Moorish tortoise ♀, Northeast Turkey
|Linnaeus , 1758|
The Moorish tortoise ( Testudo graeca ) belongs to the genus of the Palearctic tortoises ( Testudo ). Most of its huge range is in Asia and North Africa , but two of the six subspecies are also found in the European Mediterranean region. As a small to medium-sized, herbivorous tortoise, the Moorish tortoise is a popular pet in German-speaking countries , but its population is threatened in many of its ancestral home areas. The private import of these turtles is therefore (depending on the subspecies) illegal and is punished with a fine or imprisonment.
The Moorish tortoise is a small to medium-sized tortoise. Adult specimens of the Eastern European Testudo graeca ibera can reach a length of up to 35 cm (SCL, height of the back armor) and a weight of more than 5 kg (Beshkov 1997). Animals of western Mediterranean subspecies remain significantly smaller. For southeastern Spain an average length of 11.2 cm for males and 12.6 cm for females is given (Pérez 1998). The back armor ( carapace ) is highly arched, with the highest point in the area of the third vertebral, and has relatively wide vertebral shields. The belly armor ( plastron ) is significantly thickened at the front lobe and in adult animals has a weakly movable hinge at the rear end. As with most turtles, the shell, made of bone plates, is covered by thin horn shields, whereby the so-called tail shield in Testudo graeca is mostly undivided. The front legs of the Moorish tortoise are covered on the outside with four to six rows of overlapping, large horn scales and usually have five claws. In addition to a North African local form, there are two typical horn cones on the hind legs, arranged symmetrically to the right and left of the tail. The coloring and drawing of the shell is very different depending on the individual and the subspecies. Young animals usually hatch from the egg with a clear and high-contrast drawing of the shell, but in some North African local forms they are almost monochrome light brown. In older animals, the coloring and drawing become increasingly blurred. The color of the soft tissues also varies greatly, depending on the climatic conditions, from light yellow-brown to almost black. In some local forms, there is a characteristic bright spot on the top of the head.
Features distinguishing it from other species of the genus
Compared to the Greek tortoise Testudo hermanni , Testudo graeca differs through its almost always present thigh spurs and the lack of a horn nail at the tail end. Further distinguishing features are four to six conspicuous transverse rows of coarse, overlapping scales on the outer surfaces of the forearms and the hinge on the rear lobe of the belly shell, which is always present in adult animals. The Moorish tortoise can be distinguished from the wide-brimmed tortoise Testudo marginata by its more rounded shell shape and the lack of triangular plastron markings. The Egyptian tortoise Testudo kleinmanni is much smaller and lighter in color than Testudo graeca and also has a triangular plastron mark. The four-toed tortoise ( Testudo horsfieldii ) has only four, while the Moorish tortoise has five claws on its front feet.
Moorish tortoises have a pronounced sexual dimorphism . Males and females show clear secondary sexual characteristics that are expressed at around four to six years of age. Animals that have just hatched from the egg cannot be differentiated according to sex.
Adult males have a longer tail and a thicker tail base in which the penis is hidden and is only everted for mating. Males usually have their tails folded sideways under the armor, while the short tails of the females remain stretched out. Your cloaca is also closer to the body. The belly armor of the males is slightly curved inwards ( concave ). The tail shield clearly curved inwards. Females, on the other hand, are generally larger and heavier than males, although the differences in size and weight can sometimes be considerable.
For a long time, the Moorish tortoise was considered to be a species with few subspecies despite its huge distribution area over three continents with different landscapes and climatic zones. For several years, however, new varieties have been discovered and described as further subspecies (e.g. Testudo graeca pallasi ) or independent species (e.g. Testudo perses ) or even genera (e.g. Furculachelys nabeulensis ), but in some cases not generally accepted. Two new studies based on genetic studies indicate the following 10 subspecies for the Moorish tortoise.
