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Testudo

Testudo is a genus of tortoises in the family Testudinidae, native to arid and semi-arid habitats across southern Europe, North Africa, and western Asia. These small to medium-sized reptiles, typically measuring 15–35 cm in carapace length, feature high-domed shells, sturdy limbs adapted for terrestrial life, and a primarily herbivorous diet of grasses, herbs, and occasional fruits or invertebrates.^[1]^[2] The genus includes prominent species such as the spur-thighed tortoise (Testudo graeca), Hermann's tortoise (Testudo hermanni), marginated tortoise (Testudo marginata), and Russian tortoise (Testudo horsfieldii), with numerous subspecies reflecting regional adaptations and ongoing taxonomic revisions that sometimes elevate former subgenera like Agrionemys or Chersine based on phylogenetic analyses.^[3]^[4] Testudo species are noted for their longevity, often exceeding 50–100 years in captivity, and behaviors including seasonal estivation or hibernation to endure dry periods.^[5] Despite their popularity in the international pet trade—which has driven captive breeding programs—wild populations of most Testudo taxa face severe declines due to habitat fragmentation from agriculture and urbanization, overexploitation for pets and traditional medicine, and vulnerability to climate shifts, resulting in IUCN classifications ranging from vulnerable to critically endangered for key species and subspecies.^[1]^[5] Conservation efforts emphasize protected areas, trade regulations under CITES Appendix II, and reintroduction initiatives, though challenges persist from taxonomic uncertainties complicating targeted protections.^[4]^[3]

Taxonomy

Etymology

The genus name Testudo derives from the Latin noun testūdō (genitive testūdinis), signifying "tortoise" or "turtle," a term rooted in testa, meaning "shell" or "potshard," evoking the reptile's domed, protective carapace.^[6] This etymon underscores the ancient perception of the shell as an arched, impregnable enclosure, akin to a natural fortress.^[7] In Roman usage, testudo extended metaphorically to military tactics, denoting the "tortoise" formation in which legionaries raised and overlapped shields to form a continuous overhead barrier against projectiles, drawing an explicit analogy to the tortoise's defensive shell without implying any direct influence on the zoological designation.^[8] Swedish naturalist Carl Linnaeus established Testudo as a formal genus in binomial nomenclature through the tenth edition of Systema Naturae, published on October 1, 1758, wherein he classified various Old World tortoise species under it, including Testudo graeca (the spur-thighed tortoise).^[2]

Historical classification

In 1758, Carl Linnaeus established the genus Testudo within his Systema Naturae, initially including a broad array of known tortoise species based on limited morphological descriptions available at the time, such as those of Mediterranean and Eurasian forms distinguished primarily by shell shape and size.^[9] Subsequent 18th- and 19th-century classifications refined this by emphasizing geographic distribution and external morphology, leading to the exclusion of non-Mediterranean species and the recognition of distinct forms like T. graeca and T. hermanni as core members, while species from farther east, such as T. horsfieldii, were retained under Testudo despite emerging doubts about their congruence.^[5] By the mid-20th century, taxonomic debates intensified over the inclusion of T. horsfieldii (Russian tortoise), with proponents arguing for its separation into the genus Agrionemys due to key morphological differences, including the absence of a functional plastral hinge—a trait present in Mediterranean Testudo species that allows partial shell closure for defense.^[5] This reclassification was formalized in 1966 by Khozatsky and Mlynarski, who prioritized osteological evidence like plastral structure and neural bone patterns to delineate Agrionemys as a distinct lineage, reflecting a shift toward more rigorous anatomical criteria over Linnaean typology.^[5] Genetic analyses from the early 2000s, leveraging mitochondrial DNA sequences such as the 12S rRNA gene, provided empirical support for a monophyletic Mediterranean clade comprising species like T. graeca and T. hermanni, characterized by shared haplotypes indicative of common ancestry in southern Europe and North Africa.^[10] These studies, however, revealed substantial intraspecific variation and deep divergences within nominal subspecies of T. hermanni and T. graeca—often exceeding interspecies thresholds in other taxa—prompting revisions that questioned the validity of traditional subspecific boundaries established on morphology alone and highlighted hybridization risks in fragmented populations.^[11]

Recognized species and subspecies

The genus Testudo encompasses four recognized extant species: T. hermanni, T. graeca, T. marginata, and T. kleinmanni.^[12] These species are distinguished primarily by carapace morphology, scute patterns, and genetic markers, with phylogenetic studies confirming their monophyly within Testudinidae.^[13] Testudo horsfieldii (Russian tortoise), historically included, has been excluded and reclassified into the distinct genus Agrionemys based on morphological traits such as plastron shape and carapace form, corroborated by mitochondrial DNA analyses from the 2010s onward.^[14] This separation persists despite occasional mislabeling in pet trade contexts, where hybrids with Testudo species complicate identification.^[15] Testudo hermanni (Hermann's tortoise) features two subspecies differentiated by external morphology and genetics: the western T. h. hermanni, with vivid yellow-green coloration, pronounced black markings on scutes, and smaller adult size; and the eastern T. h. boettgeri, exhibiting paler tones, subdued black patches, and larger body proportions.^[16] These distinctions arise from geographic isolation and are supported by morphometric data, though captive breeding has introduced hybrid forms challenging pure-line preservation.^[17] Testudo graeca (spur-thighed tortoise) displays extensive intraspecific variation, marked by enlarged tibial spurs and debated subspecies such as T. g. ibera, which genetic studies, including mitochondrial 12S rRNA sequencing, identify as distinct from the nominate form due to deep haplotypic divergence dating to the Pleistocene.^[18] Taxonomic controversies persist over the elevation of regional variants like T. g. ibera to full species status, fueled by morphological gradients across North Africa and Eurasia, with calls for nuclear DNA integration to resolve clinal versus discrete boundaries.^[12] Testudo marginata (marginated tortoise) is defined by its characteristic flared posterior carapace margins and upturned peripheral scutes, with recognized subspecies including T. m. marginata (continental Greece) and T. m. sarda (Sardinia and Corsica), though proposals to elevate T. m. weissingeri highlight ongoing nomenclatural flux based on limited samples.^[19] Testudo kleinmanni (Egyptian tortoise), the genus's smallest member, lacks described subspecies and is typified by a highly domed, keeled carapace with bold yellow markings on dark scutes.^[20] Captive hybridization among Testudo species and subspecies, prevalent since the mid-20th century pet trade expansion, has eroded wild-type genetic integrity, prompting recommendations for molecular authentication in ex situ populations to inform reintroduction viability.^[21]

