An Annotated Bibliography of the Egyptian Tortoise, Testudo kleinmanni Lortet 1883
by James Buskirk
In the last 5 years, hundreds, possibly thousands, of Egyptian tortoises (Testudo kleinmanni) have
been exported from Cairo to the United States. Many of these captives fare badly, dying after only weeks or months in
confinement. Captive breeding has been exceptionally infrequent. It is hoped that the commercial exploitation of this
species will soon cease. Most tortoises wind up in the hands of individuals who are likely to have access to this
publication. In the meantime, in an effort to be of assistance regarding the husbandry of T. kleinmanni, the
following information is offered.
The 36 literature references below exclude species accounts found in well-known popular and scientific
books on turtles and tortoises of the world. Most of them are abridgements of works listed here, and were authored by
researchers who have never examined a specimen of T. kleinmanni, living or dead. Also omitted are two Egyptian
biochemical treatises, which used the tissue of vast numbers of sacrificed specimens. Such works have no relevance for
captive maintenance, breeding, or conservation.
In works of more than a few pages not dealing exclusively with T. kleinmanni, only the pertinent
page numbers are given. Photographs, if present, are described.
Anderson, J. 1896. Herpetology of Arabia. SSAR reprint edition 1984. On page 68 the author cites Forskål's
posthumous report (1775) that a tortoise, which Anderson believes might be T. leithii (= T. kleinmanni)
occurs in what is now northwestern Yemen. Forskål, incidentally, is inadvertently responsible for the misused,
mismatched, misunderstood trinomial Testudo graeca terrestris and died of the plague in 1763.
Anderson, J. 1898. Zoology of Egypt: Reptilia and Batrachia. Bernard Quaritch, London. Pages 28-30 provide
an accurate descriptive account, comparing this tortoise to other Mediterranean tortoises and discussing its range,
relying considerably upon Lortet (1887). Plate II provides 3 excellent life-size figures.
Arbel, A. 1984. Testudo kleinmanni. Reptiles and Amphibians. Plants and Animals of the Land of Israel, an
illustrated encyclopedia. Volume 5, pages 37-38. Ministry of Defense Publishing House/Society for the Protection of
Nature in Israel. Largely accurate, this account provides diagnostic criteria distinguishing this tortoise from
T. graeca and summarizes its natural history. Arbel is the first writer to refer to the absence of dark markings
whatsoever on some specimens' plastrons; his photo of a lumpy, captive-bred juvenile is regrettable. In Hebrew.
Bodenheimer, F. S. 1935. L. Mayer Press, Jerusalem. Three sentences on page 197 deal with the presence of
this species, as T. leithii, in the Negeb Desert--the first report from what is now Israel since Tristram
Buskirk, J. 1985. The endangered Egyptian tortoise, Testudo kleinmanni: status in Egypt and Israel. Ninth
International Herpetological Symposium on Captive Propagation and Husbandry. McKeown, Caporaso & Peterson, eds.,
35-52. Summarizes natural history literature to date and shares author's generally negative experiences with
Egyptian zoo and wildlife officials. Incorporates personal communication from two leading Israeli T. kleinmanni
researchers, Geffen and Mendelssohn. Two figures, 1 map, extensive bibliography.
Calabresi, E. 1924. Missione Zoologica del Dr. E. Festa in Cirenaica. Bollettino dei Musei di Zoologia ed
Anatomia comparata della R. Universita di Torino 38:7-8. First record of the species for Libya, with brief
discussion of sympatricity with "T. ibera". The single specimen found had symmetrical supernumerary costal shields. In
Flower, S. S. 1926. Species of land tortoises of the genus Testudo found in the countries bordering the
Mediterranean Sea. Copeia, 133. Small is beautiful: Flower's concise tabular key to the 4 species remains accurate
today and is bereft of esoteric jargon and gobbledegook.
Flower, S. S. 1929. List of the vertebrated animals exhibited in the gardens of the Zoological Society of London,
1828-1927. Vol. 3 (Reptiles), p. 29. London. Six T. kleinmanni, donated by John Anderson, were presented to
the London Zoo on 5/26/1892. Flower was unaware that Calabresi (1924) had recorded this species from Libya.
Flower, S. S. 1933. Notes on the recent reptiles and amphibians of Egypt. Proc. Zoological Society of London,
743-749. For decades, the account of this tortoise outshone all prior and subsequent works. Author describes
individual variation in great detail but discounts geographic significance. He describes sexual dimorphism and
reproduction (including captive breeding), record sizes and weight, and compares the hatchling with that of T.
graeca, among other achievements.
Geffen, E. 1985. On the biology of the Egyptian tortoise (Testudo kleinmanni) in Holot Agur, Israel.
Master's thesis, Dept. of Zoology, Tel Aviv University. Eighty-three pages in Hebrew, with 4-page English summary.
The most complete document to date on the natural history of this tortoise. Optimal air temperatures for this species
lie between 71-76°F. A list of natural food items appears on p. 60. Loaded with information!