- Armenian tortoise , Testudo graeca armeniaca, Chkhikvadze, Bakradze 1991
- This subspecies is characterized by a low armor shape reminiscent of Testudo horsfieldii and a mostly rather dark color with little contrasting markings. The maximum size is given as 20–23 centimeters. According to genetic studies, the Testudo graeca pallasi previously described as a separate subspecies was synonymous with Testudo graeca armeniaca.
- Area: western coastal area of the Caspian Sea, eastern or parts of the Central Caucasus in Armenia and Turkey
- Caspian tortoise , Testudo graeca buxtoni , Boulenger 1921
Testudo perses was incorporated into this subspecies .
- Area: Northwest and Central Iran, Eastern Caucasus
- Eurasian tortoise , Testudo graeca ibera , Pallas 1814
- This is the largest subspecies. Individual animals reach body sizes of over 30 centimeters. The armor shape is slightly flatter than other representatives of the species and round to oval. The basic color of the tank is olive in youth with clearly contrasted black markings. With increasing age, the animals usually get darker, especially on the soft tissues. Testudo ibera racovitzai and Testudo graeca nikolskii , which were previously described separately, were also incorporated into this subspecies .
- Biotope: cultural landscape, open bush landscape and meadows, light forest edges, sandy coastal strips, steppe and semi-desert.
- Area: Southeast Europe, western Asia Minor, Russian and Georgian Black Sea coast, Central Caucasus.
- Origin of name: The subspecies name "ibera" goes back to an ancient name of the Kura valley in Georgia ( Iberia ) and not the Iberian peninsula (Spain).
- Levantine tortoise , Testudo graeca terrestris Forskål 1775
- These animals stand out due to the predominantly light, sometimes striking yellow color ("Golden Greek") of their armor and soft tissues. Dark pigment spots are seldom black, but mostly rather brown and reduced in spread. The animals are extremely high back. There is a characteristic yellow mark on the head. The maximum size is usually 16 centimeters. Testudo ibera var. Bicaudalis , Testudo floweri , Testudo graeca anamurensis and Testudo antakyensis were integrated into this subspecies .
- Biotope: poorly vegetated steppe, scrubland, dry forests
- Area: southern and eastern Asia Minor, Levant
- Persian tortoise , Testudo graeca zarudnyi, Nikolsky 1896
- The elongated shell shape with a sawn and broadened rear edge resembles the wide-brimmed tortoise Testudo marginata . The coloring of the animals is olive to brown, the clear spots of the youth stage later only blurred. It is a large subspecies with armor lengths over eight inches.
- Biotope: Rocky hills and plains up to a height of 2500 meters
- Area: East Iran
- Origin of the name: This subspecies was named in honor of the Russian zoologist Nikolai Alexejewitsch Sarudny .
- Cyrenaica tortoise , Testudo graeca cyrenaica , Pieh and Perälä 2002
- The Cyrenaica tortoise has narrow front extremities with only weak defensive scales. The armor is waisted, has a narrow front opening and rear marginalia that are skirted upwards. The supracaudal scale is comparatively strongly curved inwards in male animals.
- Area: Libya
- Holotype: Museum für Tierkunde, Dresden, MTKD 31880
- Terra typica: Derna 32 ° 46'N, 22 ° 39'E (Darnah, Cyrenaica, Eastern Libya)
- Origin of name: the name refers to the Cyrenaica region in northeastern Libya
- Moroccan tortoise , Testudo graeca morokkensis , Pieh and Perälä 2002
Testudo graeca marokkensis has a comparatively flatter back armor than the nominate form . A multitude of fine lines, lines and speckles contrast with the ocher-colored to yellow basic color of the armor; the shields also have a dark, partially torn border and central areole spots. The belly armor is lighter than the back, but also has black drawing elements. The head is dark with several light scales, the extremities are predominantly dark with individual lighter scales. Genetic studies have shown that Testudo graeca lamberti is very closely related to Testudo graeca marokkensis and should not be understood as an independent subspecies.