Morphology and physiology

External features

Species of the genus Testudo are characterized by a high-domed, rigid carapace formed from fused dermal bones overlain by keratinous scutes, providing robust protection against predators. The dorsal aspect typically comprises 13 scutes—five unpaired vertebral scutes aligned centrally and eight paired costal scutes—framed by 22–24 marginal scutes along the periphery.^[22] Adult straight carapace lengths range from 15 to 35 cm across the genus, with variation by species; for instance, T. marginata often exceeds 30 cm and features flared posterior marginal scutes that enhance enclosure and deter attacks.^[23]^[24] The plastron, the ventral shell component, is generally flat and composed of six paired scutes fused to the carapace via a bridge, though four of the five recognized Testudo species exhibit a weak kinetic hinge between the abdominal and femoral scutes, enabling partial closure of the posterior shell for added security.^[25] Limbs are columnar and scaled with large, overlapping epidermal plates adapted for terrestrial locomotion; T. graeca, for example, possesses prominent, in-turned spurs on the thighs, a trait less pronounced or absent in other congeners like T. hermanni.^[26] Marked sexual dimorphism manifests externally, with females attaining larger overall sizes than males—evident in metrics such as carapace length and mass.^[27] Males typically exhibit longer, thicker tails housing the reproductive organs, concave plastrons to facilitate mounting during copulation, and elongated foreclaws suited for digging and combat, alongside shape differences in both carapace and plastron contours.^[28]^[29]

Internal adaptations

Testudo species demonstrate internal physiological adaptations optimized for water conservation, primarily through excretory and osmoregulatory mechanisms that reduce obligatory water loss in arid conditions. As uricotelic organisms, they convert nitrogenous wastes into uric acid, a semi-solid paste that precipitates in the cloaca with minimal associated water, contrasting with the aqueous urine of ureotelic or ammonotelic taxa and thereby conserving body fluids during dehydration. This process is supported by renal function producing low-volume, concentrated uric acid suspensions, with overall urinary output suppressed to as little as 0.1-0.5% of body weight daily under normal conditions, further minimized during water scarcity.^[30] The urinary bladder functions as a dynamic reservoir, capable of expanding to store up to 5-10% of body volume in urine during hydrated periods, from which water and sodium are selectively reabsorbed via mucosal epithelia responsive to hormonal signals like arginine vasotocin. In Testudo graeca, permeability studies confirm the bladder's role in post-renal water recovery, preventing desiccation by recycling fluids into the bloodstream during droughts, while retaining uric acid and potassium for delayed elimination. Cloacal epithelia complement this by facilitating additional water reabsorption from stored wastes or environmental moisture when tortoises immerse in temporary water sources, though uptake rates are modulated to avoid osmotic imbalances. These mechanisms enable prolonged estivation without external water intake, with bladder contents serving as an internal buffer against hypernatremia.^[31]^[32] Metabolic adaptations further enhance drought tolerance by inducing hypometabolism during estivation, a summer torpor state triggered by high temperatures and aridity. In Testudo hermanni, standard metabolic rate, measured via oxygen consumption, exhibits strong thermal dependence but depresses significantly in dormancy, dropping to 10-20% of active levels at ambient temperatures of 20-28°C, allowing reliance on endogenous fat reserves without feeding. Empirical data from respirometry indicate Q10 values of 2.5-3.5 across 8-28°C, with seasonal reductions in ventilatory rate and cardiac output conserving energy and limiting evaporative respiratory losses. This metabolic slowdown, coupled with elevated urea tolerance to avoid toxic accumulation from protein catabolism, permits survival without food for 3-6 months, as evidenced by field observations and fasting trials where plasma osmolality stabilizes despite prolonged inanition. Such responses underscore causal linkages between reduced cellular respiration and extended viability in resource-poor environments.^[33]^[34]