Geffen E. & H. Mendelssohn 1986. The sand tortoise in Israel--present and future. Teva v'Aretz (Nature & Land)
6:26-27. Presents the needs for conservation measures on behalf of this species after listing natural and human
threats to its survival in Israel. Map, photos. In Hebrew.
Geffen E. & H. Mendelssohn 1988. Home range use and seasonal movements of the Egyptian tortoise (Testudo
kleinmanni) in the northwestern Negev, Israel. Herpetologica, 44:354-359. Interprets use of home range of 9
tortoises equipped with radio transmitters over 4- to 18-month periods.
Geffen E. & H. Mendelssohn 1989. Activity patterns and thermoregulatory behavior of the Egyptian Tortoise
Testudo kleinmanni in Israel. J. Herpetology 23:404-409. Self-explanatory title, further discussing the
decline of this species throughout its range. This species is the only known temperate chelonian which is active in
winter and dormant in summer.
Geffen E. & H. Mendelssohn 1991. Preliminary study on the breeding pattern of the Egyptian tortoise, Testudo
kleinmanni, in Israel. Herpetological J., 1:574-577. A portable x-ray generator was used to confirm gravid
status of females after which gravid individuals were kept in a natural enclosure and released after nesting. Females
lay 2-3 clutches per year of 1-3 large eggs at 20-30 day intervals.
Groombridge, B. 1982. Egyptian Tortoise Testudo kleinmanni Lortet 1883. The IUCN Amphibia-Reptilia red
data book, Part 1, 133-136. Gland, Switzerland. "Indeterminate" status is given to this species. A critical
analysis of geographic records is followed by a discussion of habitat, density, predation, and the need for
conservation, based largely on information provided by H. Mendelssohn (see below).
Günther, A. 1869. Report on two collections of Indian reptiles. Proc. Zool. Soc. London, 500-507, figs. 1-4,
pls. xxxviii-xl. Original description of this tortoise as Testudo leithii, based on a specimen brought to
Günther from what is now Pakistan. How it got to Pakistan is still a mystery, as is its true origin.
Hoath, R. 1990. Wild for sale. Cairo Today--the magazine of Egypt. October 1990:60-64. In this alarming
account of the domestic exploitation of Egyptian wildlife, the author includes 2 disturbing photos of tortoises for
sale, piled several deep.
Khalil, F. & G. Haggag 1955. Ureotelism and uricotelism in tortoises. Journal of Experimental Zoology
130:423-432. By analyzing the quantity of nitrogen in tortoise urine, the authors established that this species
can excrete either urea, uric acid, or both as the main end-product of protein metabolism. The Egyptian tortoise
favors the excretion of uric acid when faced with dehydration, an adaptation of other arid-adapted tortoises as well.
Lortet, L. Testudo kleinmanni 1883. Archives de la Muséum d'Histoire Naturelle de Lyon 3:188. Lortet
was the first scientist to learn where this species actually comes from--the environs of Alexandria, Egypt--and in
describing this tortoise was apparently unaware of Günther's prior description of T. leithii. However, the
latter binomial remained in common usage until Williams showed that the term rightly belonged to a fossil turtle
described in 1852 (see Loveridge & Williams, 1957). Lortet received his first specimens of T. kleinmanni in
1875. In French.
Lortet, L. 1887. Observations sur les tortues terrestres et paludines du bassin de la Méditerranée. Arch. Mus.
Hist. Nat. Lyon 4:1-26. Exquisite colored figures of this and other Mediterranean tortoises accompany an
informative, factual account which first points out the winter activity of this species. A masterpiece. In French.
Loveridge, A. & E. E. Williams 1957. Revision of the African tortoises and turtles of the suborder Cryptodira.
Bull. Mus. Comp. Zool. 115:276-283. This excellent account relies heavily on Flower's works and was
state-of-the-art until Geffen and Mendelssohn began to publish their findings in the 1980s. Two good photos of a
preserved specimen are included as well as excellent drawings of the skull, bony carapace and plastron, and 3 variants
in head scalation. The presentation of Pseudotestudo as a new monospecific subgenus for this tortoise is
preceded by a discussion and key on pp. 257-261.
Marx, H. 1968. Checklist of the reptiles and amphibians of Egypt. Technical Report, p.44, map 35. U.S. Naval
Medical Research Unit No. 3. FPO New York. Marx's team collected 13 specimens, 3 of them from new sites for this
species, one of which (Bir Gindali) apparently is an introduction.
Mendelssohn, H. 1981. Biology and conservation problems for Testudo (Pseudotestudo) kleinmanni
Lortet 1883 in Israel. This unpublished report forms the basis for Groombridge's account of the species (1982).
Mendelssohn, H. & E. Geffen 1987. The Egyptian tortoise in Israel. Israel Land and Nature 12:153-157.
Essentially, this excellent review of the species is an English translation of Geffen and Mendelssohn 1986. Includes
an x-ray of a gravid female and a map which shows only a small part of this species' actual range in Libya.