- Biotope: arid landscape with steppe climate
- Area: Northern Morocco, on both sides of the Rif Mountains
- Holotype: State Museum for Natural History, Stuttgart, SMNS 7602
- Terra typica: Tarmilete, 33 ° 23 'N, 06 ° 04' W, north-west Morocco
- Origin of the name: The name goes back to the distribution area in Morocco.
- Moorish tortoise , Testudo graeca graeca , Linnaeus 1758
- The carapace drawing by Testudo graeca graeca consists of solid drawing elements on a yellow background. The vortex shields have a dark edge and also dark areole spots . The local form described as Testudo graeca whitei belongs genetically to this subspecies.
- Area: Algeria, Eastern Morocco, Spain (probably introduced here)
- Terra typica: Oran (approx. 35 ° 41'N 0 ° 38'W), Algeria
- Sousstal tortoise , Testudo graeca soussensis , Pieh 2001
- The basic color of the back armor is ocher, with little contrast, thin-line line drawing and almost completely missing drawing on the plastron. The soft tissues are brownish to sand-colored, sometimes also yellowish-orange, with lighter, but less contrasting scales on the head. Young animals have little or no drawing. It is the largest Moroccan tortoise species with a back shell length of up to 25 cm (SCL). The thigh spurs otherwise present in Graeca animals are often missing in Testudo graeca soussensis , especially in animals from the actual Sousstal .
- Biotope: Opportunistic settlement of cultivated areas, particularly occurring in thorn hedges, which offer protection and food
- Area: South Morocco
- Holotype: Museum für Tierkunde, Dresden, MTKD 33842
- Terra typica: Area of Agadir, 30 ° 28'N, 9 ° 55'W, Sousstal, southwest Morocco
- Name origin: the name goes back to the southern Moroccan Sousstal.
- Tunisian tortoise , Testudo graeca nabeulensis , Highfield 1990
- On a yellow basic color, this subspecies has vividly contrasting, black drawing elements. The skin of the soft tissues and the scales on the extremities are sand-colored to yellow, sometimes with black tips. The head shows a characteristic, bright yellow markings. The Tunisian tortoise is the smallest Graeca subspecies. Females typically reach a back armor length of a little over 13 cm and a weight of approx. 600 g. Males remain significantly lighter with a shell length of just under 12 cm and an average of 375 g.
- Area: Tunisia, East Algeria, Sardinia and Sicily (probably introduced)
- Terra typica: Nabeul, Tunisia
- Origin of name: the name refers to the terra typica
Way of life in nature
Moorish tortoises are exclusively diurnal reptiles that depend on the supply of solar energy to digest their food. As poikilothermal (cold-blooded) animals, they can not generate the necessary body heat themselves, but have to influence it according to their needs by moving from shady to sunny places. The body temperatures of active animals are given in the literature as 22 to 37 ° C.
For the proper digestion of their fiber-rich food, they even need body temperatures of over 30 ° C for a few hours a day, which they can reach, for example, by sunbathing in the morning even when the air temperature is significantly lower. In the cooler season, the animals usually only appear in the late morning, sunbathe and then disappear again. Above 40 ° C they endanger their lives and bury themselves in the cooler earth or look for crevices and structures of other animals. In the summer months they therefore show a two-phase behavior, with activity only in the cooler morning and afternoon. In hot, arid habitats, a longer summer sleep ( estivation ) is even necessary in summer , in southern Morocco from around June to September. The metabolism comes to a standstill below 8 ° C. Breathing and heart rate are also greatly reduced. In some parts of its range, the Moorish tortoise therefore often falls into months of rigid winter ( hibernation ) and is partly inactive and buried from September to March. In addition, little is known about the way of life of the various subspecies of the Moorish tortoise. Only a small Spanish population is being researched more intensively. The following information therefore often only refers to the nominate form of the Moorish tortoise as it occurs in the Coto de Doñana National Park in southern Spain .