Distribution and ecology

Geographic range

The genus Testudo exhibits a natural distribution primarily confined to the Mediterranean Basin, encompassing southern Europe, North Africa, and adjacent regions of Southwest Asia. Fossil evidence from Pleistocene deposits indicates historically broader ranges across parts of Europe, including the Iberian Peninsula and Greece, reflecting post-glacial expansions followed by fragmentation into isolated populations.^[35]^[36] Testudo hermanni occupies southern Europe, ranging from the Iberian Peninsula (including Spain and the Balearic Islands) through France, Italy (with Sicily and Sardinia), the Balkans (Slovenia to Greece, including Aegean islands), and marginally into western Turkey.^[37] T. marginata is more restricted, primarily in central and southern Greece, extreme southern Albania, and Italy's Sardinia, with occurrences on Balkan islands.^[38] T. graeca spans a wider arc from North Africa (Morocco to Libya and Egypt) across southern Europe (coastal Spain, Italy, Balkans) to Southwest Asia (Turkey, Syria, Israel, and Caucasus regions).^[39]^[40] Testudo kleinmanni has the most limited current extent, with remnant populations confined to disjunct sites in northern Libya and possibly southern Israel, following a historical distribution along the southeastern Mediterranean coast including northern Egypt.^[41] The species' range has contracted by approximately 90% over the past three generations (roughly since the mid-20th century), based on field surveys and IUCN assessments documenting near-extirpation in former core areas like Egypt.

Habitat requirements

Species of the genus Testudo primarily occupy dry, open habitats such as scrublands, grasslands, and cleared oak woodlands, where annual rainfall typically ranges from 200 to 500 mm, supporting semi-arid conditions conducive to their ectothermic physiology.^[42] These environments feature mosaic landscapes with meadows, roadsides, and abandoned fields, providing access to low-lying vegetation for foraging while allowing unobstructed solar exposure.^[43] Loose, sandy soils predominate, facilitating burrowing for shelter, thermoregulation, and overwintering, with preferences for slight slopes oriented south or southeast to maximize insolation during nesting and activity periods from April to June.^[44]^[43] Microhabitat selection emphasizes sunny basking sites interspersed with vegetative cover, enabling individuals to regulate body temperatures near 30°C through behavioral adjustments like morning and midday exposure to sunlight, particularly vital in cooler months such as October.^[45] Shrub-dominated scrubs and dune grasslands serve as key refugia, with seasonal shifts toward sclerophyllous vegetation for enhanced thermoregulation and feeding on tender shoots.^[46] Field observations indicate higher population densities in simplified, semi-open structures like re-colonizing shrublands and agricultural margins, reflecting adaptability to disturbed edges over uniform dense cover.^[47] While capable of persisting in marginal semi-arid agro-ecosystems, Testudo species exhibit vulnerability in dense forests lacking sufficient light penetration or in wetlands, where excessive humidity and shaded understories hinder basking efficacy and access to preferred herbaceous resources.^[43] Such preferences underscore their reliance on xeric Mediterranean biomes, with avoidance of closed-canopy habitats limiting occupancy to areas supporting diurnal activity patterns.^[46]

Behavior and life history

Daily and seasonal activity

Testudo species display predominantly diurnal activity, with individuals emerging from burrows or shelters in the early morning to bask and become active, peaking in mid-morning to early afternoon during spring and cooler months, before retreating in the late afternoon or evening as temperatures rise.^[48]^[49] Activity patterns can shift to bimodal in warmer periods, with a midday lull for shade-seeking to regulate body temperature and avoid desiccation.^[49] Seasonally, northern populations such as T. hermanni enter brumation (winter torpor) from mid-October to late March, burying in soil or vegetation to withstand cold and low resource availability.^[50] In southern, arid ranges, species including T. graeca and T. kleinmanni exhibit aestivation during hot, dry summers—typically June to September—to minimize water loss and heat stress, emerging with autumn rains.^[51]^[27] These tortoises maintain solitary lifestyles year-round except for transient mating encounters, showing philopatry with radio-tracking revealing restricted dispersal; home ranges average 4.6–7.4 hectares for T. hermanni, with males often covering slightly greater distances annually than females, though total movements rarely exceed 1 km per year in undisturbed habitats.^[52]^[53]

Diet and foraging

Testudo species exhibit strict herbivory, deriving nutrition almost exclusively from plant matter including grasses, broadleaf weeds (forbs), succulents, and opportunistic consumption of flowers, fruits, and seeds.^[54]^[55] Their diet emphasizes high-fiber, low-protein vegetation such as dandelion (Taraxacum officinale), plantains (Plantago spp.), and legumes, which provide essential minerals while minimizing risks of renal strain from excessive protein or sugars.^[56]^[57] Foraging strategies prioritize selective grazing on calcium-rich plants, including species like Plantago with calcium-to-phosphorus ratios exceeding 20:1, to support shell and skeletal integrity in environments where mineral availability varies.^[58] Seasonal dietary shifts occur in response to resource phenology; for instance, Testudo hermanni consumes more legume leaves and grasses during spring flushes of tender growth, transitioning to flowers and unripe fruits like those of Ruscus aculeatus in summer.^[57] In Testudo marginata, preferences lean toward diverse herbaceous plants, including legumes for moderate protein intake and aromatic Mediterranean flora such as thymes, alongside grasses in arid habitats.^[55]^[57] Digestive adaptations enhance foraging efficiency on fibrous diets, with low metabolic rates permitting sustenance from sparse, low-digestibility vegetation through prolonged digesta retention.^[59] Hindgut microbial fermentation breaks down cellulose in species like Testudo graeca, yielding volatile fatty acids for energy and enabling utilization of otherwise indigestible plant fibers, as evidenced by studies on gut pH and retention times.^[60]^[56] This process supports survival in seasonal Mediterranean ecosystems where fresh forage is intermittent.^[61]

