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Centrochelys

Centrochelys is a monotypic genus of tortoise in the family Testudinidae, consisting solely of the species Centrochelys sulcata, commonly known as the African spurred tortoise or sulcata tortoise.^[1] It is the largest tortoise species native to mainland Africa and the third-largest tortoise species worldwide, with males reaching up to 35 inches (89 cm) in length and females up to 26 inches (66 cm).^[2] Native to the semi-arid Sahel region along the southern edge of the Sahara Desert, this species inhabits dry savannas, grasslands, shrublands, and woodlands, where it digs extensive burrows for shelter during extreme heat or drought.^[2] Classified as Endangered on the IUCN Red List (as of 2021), Centrochelys sulcata faces declining populations due to habitat degradation, overcollection for the pet trade, and human-wildlife conflicts.^[3] The genus Centrochelys derives its name from the Latin word centrum (meaning "center" or "midpoint") combined with the Greek chelys (meaning "tortoise"), while the species epithet sulcata refers to the distinctive grooved or sulcate pattern on its carapace.^[1] First described as Testudo sulcata by John Frederick Miller in 1779, the species was previously included in the genus Geochelone but has since been placed in the distinct genus Centrochelys based on phylogenetic analyses to reflect its position relative to other African tortoises.^[1] The carapace is high-domed and features prominent growth rings with deep sulci, particularly pronounced in adults, and the species is notable for its large spurs on the thighs and forelimbs, which aid in digging.^[2] Adults exhibit sexual dimorphism, with males having a concave plastron, longer tails, and more aggressive behavior during breeding, which occurs in the rainy season from February to March.^[2] Centrochelys sulcata is distributed across a broad but fragmented range in sub-Saharan Africa, from Senegal in the west through Mauritania, Mali, Niger, Chad, Sudan, Eritrea, Ethiopia, and south to Cameroon and South Sudan.^[1] It prefers arid to semi-arid environments with sparse vegetation, where it forages primarily on grasses, succulents, and leaves, showing a particular preference for morning glory vines.^[2] The tortoise's lifespan can exceed 50 years in the wild and up to 150 years in captivity, with sexual maturity reached around 15 years of age; females lay clutches of 15–20 eggs that incubate for 120–212 days depending on temperature, which also determines offspring sex.^[2] Despite its adaptability to harsh conditions through burrowing, the species has been introduced to regions like the United States and the British Virgin Islands, where it sometimes establishes feral populations.^[1] Conservation efforts for Centrochelys sulcata are challenged by its extensive range across politically unstable areas, but key threats include agricultural expansion leading to habitat loss, illegal poaching for bushmeat and traditional medicine, and the international pet trade, which has prompted CITES Appendix II listing to regulate commerce.^[3] Population estimates are unavailable due to the remote habitats, but overall trends indicate a significant decline—leading to its reassessment as Endangered in 2021—prompting recommendations for protected area expansion and community-based management in the Sahel.^[3] Research highlights the species' ecological role in seed dispersal and soil aeration through its burrowing activities, underscoring the need for sustained conservation to prevent further vulnerability.^[3]

Taxonomy

Etymology

The genus name Centrochelys derives from the Latin centrum (center or midpoint) and the Greek chelys (tortoise).^[1] The species epithet sulcata originates from the Latin sulcatus (grooved or furrowed), describing the distinctive deep grooves and furrows on the carapace scutes.^[4] Originally described as Testudo sulcata by John Frederick Miller in 1779, based on an illustration in his work Icones Avium, the taxon was later transferred to the monotypic genus Centrochelys by John Edward Gray in 1872 to reflect its distinctive limb morphology within the Testudinidae.^[5]^[6]

Classification

Centrochelys is classified within the kingdom Animalia, phylum Chordata, class Reptilia, order Testudines, suborder Cryptodira, superfamily Testudinoidea, family Testudinidae, subfamily Testudininae, and genus Centrochelys.^[1]^[7] The genus contains a single extant species, Centrochelys sulcata, which serves as the type species, originally described as Testudo sulcata by Miller in 1779.^[1]^[8] Molecular phylogenetic analyses position Centrochelys as basal within Testudininae, forming a sister group to a diverse clade that includes genera such as Stigmochelys, Chelonoidis, and Geochelone.^[9] No subspecies are currently recognized for C. sulcata, although molecular studies have identified three distinct mitochondrial DNA haplotypes corresponding to eastern, western, and central African populations, which have not been formalized taxonomically due to low genetic divergence.^[1]