Distribution area and habitat
The Moorish tortoise colonizes a huge, but very fragmented range, with large gaps between the individual occurrences. It extends over 27 national areas, from the Atlantic coast in western Morocco to the eastern parts of Iran. It extends over 6000 kilometers in an east-west direction and 1600 kilometers in a north-south direction. The species lives under very different climatic conditions, the comparatively balanced Mediterranean climate and the extreme weather conditions of the Asian steppes with scorching summers and months of frost in winter. The populated habitats also differ greatly, from humid swamp areas to heather and grasslands, light forests, sandy dune landscapes to sparse, dry steppes and semi-deserts. The IUCN classifies the Moorish tortoise as "Endangered".
This type of turtle feeds mainly on plants. A large number of annual and perennial plants from a wide range of plant families are recorded. In the Doñana an uptake of 86 plant species from 26 families was observed, mainly representatives of the sweet grasses (Gramineae), daisy family (Asteraceae), legumes (Leguminosae), buttercups (Ranunculaceae) and carnations (Caryophyllaceae). If there is sufficient food available, usually in spring, there are clear food preferences for certain plant species. In Dagestan, a share of 97 percent of the comparatively protein-rich legumes was found in the stomach contents of 62 animals. These are, for example, wild vetch and clover. In addition, the more nutritious parts are preferred, buds, flowers, fruits with seeds. If there is a lack of food, especially during the hot summer months, dried out, poorly nutritious plants and parts of plants, sometimes even poisonous plants, are ingested and used. In particular, remains of invertebrates (e.g. snails and insects) and the faeces of other animals were found in the stomach contents and faeces . Females ingest more animal components than males and young animals.
Mating and laying eggs
In the summer preceding mating, the males begin to produce sperm and the females begin to grow follicles. Immediately after the winter rigor, from mid-February to early May, the males begin courtship. With great zeal they pursue the females, circle them, bite them in the limbs, ram them with violent thrusts and ride on them. In intermittent copulation, the male opens its mouth, the red fleshy tongue becomes visible and it emits beeping sounds. After riding on, the female stops, pushes her forefeet into the earth and swings her front body. The rhythm of the tank movement corresponds to the rhythm in which the male utters his mating sound. A single successful mating is enough for several clutches.
Depending on the region, usually from April to June, one to three clutches with up to eight (on average about five) hard-shell eggs are deposited. To do this, the female digs a pit about ten centimeters deep in a carefully selected place, carefully places the eggs in it with her hind legs and bury them. No further brood care takes place. The sun does the hatching. Depending on the region and climatic conditions, the development of the embryos takes from 60 to over 100 days, with the young often remaining in the nest cavity for a while after hatching. The first hatchlings appear on the surface of the earth during the summer dry season, in the Caucasus from the end of July, in southwest Spain from mid-August. However, it is also assumed that the last hatchlings overwinter in the nest (Inozemtsev 1994).
The mean nesting temperature under natural conditions in Spain is given as around 28 ° C, with extreme values of 10 and 48 ° C. The daily fluctuation averages around 13 ° C (Díaz-Paniagua 2006). In Moorish tortoises, the sex of the developing young is determined by the incubation temperature. Animals hatched under laboratory conditions were mostly males at constant temperatures below 30 ° C, and mostly female at temperatures above 31 ° C.
Hazard and protection
By the IUCN, the thighed tortoise is on the Red List of Threatened Species as endangered ( vulnerable out).
The species is listed by the European Union in Annexes II and IV of the Habitats Directive and is therefore considered to be strictly protected species of community interest, for whose preservation special protected areas must be designated by the member states.
- The oldest known Moorish tortoise, Timothy , a female of the subspecies T. g. ibera was an estimated 160 years old. She first lived for 40 years as a naval mascot on several British ships, took part in the siege of Sevastopol in the Crimean War on board the HMS Queen and later lived at Powderham Castle near Exeter .
- The Natural History Museum of Geneva houses a two-headed Moorish tortoise. The animal hatched on September 3, 1997 and was named after the Roman god Janus .