Reproduction and development

Mating in Testudo species typically occurs in spring following emergence from brumation, with males initiating courtship by pursuing females, circling them, ramming their carapaces, biting their limbs, and producing vocalizations such as grunts or moans to stimulate mounting.^[62] Females oviposit 1–3 clutches annually between May and July, depending on species and environmental conditions; clutch sizes range from 1–7 eggs, averaging 3–5 for T. hermanni and T. graeca, with smaller clutches of 1–3 for T. kleinmanni.^[63]^[64] Eggs are laid in flask-shaped nests excavated in loose soil to depths of 5–15 cm, covered and left to incubate for 60–120 days under natural temperature fluctuations.^[27]^[65] Incubation temperature determines hatchling sex via temperature-dependent sex determination, with pivotal temperatures around 30°C yielding balanced sex ratios; lower temperatures produce more males, while higher ones (>32°C) favor females, though wild nest microclimates often result in variable outcomes.^[66] Hatchlings emerge in late summer or autumn, fully independent but vulnerable, with annual juvenile survival rates as low as 52% for T. hermanni due primarily to predation and desiccation.^[67] Growth is indeterminate but slows after maturity, with individuals reaching sexual maturity at 6–15 years, males typically earlier than females based on carapace length thresholds of 100–140 mm; mark-recapture data indicate adulthood in 10–15 years under optimal conditions.^[68]^[69] Adult sex ratios in wild populations are frequently male-biased (e.g., >1.5:1 in dense Greek T. graeca sites), attributable to earlier male maturation, higher female energetic costs from reproduction, and differential survival favoring males in some habitats.^[70]^[71] Fecundity remains low, with lifetime reproductive output limited by infrequent breeding and high early-life mortality, contributing to slow population recovery rates observed in long-term studies.^[72]

Conservation and threats

Population statuses

Testudo kleinmanni is assessed as Critically Endangered by the IUCN, with a global population estimated at around 3,000 individuals, including fewer than 2,000 in Israel, and experiencing continued declines.^[73]^[74] Testudo hermanni is classified as Vulnerable, with fragmented populations across its range; for instance, localized estimates in Corsica indicate densities supporting hundreds of individuals per site, though island-wide totals remain under 10,000 in key areas amid decreasing trends.^[75] Testudo graeca holds a Vulnerable status overall, with subspecies clades showing variable stability—relatively common in North African core ranges but declining in peripheral European and eastern populations due to fragmentation.^[76]^[77] In contrast, Testudo marginata is listed as Least Concern, with dense populations reported in parts of its Greek and Italian range, such as encounters of over 30 individuals in short surveys. Across the genus, long-term monitoring reveals patchy data but confirms overall declining trajectories, with some insular T. hermanni populations in the Balearic Islands exhibiting stability as of 2020s assessments.^[78]^[79]

Primary threats

Habitat loss and fragmentation constitute the dominant threats to Testudo species, driven primarily by agricultural expansion, urbanization, and overgrazing by livestock across Mediterranean and North African ranges. These pressures have resulted in rapid degradation of open woodlands, scrublands, and grasslands essential for foraging and shelter, with long-term monitoring revealing heavy population declines and habitat shifts in species like T. hermanni over periods spanning 28 years in central Italy.^[80] For T. kleinmanni, urban development, cultivation, overgrazing, and associated desertification have imposed enormous strain on desert-edge habitats in Egypt and Libya, exacerbating fragmentation and reducing available refugia.^[81] Illegal collection for the pet trade inflicts severe direct mortality, particularly on T. graeca and T. kleinmanni, with intensive commercial harvesting persisting despite CITES Appendix I and II listings. Smuggling operations, often involving wild-caught juveniles from source countries like Morocco and Iran, continue to deplete local populations, as evidenced by ongoing online trafficking and high-value sales that incentivize poaching.^[82]^[83] Compounding these are incidental threats such as road vehicle collisions, which elevate adult and juvenile mortality in fragmented landscapes, and predation by introduced mammals like foxes in altered ecosystems.^[84] Although climatic variability, including intensified droughts, influences survivorship and reproduction in arid habitats, data from range-wide assessments underscore anthropogenic land-use changes as the overriding causal factor in observed declines, outpacing natural stressors in impact.^[73]^[85]

Conservation strategies

Species of the genus Testudo have been listed under CITES Appendix II since 1975, regulating international trade to prevent overexploitation while allowing exports of captive-bred specimens.^[86] This has contributed to reductions in wild-caught exports, particularly through ranching and captive propagation programs that supply the pet trade legally, thereby alleviating pressure on natural populations; for instance, European programs for T. hermanni have propagated thousands of individuals annually since the 1990s.^[87] ^[88] Habitat restoration efforts emphasize protected areas, such as EU Natura 2000 sites covering significant portions of T. hermanni range (e.g., 29% of area of occupancy in some regions), which incorporate measures like vegetation management and fire prevention to maintain suitable scrubland and open woodland.^[84] These initiatives have stabilized local densities in core sites by limiting fragmentation, though population-level increases remain limited without broader landscape connectivity.^[89] Head-start programs, rearing juveniles in captivity before release, have been implemented by organizations like SOPTOM in France and CARAPAX in Italy, yielding releases of 200–300 T. hermanni annually from the Albera project since the early 2000s, with short-term survival improved via size-at-release thresholds exceeding natural hatchling vulnerabilities. ^[90] Reintroduction efforts, often paired with head-starting, face variable outcomes; while some translocations succeed through habitat matching (e.g., microhabitat adaptation in French sites), others report high post-release mortality—up to 40% uncertainty in long-term viability across reptile projects—due to suboptimal site selection, such as inadequate thermal refugia or predation exposure.^[91] ^[92] Criticisms highlight overemphasis on public reserves, which cover fragmented habitats and overlook incentives for private landowners to sustain tortoises on agricultural edges where much Testudo range persists, potentially undermining scalability.^[93] Overall, while CITES and captive programs have curbed trade declines, integrated strategies addressing private land stewardship and refined reintroduction protocols are needed for sustained recovery.^[94]