Mendelssohn, H. & E. Geffen 1987a. Egyptian tortoise needs refuge in Israel. Oryx 21:250. A summary of the
above work, this is intended for a broader audience beyond Israel.
Obst, F. J. & W. Meusel. 1974. Die Landschildkröten Europas. A. Ziemsem Verlag, Wittenberg Lutherstadt, DDR.
This valuable book (out of print) emphasizes captive husbandry and breeding of Mediterranean tortoises, and has photos
of a T. kleinmanni on p.11, p. 20, and a color plate after p.16. The account of this species is very brief
(pp.11-12). The range map for Testudo species is a joke. In German.
Perry, G. 1984. Guide to the turtles of Israel. HARDUN, J. Israel Herpetological Information Center 2:66-67.
Four sentences cover T. kleinmanni, with 3 misspellings in the last one. Anderson's 1898 life-size color print
of this tortoise is reproduced on the back cover, and the lengthier Hebrew account of T. kleinmanni in Israel
appears on p. 17. A good photo contrasting the plastrons of 2 T. kleinmanni with that of a juvenile T.
graeca appears on p. 18.
Philippen, H.-D. & C. Serbent. 1987. Zur Biologie und Ökologie von Testudo (Pseudotestudo) kleinmanni--der
Ägyptischen Landschildkröte. Die Schildkröte 1: 3-20. Leave it to the Germans to publish the first article focused
on captive husbandry, despite the title. While drawing considerably on Flower's and Groombridge's species accounts,
those of Lortet and Anderson are ignored. References to two previous German accounts on captive husbandry are given.
The finicky nature of this species in captivity is discussed but pp. 13-15 provide useful guidance for feeding and
housing. There are 4 photos, 2 of which show advocated enclosures. In German.
Pritchard, P. C. H. 1985. Testudo kleinmanni in Egypt and Israel. IUCN Freshwater Turtle & Tortoise
Newsletter 1:3. Personal communication from yours truly and my (yet unpublished) 1985 work form the basis of these
3 paragraphs. The figure of 157.7 mm as the average carapace length of an adult female T. kleinmanni is a
typographical error, as the record size for this species is only 127 mm (Flower, 1933).
Schleich, H. H. 1984. Reptilien und Amphibien des Kouf National Parkes in NE-Libyen. Landesverband für Amphibien-
und Reptilienschutz Bayern e. V. Mitteilungen 4(1):9-10. Essentially this short paper is an abstract or prospectus
for his much longer 1989 paper, listing T. kleinmanni from semi-desert and T. graeca from wadis and
plateaus. In German.
Schleich, H. H. 1984a. Merkmalsausbildungen an Landschildkröten in Nordost Libyen (Testudines: Testudinidae). On
characteristics of northeast Libyan tortoises. Herpetozoa 1:97-108. This is the first contemporary work which
deals with T. kleinmanni in Libya. Unfortunately the author presents inaccuracies with regard to climatology,
diagnostic plastron markings, and geography. He found 3 specimens (1.2), two of which lack dark markings whatsoever on
the plastron--perhaps the most significant contribution of this account. Among the excellent photos are 2 fine ones of
T. kleinmanni, one showing the immaculate yellow plastron alluded to. If Schleich is correct, the habitat use
of this species in Libya (favoring crevices and bushes as cover sites) differs from that in Israel, where rodent
burrows are chiefly utilized. In German.
Tristram, H. B. 1884. The Survey of Western Palestine. Vol 4, pp. 156-157. Palestine Exploration Fund, London.
To his credit, Tristram was apparently the first naturalist to use Lortet's binomial T. kleinmanni, but his
geographic grasp of its occurrence in what is now southern Israel was inaccurate. Flower (1933) may himself have
misjudged Tristram in his identification of tortoises from the northern Negev, and excludes kleinmanni/leithii
from what is now Israel. Bodenheimer (1935) was on the right track, but was Tristram?
Werner, Y. 1982. Georg Haas, 1905-1981. Obituary, Editorial news and notes, Copeia (2):491-496. Haas first
"rediscovered" T. kleinmanni in Israel in 1963, setting the stage for the field work of Mendelssohn and Geffen.
Werner, Y. 1982a. Herpetofaunal survey of the Sinai Peninsula (1967-77), with emphasis on the Saharan sand
community. U.S.F.W.S., Wildlife Research Report 13:153-161. Very little discussion of T. kleinmanni in
Sinai, where few specimens were found by Israelis.
Zavattari, E. 1930. Erpetologia della Cirenaica. Archivio Zoologico Italiano 14:263. Professor Zavattari
deserves an "A" for honesty, as he confesses that the tortoises which he found in the interior plateau of Cyrenaica
(NE Libya) somehow disappeared before he could identify them as either "T. leithii" or "T. ibera". Like
Anderson (1898), he makes a troublesome reference to the species occurring beyond Egypt in Syria, unsubstantiated to
date by specimens. In Italian.
Originally published in the Tortuga Gazette 29(1): 1-4, January 1993