Kleinmann's tortoise (''Testudo kleinmanni''), also called commonly the Egyptian tortoise, Leith's tortoise, and the Negev tortoise, is a critically endangered

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...

of neck-hiding tortoise in the

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Testudinidae. The species is native to

Egypt Egypt ( ar, مصر , ), officially the Arab Republic of Egypt, is a List of transcontinental countries, transcontinental country spanning the North Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via a land bridg ...

Libya Libya (; ar, ليبيا, Lībiyā), officially the State of Libya ( ar, دولة ليبيا, Dawlat Lībiyā), is a country in the Maghreb region in North Africa. It is bordered by the Mediterranean Sea to the north, Egypt to the east, Suda ...

, and

Israel Israel (; he, יִשְׂרָאֵל, ; ar, إِسْرَائِيل, ), officially the State of Israel ( he, מְדִינַת יִשְׂרָאֵל, label=none, translit=Medīnat Yīsrāʾēl; ), is a country in Western Asia. It is situated ...

. The species was once more widespread, but its numbers are now dwindling. The species is nearly extinct in Egypt, and complete extinction in the wild is a looming threat unless more actions are taken to protect this species.

Etymology

The specific name, ''kleinmanni'', is in honor of Edouard Kleinmann, a French stockbroker who collected the

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...

in 1875.Beolens, Bo; Watkins, Michael; Grayson, Michael (2011). ''The Eponym Dictionary of Reptiles''. Baltimore: Johns Hopkins University Press. xiii + 296 pp. . (''Testudo kleinmanni'', p. 143; ''T. werneri'', p. 282). 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. Female tortoises are larger than the males; males are more slender and have a longer tail. The plastron has a high dome, and ranges in color from ivory to pale gold to dark brown to pink or dull yellow. This colouring strongly follows

Gloger's rule Gloger's rule is an ecogeographical rule which states that within a species of endotherms, more heavily pigmented forms tend to be found in more humid environments, e.g. near the equator. It was named after the zoologist Constantin Wilhelm Lambe ...

, as it helps to regulate the impact of sunlight. 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

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 ...

is light yellow, often with two dark triangles on each abdominal

scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "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 birds. The term is also used to describe the anterior po ...

. The tortoise's 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 (plural: 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 ...

''Pseudotestudo'' is invalid, based on immature characters.Perälä (2003) 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 he ...

in the genus ''Chersus''. These clearly share a common ancestor with the

common tortoise The Greek tortoise (''Testudo graeca''), also known commonly as the spur-thighed tortoise, is a species of tortoise in the family Testudinidae. ''Testudo graeca'' is one of five species of Mediterranean tortoises ( genera '' Testudo'' and '' ...

(Greek tortoise). The former two are somewhat more similar to each other than to the Greek tortoise regarding

DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. T ...

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 north by Western and Southern Europe and Anatolia, on the south by North Africa, and on ...

, or the supposed "

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...

" 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 com ...

Negev tortoise

The

Negev The Negev or Negeb (; he, הַנֶּגֶב, hanNegév; ar, ٱلنَّقَب, an-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 sout ...

subpopulation 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 o ...

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 barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...

s and semiarid

habitat In ecology, the term habitat summarises the array of resources, physical and 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 ...

s, usually with compact sand and gravel plains, scattered rocks, shallow, sandy

wadi Wadi ( ar, وَادِي, wādī), alternatively ''wād'' ( ar, وَاد), North African Arabic Oued, is the Arabic term traditionally referring to a valley. In some instances, it may refer to a wet (ephemeral) riverbed that contains water ...

s, dry woodlands, shrubby areas, and coastal

salt marsh A salt marsh or saltmarsh, 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. It is domin ...

habitats. In captivity, it eats grasses, fruits, and vegetables, but the diet of ''T. kleinmanni'' in the wild is unknown. 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 Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...

burrow 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 locomotion. Burrows provide a form of 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 choic ...

similar to the call of the

mourning dove The mourning dove (''Zenaida macroura'') is a member of the dove family, Columbidae. The bird is also known as the American mourning dove, the rain dove, and colloquially as the turtle dove, and was once known as the Carolina pigeon and Caroli ...

Eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...

are laid in shallow bowls beneath bushes, or in vacant burrows. Each

clutch A clutch is a mechanical device that engages and disengages power transmission, especially from a drive shaft to a driven shaft. In the simplest application, clutches connect and disconnect two rotating shafts (drive shafts or line shafts). ...

contains one to five eggs, which hatch in the summer or early autumn.

Status and conservation

Once found in Egypt,

and

, the

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. 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) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...

. 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

, ''T. kleinmanni'' (previously known as ''T. werneri'') populations are known from northwestern

and the Yemin Plain, with lone individuals occasionally found near

Yeruham Yeruham ( he, יְרוֹחַם, ''Yeroham'') is a town ( local council) in the Southern District of Israel, in the Negev desert. It covers 38,584 dunams (~38.6 km²), and had a population of in . It is named after the Biblical Jeroham. Un ...

. 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, ecological or cultural values. There are several kinds of protected areas, which vary by level of protection depending on the ena ...

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, fl ...

of the western Negev. In 2010, the population in

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 the world's most comprehensive inventory of the global conservation status of biolo ...

, Kleinmann's tortoise is classified as critically endangered (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 and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...

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), 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 siz ...

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-sustainin ...

purposes should be avoided, as the ecological parameters of habitat differ.

DNA fingerprinting DNA profiling (also called DNA fingerprinting) is the process of determining an individual's DNA characteristics. DNA analysis intended to identify a species, rather than an individual, is called DNA barcoding. DNA profiling is a forensic t ...

of individuals to help maintain heterozygosity in captive and reintroduction populations can be performed during routine stool analyses. In this context, notably, there may be a

uniquely found in the Egyptian tortoise.Široký & Fritz (2007) 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 ( ; it, Genova ; lij, Zêna ). is the capital of the Italian region of Liguria and the sixth-largest city in Italy. In 2015, 594,733 people lived within the city's administrative limits. As of the 2011 Italian census, the Province of ...

, 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,

fractured during smuggling.

References

External links

* (''Testudo werneri'', new species). * Listed as Critically Endangered (CR A2abcd+3d v3.1). * * *
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 endangered by the pet trade Species endangered by agricultural development Species endangered by grazing Taxa named by Louis Charles Émile Lortet Reptiles described in 1883