Human interactions

Role in pet trade

Species of the genus Testudo, particularly T. hermanni and T. graeca, are popular in the pet trade due to their compact size (typically 15-25 cm in adulthood) and relative hardiness in temperate climates, making them suitable for outdoor enclosures in regions like Europe and the United States.^[95] Legal captive-bred specimens, such as those from registered breeders in the EU, help alleviate pressure on wild populations by providing alternatives to poaching; for instance, hundreds of T. h. boettgeri are imported annually into the US from European captive sources, often documented under CITES Appendix II permits.^[88] ^[96] Captive breeding programs have expanded since the 1990s, with EU regulations emphasizing source verification to promote sustainability; however, challenges include risks of hybridization between subspecies (e.g., western T. h. hermanni and eastern T. h. boettgeri), which can dilute genetic purity and complicate conservation reintroductions.^[97] High juvenile mortality rates, often exceeding 20-30% in the first year, stem from inadequate husbandry such as insufficient UVB lighting, improper temperature gradients (requiring 30-35°C basking spots), or suboptimal diets leading to metabolic bone disease.^[98] Despite these issues, well-managed captives achieve longevity of 50-100 years, far surpassing wild averages affected by predation and habitat loss.^[99] Bans on wild imports, intensified post-1990s via CITES and EU wildlife trade regulations (e.g., prohibiting commercial Annex A species like T. hermanni), aim to curb overexploitation but have sparked debate over stifling verified legal trade while failing to eliminate black markets, where mislabeled wild-caught animals persist online and via smuggling.^[100] ^[95] Evidence from trade analyses indicates that robust captive breeding sustains pet demand more effectively than wild-only protections, reducing poaching incentives in source countries like Turkey and North Africa, though illegal trade volumes remain substantial (e.g., thousands of undeclared T. graeca annually).^[101] ^[102]

Cultural and historical significance

In ancient Greek literature, species of the Testudo genus, such as Testudo graeca, symbolized perseverance through Aesop's fable "The Tortoise and the Hare," where the tortoise's steady determination defeats the hare's overconfidence, teaching that consistent effort triumphs over impulsiveness.^[103] This narrative, attributed to Aesop around the 6th century BCE, drew from observations of Mediterranean tortoises' deliberate movement and endurance.^[103] The Romans associated Testudo with defensive resilience, naming their interlocking shield formation testudo after the tortoise's protective shell, a tactic used in sieges from the 1st century BCE onward to shield against arrows and projectiles.^[104] Pliny the Elder, in his Natural History (circa 77 CE), described a tortoise's role in the mythical death of playwright Aeschylus (d. 455 BCE), where an eagle dropped it on his bald head, mistaking it for a stone, highlighting the animal's cultural presence in Greco-Roman anecdotes about fate and nature.^[105] Elites in both cultures kept tortoises as pets or omens, valuing their longevity—some reportedly living over 100 years—as emblems of stability, though records emphasize symbolic rather than practical utility.^[106] In modern contexts, the Testudo name evokes the genus in the University of Maryland's mascot, adopted in 1933 as a bronze statue of a diamondback terrapin (Malaclemys terrapin), symbolizing tenacity for the institution's athletic teams; the choice, proposed by coach H. Curley Byrd in 1932, linked to regional reptile imagery despite the species difference.^[107] Unlike more prominent reptiles in heraldry or folklore, Testudo species hold limited folkloric roles in Mediterranean traditions, occasionally representing wisdom or earth-bound endurance, but without widespread heraldic adoption.^[108] Early scientific interest focused on anatomy, contributing to herpetological taxonomy since Carl Linnaeus's classification of the genus in 1758, underscoring its role in foundational studies of reptilian morphology.^[109]

