
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
fractured during smuggling.


Threats

Kleinmann's tortoise confronts a range of anthropogenic challenges affecting its conservation. These threats include agricultural practices, overgrazing, industrial development, and, notably, the illegal pet trade. The species suffered a substantial setback when Egyptian subpopulations faced extinction, leading to the utilization of the Libyan stock in both national and international pet markets, a practice that eventually became illegal. Kleinmann's tortoise encounters threats from agricultural activities, overgrazing, and industrial development. These pressures contribute to habitat degradation, posing challenges to the species' survival and exacerbating its vulnerability. A critical concern for Kleinmann's tortoise is the illegal pet trade, which originated with the use of Libyan stock after Egyptian subpopulations were depleted. The illicit trade persists within Egypt and Libya. This trade has potentially devastating consequences for the global population, with collection pressure reported to be higher in the eastern regions. Kleinmann's tortoise faces heightened vulnerability due to its low annual biomass production, resulting in a high sensitivity to population disturbance. The species exhibits poor recovery abilities from activities impacting the population size negatively.


References


External links


ARKive species informationBristol Zoo species informationRome zoo breeds rare Egyptian tortoises
{{Taxonbar, from=Q253781 Testudo (genus) Turtles of Africa Critically endangered fauna of Africa Species that are or were threatened by the pet trade Species that are or were threatened by agricultural development Species that are or were threatened by grazing Taxa named by Louis Charles Émile Lortet Reptiles described in 1883