- The Latin name of the species, Testudo graeca , means "Greek tortoise", but it is to be distinguished from the Greek tortoise ( Testudo hermanni , also known as Testudo graeca in old literature ). Linnaeus is said to have chosen the name because the appearance of the armor shields reminded him of a Greek mosaic .
- U. Fritz, AK Hundsdörfer, P. Široký, M. Auer, H. Kami, J. Lehmann, LF Mazanaeva, O. Türkozan, M. Wink: Phenotypic plasticity leads to incongruence between morphology-based taxonomy and genetic differentiation in western Palaearctic tortoises (Testudo graeca complex; Testudines, Testudinidae). In: Amphibia-Reptilia. 28 (1), 2007, pp. 7–121 ( summary ( memento of the original dated October 2, 2007 in the Internet Archive ) Info: The archive link has been inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. )
- James F. Parham, Oğuz Türkozan, Bryan L. Stuart, Marine Arakelyan, Soheila Shafel, J. Robert Macey, Yehudah L. Werner, Theodore J. Papenfuss: Genetic Evidence for Premature Taxonomic Inflation in Middle Eastern Tortoises. In: Proceedings of the California Academy of Sciences. Fourth Series, Vol. 57, No. 3, 2006, , pp. 955-964
- Uwe Fritz, D. James Harris, Soumia Fahd, Rachid Rouag, Eva Graciá Martínez, Andrés Giménez Casalduero, Pavel Široký, Mohsen Kalboussi, Tarek B. Jdeidi, Anna K. Hundsdörfer (2009): Mitochondrial phylogeography of Testudo graeca in the Western Mediterranean: Old complex divergence in North Africa and recent arrival in Europe.- Amphibia-Reptilia 30 (2009): 63-80 Summary ( page no longer available , search in web archives ) Info: The link was automatically marked as broken. Please check the link according to the instructions and then remove this notice.
- Highfield, http://www.tortoisetrust.org/articles/furculachelys.html , accessed February 18, 2009
- Julia Voss : An uprising without heraldic animal , FAZ , July 6, 2013, p. 31. There are also the comments made by Edhem Eldem in an interview.
- Rory Knight Bruce: Timothy the Tortoise. Orion, London 2004, ISBN 0-7528-6872-1
- JANUS fête son dixième anniversaire ( Memento of the original of March 7, 2009 in the Internet Archive ) Info: The archive link has been inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. , Geneva City Archives (French)
- Peter CH Pritchard : Encyclopedia of Turtles . TFH Publications Inc. Ltd., Neptune 1979, ISBN 0-87666-918-6 , p. 403 u. 398
- Markus Baur: The keeping and care of European tortoises. Transcript of a lecture. Zoo, Karlsruhe 1999.
- James R. Buskirk, Claudia Keller, Ana C. Andreu: Testudo graeca LINNAEUS, 1758 - Moorish tortoise. Handbook of the reptiles and amphibians of Europe. Vol. 3 / IIIA. Tortoises (Testudines). TI (Bataguridae, Testudinidae, Emydidae). Aula-Verlag, Wiebelsheim 2001, ISBN 3-89104-004-0 , pp. 125-178
- C. Díaz-Paniagua, AC Andreu: Tortuga mora - Testudo graeca. In: LM Carrascal, A. Salvador, (Ed.): Enciclopedia Virtual de los Vertebrados Españoles. Museo Nacional de Ciencias Naturales, Madrid 2005
- Walter Kirsche: The tortoises of Europe. 2nd Edition. Mergus, Melle 1998, ISBN 3-88244-009-0
- Carolus Linnaeus: Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Stockholm 1758 (original description by Testudo Graeca)
- Alexander Pieh, Hans-Dieter Phillipen: Mediterranean tortoises In: Draco. 8 (4), Münster 2007, pp. 7-22
- Wolfgang Wegehaupt: Sardinia, the island of the European turtles. Wegehaupt, Kressbronn 2004, ISBN 3-9809774-1-2