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[32]
Role of urinary and cloacal bladders in chelonian water economy
The bladder thus acts both as a store for nitrogenous waste and K+ and as a water reservoir during droughts.Missing: Testudo | Show results with:Testudo
[33]
Oxygen consumption in the tortoise, Testudo hermanni G., subjected ...
The standard metabolic rate presents an important and constant thermal dependence in the range 8-18-28 degrees C.Missing: estivation | Show results with:estivation
[34]
Aestivation in Nature: Physiological Strategies and Evolutionary ...
Aestivation is a common hypometabolic survival strategy where metabolic rate is reduced to 1–20% of the resting metabolic rate, and this inhibition is believed ...Missing: hermanni | Show results with:hermanni
[35]
The fossil record of turtles and tortoises (Reptilia: Testudines) in ...
Dec 18, 2020 · In Greece, turtle fossils have been identified in at least 68 continental and insular localities, ranging from the early Miocene to Holocene deposits.
[36]
The quaternary fossil record of the genus Testudo in the Iberian ...
We focus on two topics: (1) the presence or absence of the genus Testudo from the Early Pleistocene to the Holocene in SW Europe; and (2) the relationship ...
[37]
Testudo hermanni GMELIN, 1789 - The Reptile Database
boettgeri: coastal regions of the former Yugoslavia, through Albania, Greece, Bulgaria, to the Black Sea. Type locality: Orsova, Banat. Type locality: unknown ( ...
[38]
http://reptile-database.reptarium.cz/species?genus=testudo&species=marginata
[39]
http://reptile-database.reptarium.cz/species?genus=testudo&species=graeca
[40]
[PDF] European Red List of Reptiles - IUCN Portal
The Spur-thighed Tortoise Testudo graeca is considered to be Vulnerable (VU) at the European and EU level as it has declined by more than 30% over the last ...
[41]
Assessment of the threatened status of Testudo kleinmanni Lortet ...
Aug 7, 2025 · Testudo kleinmanni as currently defined had a historical distribution in northern coastal Egypt with two disjunct populations in Libya. Very ...
[42]
[PDF] Assessing changes in habitat quality due to land use changes in the ...
Testudo graeca appears to be an appropriate case-study to explore the effect of changes in habitat quality: positive shifts are expected at intermediate ...Missing: preferences | Show results with:preferences<|separator|>
[43]
Mediterranean tortoise: everything about its characteristics, habitat ...
Aug 12, 2025 · This animal needs Mediterranean mosaic landscapes: scrublands, meadows, roadsides, cleared holm oak forests and abandoned crops. It avoids dense ...
[44]
[PDF] Observations on ecological changes threatening a population of ...
Geology. -. The Souss Valley developed during the. Tertiary period when a variety of sedimentary rocks of predominantly calcareous lithologies was laid down.
[45]
Temperature regulation in the tortoise Testudo hermanni, studied ...
Aug 5, 2025 · Behavioural thermoregulation has been studied in the terrestrial tortoise Testudo hermanni in south-eastern France by implanting temperature-sensitive probes ...
[46]
Seasonal habitat preference by the flagship species Testudo hermanni
Hermann's tortoises are located in the xeric areas of the Mediterranean region, characterized by thermo- and meso-Mediterranean climate [9], mainly in the ...Missing: genus | Show results with:genus
[47]
Habitat Selection by the Spur-thighed Tortoise Testudo graeca in a ...
Results show that, at a landscape scale, the species selects simplified vegetation structures and includes in its home range re-colonisation shrubland and small ...Missing: genus | Show results with:genus
[48]
[PDF] Activity patterns of the spur-thighed tortoise, Testudo graeca ...
The results show that the species displays a unimodal daily activity pattern during spring, with the highest activity during the mornings, and the annual ...
[49]
[PDF] TESTUDO MARG/NA TA - The British Herpetological Society
In May the daily activity patterns of T. hermanni were unimodal, in June and August they were bimodal. I n. August during the afternoon the activity increased ...<|separator|>
[50]
[PDF] Hibernation observation
In general, it is known that in most cases Testudo hermanni and Testudo graeca hibernate from the middle of October, until the end of March. In periods of warm ...
[51]
[PDF] Aestivation observed in Testudo graeca ibera PALLAS, 1814 in ...
Oct 27, 2010 · Aestivation was observed between June and September in Spain and Morocco (DÌAZ-PANIAGUA et al. 1995; PÉREZ et al.
[52]
Activity and home range of Testudo hermanni in Northern Italy
Aug 5, 2025 · Home range size, 7.4 ha for females and 4.6 ha for males in our study area, was from three to seven times larger than that of all other ...
[53]
Annual variation of activity and daily distances moved in adult Spur ...
Aug 10, 2025 · Annual variation of activity and daily distances moved in adult Spur-thighed tortoises, Testudo graeca, in southwestern Spain. January 1995 ...
[54]
Feeding Tortoises: Avoiding Dietary Disasters
Apr 28, 2024 · The diet of Mediterranean tortoises (Testudo species) in the wild consists almost entirely of herbaceous and succulent vegetation, including ...
[55]
Ernährung im Habitat - Testudo-Farm.de
European tortoises eat plant seedlings, leaves, stems, flowers, fruits, seeds, bark, roots, and dried plants. They also eat dandelion-like plants, spicy ...
[56]
Dietary Fibre in the diet of the Herbivorous Tortoise Testudo graeca ...
Mar 30, 2024 · For many years, the Tortoise Trust has recommended a diet based upon “High fibre, low protein, low digestibility and high calcium content” ...
[57]
Seasonal Changes in the Diet of Testudo hermanni ... - Allen Press
Sep 1, 2011 · Herbivory is the dominant feeding strategy in tortoises, and dietary shifts are common in response to changes in resource availability.
[58]
http://www.tortoisetrust.org/articles/Feeding_FAQ.htm
[59]
Herbivorous reptiles and body mass: Effects on food intake, digesta ...
Here, we test these concepts in herbivorous reptiles, adding data of an experiment that measured food intake, digesta retention, digestibility and gut capacity ...
[60]
[PDF] Effects of ingesta fiber particle size on hindgut digestion and ...
Fasting stomach pH of the herbivorous Greek tortoise (Testudo graeca) has ... Digestive efficiency and gut morphology of omnivorous and herbivorous. African ...
[61]
[PDF] Feeding herbivorous reptiles and primates I. Digestive anatomy ...
Fermentation prior to enzymatic digestion and absorption: Fermentation after enzymatic digestion and absorption: ... Testudo graeca. 0. 1000. 2000. 3000. 4000.
[62]
Vocalizations and courtship intensity correlate with mounting ...
The courtship and mounting behaviour of tortoises is elaborate, and based on a multiple signalling system involving visual, olfactory and acoustic signals.Missing: ramming | Show results with:ramming
[63]
Clutch frequency, egg and clutch characteristics, and nesting activity ...
Clutch size varied from 1 to 7 eggs and averaged 3.5 eggs. Carapace length and mass of females were positively correlated with the starting date and duration of ...Missing: behavior incubation period
[64]
[PDF] TESTUDO KLEINMANN!,
Females laid 2-3 clutches per year, each consisting of 1-3 large eggs. The internesting period was estimated to be 20-30 days.
[65]
https://reptilesmagazine.com/egyptian-tortoise-breeding-observations/
[66]
[PDF] INCUBATION PERIOD AND SEX RATIO OF TESTUDO hermanni ...
Information on fertility, clutch size, incubation period and sex ratio are presented. Special attention is being given to the effect of incubation ...Missing: mating behavior
[67]
[PDF] Survival-rates-and-demography-of-the-Hermanns-Tortoise-Testudo ...
Annual survival probabilities are strongly age-dependent (mean and 95% confident interval):. 52% [0.29-0.75] for juveniles, 88 % [0.79 0.93] for immatures, 95 % ...
[68]
Variation of adult body size of the tortoise Testudo hermanni in ...
Feb 28, 2006 · The age at maturity (α) increased with body size in the 17 populations, while the Bertalanffy growth constant (k) decreased with body size. The ...
[69]
Age-size relationship of male (above) and female þelow) Testudo ...
Males reach their sexual maturity earlier than females but juvenile growth rates do not differ between sexes.
[70]
Population density and adult sex ratio of the tortoise Testudo ...
Feb 28, 2006 · The sex ratios (adult males: adult females) of most dense Greek populations were more extreme (over 1.5:1) than could be accounted for by the ...
[71]
[PDF] Testudo hermanni (Gmelin 1789) – Hermann's Tortoise
Dec 31, 2011 · The maximum number of clutches laid annually is 3, but most frequent are 1 to 2 clutches per year. Maximum clutch size is 7 eggs for T. h.Missing: fecundity | Show results with:fecundity
[72]
Adult survival and recruitment and the explanation of uneven sex ...
Aug 5, 2025 · Survival, recruitment and dynamics of adult Testudo hermanni at Alyki (N Greece) were studied from 1980-1988. Recruitment of adult males was ...
[73]
Threatened Species Status With Section 4(d) Rule for Egyptian ...
Nov 9, 2021 · We have determined that the Egyptian tortoise is likely to become endangered throughout all of its range in the foreseeable future, meeting the ...Missing: contraction | Show results with:contraction
[74]
Egyptian Tortoise (Testudo kleinmanni) IUCN Red List ... - Facebook
Jan 14, 2025 · Their population in Israel is estimated at 2000 individuals only and is considered one of the world's rarest tortoises.
[75]
Survival rates and demography of the Hermann's Tortoise Testudo ...
However, due to the loss of one study animal to poachers the survival rate was only 76%. Comparison of the annual survival rate obtained in this study with ...Missing: mortality | Show results with:mortality<|separator|>
[76]
Testudo graeca (Eastern Subspecies Clades), 120
Status. – IUCN 2022 Red List: Vulnerable (VU A1cd, assessed 1996); Regional (Europe): Vulnerable (VU A2bcde+4bcde, assessed 2009); CITES: Appendix II, as ...
[77]
Testudo graeca (Western Subspecies Clade), 117
Population stability depends mainly on adult survival. The species is still relatively common and widespread in much of North Africa. However, European ...
[78]
Assessment of the Threatened Status of Testudo werneri Perälä ...
May 1, 2006 · Current Red List Status. Endangered (A1abcd) as Testudo kleinmanni (sensu lato). Listed nationally in Israel as Endangered (EN, Acd, B) ( ...
[79]
Conservation of the endangered Mediterranean tortoise Testudo ...
This study focuses on the genetic diversity and demographic history of one of the most endangered reptiles in Europe, the western Hermann's tortoise Testudo ...
[80]
A 28-years-long monitoring study suggests heavy decline and ...
Aug 6, 2025 · We report the results of a 28-years-long (1992-2019) monitoring of three distinct populations of a tortoise, Testudo hermanni, in Central Italy.
[81]
[PDF] Egyptian tortoise - CITES
The Egyptian tortoise(Testudo kleinmanni) is a terrestrial species found in the relatively arid zone along the southeastern coast of the Mediterranean Sea.
[82]
[PDF] Trade in spur-thighed tortoises Testudo graeca in Morocco
While numbers of tortoises collected for the pet trade are lower ... Van Uhm, D.P. (2016): The Illegal Wildlife Trade: Inside the World of Poachers, Smugglers and ...
[83]
[PDF] Investigating the Illegal Online Trade of Spur-thighed Tortoises on ...
Apr 30, 2025 · These high prices might increase global trafficking of T. graeca from Iran due to the greater profit to poachers compared to domestic sales. To ...
[84]
The current distribution and status of the Hermann's tortoise ...
Jul 6, 2023 · Introduction. Hermann's tortoise, Testudo hermanni is a western Palearctic tortoise species, distributed in southeastern Europe ( Cheylan 2001 ...
[85]
[PDF] CONSERVATION BIOLOGY OF FRESHWATER TURTLES AND ...
Jan 30, 2023 · SYNONYMY. – Testudo graeca Linnaeus 1758, Testudo terrestris Forskål 1775, Testudo ecaudata. Pallas 1814, Testudo ibera Pallas 1814, Testudo ...
[86]
[PDF] International Testudo Trade - ResearchGate
The genus Testudo was included in Appendix II of. Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) as early as 1975.
[87]
CONSERVATION PROGRAMMES FOR HERMANN'S TORTOISES ...
This paper reports the conservation programme that is currently elaborated and implemented at Soptom (CRCC) for Testudo hermanni hermanni in France.
[88]
Hermann's Tortoise-World Turtle Day 2021
May 23, 2021 · Organizations in both France (SOPTOM) and Italy (CARAPAX) are head-starting juveniles for wild release in combination with rehabilitating ...
[89]
HERMANN'S TORTOISE IN THE PLAINE DES MAURES (FRANCE)
Hermann's tortoise is one of Europe's most endangered reptiles (see centrefold, Colour Plate I). Its numbers have continued to decrease dramatically over the ...
[90]
The Albera Tortoise Project | hermannihavenhome
Testudo hermanni populations on the Balearic Islands were introduced by sailors around 3000 years ago, presumably as a fresh food source (Mayol 1985, Pinya ...
[91]
Release captive-bred reptiles into the wild: Tortoises, terrapins, side ...
All five released captive-bred subadult ploughshare tortoises survived at least one year in the wild, and settled within 138–523 m of the release site.
[92]
[PDF] Global Re-introduction Perspectives: 2013
Livoreil (2009) Distribution of the endangered Hermann's tortoise Testudo hermanni hermanni in Var, France, and recommendations for its conservation. Oryx, 43(2): ...
[93]
Home range area in the tortoise Testudo hermanni in relation to ...
Aug 6, 2025 · The need for reserves to include different vegetation types makes conservation of T. hermanni in France more difficult. Conversely, the need ...
[94]
Global Testudo Trade: Update and Recent Trends - Allen Press
Jan 6, 2025 · ... IUCN Red List and assessed in 2003 and 2022 as “Critically Endangered”. The trade in T. marginata has similarly been small, with a maximum ...
[95]
Illegal Imports - Tortoise Trust Web
Tortoises are illegally imported, often labeled "captive-bred" with false CITES documents, and are likely wild-caught, not from genuine breeding programs.
[96]
[PDF] p. 1 AC23 Doc. 8.4 CONVENTION ON INTERNATIONAL TRADE IN ...
Aug 11, 2007 · Testudo species including the Spur-thighed Tortoise are popular in the pet trade. All species of. Testudinidae have been included in the CITES ...
[97]
Extent of introgressive hybridization in the Hermann's tortoise ...
May 19, 2022 · The Western subspecies of Hermann's tortoise (Testudo hermanni hermanni: WT) is threatened by habitat loss, fragmentation, wildfires, illegal ...Missing: keeping | Show results with:keeping
[98]
Nutrition and husbandry conditions of Palearctic tortoises (Testudo ...
Apr 2, 2018 · A survey was developed to evaluate the current population structure, husbandry conditions, diet regime, and health status of Testudo species in captivity.Missing: challenges mortality hybridization<|separator|>
[99]
Hermann's Tortoise Lifespan
Both the Eastern and Western Hermann's tortoise lifespan is anywhere from 50 to 100 years. It is quite possible that they can live even longer than 100 years.Missing: longevity | Show results with:longevity
[100]
[PDF] REGULATIONS - EUR-Lex - European Union
May 17, 2023 · (1). Regulation (EC) No 338/97 regulates trade in animal and plant species listed in the Annex to that Regulation. ... Testudo hermanni (II).
[101]
TRAFFIC | Captive breeding
Many reptiles, amphibians and fish are captive bred for the pet trade, which can contribute to the protection of wild populations.Missing: Testudo | Show results with:Testudo
[102]
(PDF) International Testudo Trade - ResearchGate
Aug 7, 2025 · ... illegal wildlife trade, and certain EU countries function as transit points for illegal wildlife trafficking. Like Europe, the Middle East ...
[103]
A Summary and Analysis of Aesop's 'The Hare and the Tortoise' Fable
Feb 26, 2021 · The fable of the hare and the tortoise usually runs something like this. A hare was making fun of a tortoise for moving so slowly.
[104]
Testudo - Roman Military Research Society
May 7, 2025 · Testudines, is based on the Latin word · testudo meaning “tortoise” and was coined by German naturalist August Batsch in 1788. The word “turtle”, ...
[105]
Greek tortoise in Ancient Stageira - My Favourite Planet
An anecdote related by the Roman author Pliny the Elder (23-79 AD) claims that a tortoise was involuntarily responsible for the death of the Athenian playwright ...Missing: significance | Show results with:significance
[106]
The armoured tortoise inspired the resilient Roman testudo
Oct 7, 2013 · The Romans used to drive horses and carts over their tortoises – their human ones, that is! The testudo (tortoise) was one of their classic weapons.
[107]
Testudo | macmil - Digital Exhibitions
The original statue, which now stands in front of McKeldin Library, was the gift of the Class of 1933 whose members felt it was high time that the University ...
[108]
[PDF] The cosmic symbolism of tortoises and turtles
The Greek, and later the Romans, had a certain tradition, which associated the constellation of Lyre with a tortoise (Scherer 1953). They referred the. Kithara ...<|separator|>
[109]
[PDF] A Manual of Elementary Geology; Or, The Ancient ... - Darwin Online
... Adolphe Brongniart when founding his classification of the fossiliferous strata in reference to their imbedded fossil plants, has placed the cretaceous ...