Extinct species

Several extinct species have been attributed to the genus Centrochelys, primarily from Pleistocene and Holocene fossil records in the Macaronesian islands of the Atlantic Ocean, with additional historical placements in the Mediterranean. These taxa often exhibit island gigantism, characterized by larger body sizes compared to mainland relatives, adapted to isolated volcanic environments. Fossil evidence includes bones, carapaces, and eggshells, suggesting terrestrial habits in arid or semi-arid habitats. Most extinctions occurred between approximately 1,000 and 5,000 years ago, coinciding with human colonization, habitat alteration, and climate fluctuations in the Holocene, though some predate human arrival due to volcanic activity.^[10]^[11] In the Canary Islands, Centrochelys burchardi is known from Middle Pleistocene deposits (ca. 670,000–200,000 years ago) on Tenerife, where remains including carapaces (80–100 cm in length), limb bones, and fossilized eggshells have been recovered from volcanic quarries such as Adeje. This species displayed gigantism typical of insular testudinids, with robust limbs suited for foraging in low-vegetation landscapes. Its extinction likely predates human settlement around 2,500 years ago, attributed to recurrent volcanic eruptions and aridification rather than anthropogenic factors.^[11]^[12] Similarly, Centrochelys vulcanica from Gran Canaria shares a comparable temporal and ecological context, with fossil eggshells and shell fragments indicating a large-bodied form (carapace up to 90 cm) endemic to the island's volcanic terrains; it also went extinct in the Pleistocene, possibly due to the same geological instability.^[11]^[13] Further west in the Cape Verde archipelago, two unnamed Centrochelys species are documented. One from Sal Island, represented by subfossil remains in the Pedra de Lume volcanic crater, dates to the Pleistocene and exhibits gigantism with estimated carapace lengths exceeding 60 cm; this population became extinct in "deep time" prior to human contact, likely from volcanic burial or climatic shifts. The second, from Maio Island, persisted into the Holocene and vanished shortly after Portuguese colonization around 1,000 years ago, with bones and eggshells linked to human-induced habitat loss and predation.^[10]^[14] A 2021 ancient DNA study revealed that the previously named Centrochelys atlantica (based on 19th-century type material from Sal) actually represents misidentified specimens of the extant South American red-footed tortoise Chelonoidis carbonarius, leaving the true Sal species without a formal binomial.^[14] Taxonomic revisions in the 2020s have refined the genus boundaries. For instance, Centrochelys robusta, originally described from Pleistocene cave deposits on Malta, was reassigned to the distinct extinct genus Solitudo as S. robusta based on morphological analyses of humeri and femora, which show unique insular adaptations diverging from African Centrochelys lineages. This move highlights ongoing debates over Mediterranean Pleistocene testudinids, with potential additional unnamed Centrochelys-like species in Sicilian and Menorcan fossils awaiting genetic confirmation, though many are now classified under Solitudo due to phylogenetic distinctions.^[13]^[15]

Description

Morphology

The African spurred tortoise, Centrochelys sulcata, exhibits a robust morphology adapted to arid environments, characterized by a heavily armored shell and sturdy limbs suited for digging. The carapace is strongly convex and oval-shaped, typically sandy yellow-brown in color with darker growth lines and borders along the scutes, featuring deep sulci that create a furrowed, spur-like pattern.^[16] These sulci separate the scutes, including five vertebral, four paired costal, and eleven marginal scutes per side, with the posterior marginals upturned and serrated except along the bridge; there is no nuchal scute, and the supracaudal is single.^[16] The plastron is large, composed of six pairs of yellow-brownish scutes, and is flat in females but concave in males, providing a stable base for the tortoise's weight during locomotion and burrowing activities.^[16] The head is moderately large and covered in large scales, with a slightly hooked upper jaw that lacks a pronounced beak, enabling efficient herbivory through strong, crushing bites.^[17] The limbs are thick and powerful, with the forelimbs bearing large, overlapping scales and five claws, ideal for excavating burrows, while the hindlimbs have four claws and are more rounded with distinctly pronounced scales, including prominent spurs on the thighs that give the species its common name.^[16] These spurs and the elephantine hindlimbs contribute to the overall rugged build, enhancing traction in loose substrates.^[17] Sexual dimorphism is evident in adults, with males possessing a concave plastron, forked or diverged gular scutes, widely flared anal scutes, and a longer, thicker tail, alongside generally larger body proportions compared to females.^[16] Females, in contrast, have a flat plastron, unforked gular scutes, narrower anal scutes, and a shorter, blunter tail, with a more rounded overall shell shape that supports egg-laying.^[16] Hatchlings show minimal dimorphism and are uniformly yellow, darkening to the adult coloration with age.^[16]

Size and lifespan

The African spurred tortoise (Centrochelys sulcata), the only extant species in the genus, displays pronounced sexual dimorphism in body size. Adult males average 70-80 cm in straight carapace length (SCL), with maximums reaching 86 cm SCL and 101 cm curved carapace length (CCL), and weights up to 100 kg, though some exceed 120 kg. Females are notably smaller, attaining up to 57.8 cm SCL and averaging 37 kg (range 30-50 kg). This makes C. sulcata the third-largest tortoise species globally, surpassed only by the Galápagos tortoise (Chelonoidis nigra) and Aldabra giant tortoise (Aldabrachelys gigantea).^[18]^[19]^[20] Growth rates are rapid in the early life stages, particularly during the first five years, where juveniles increase significantly under favorable conditions, before decelerating significantly after sexual maturity at 10-15 years of age, when individuals typically measure 35-50 cm SCL and weigh 15-20 kg. Growth rates exhibit high plasticity, with captive individuals growing 1.4–2.6 times faster than wild ones due to dietary differences.^[18]^[21]^[19] Lifespan varies by environment: in the wild, C. sulcata is estimated to exceed 75 years, while in captivity, individuals commonly surpass 100 years, with the oldest recorded specimen over 120 years old.^[18]^[19]^[22]

Distribution and habitat

Geographic range

The African spurred tortoise (Centrochelys sulcata), the sole extant species in the genus Centrochelys, occupies the southern fringes of the Sahara Desert and the Sahel region across sub-Saharan Africa. Its current distribution spans from Senegal and Mauritania in the west, extending eastward through Mali, Burkina Faso, Niger, northern Nigeria, Chad, and Central African Republic, to Sudan, Eritrea, Ethiopia. Marginal populations occur in northern Algeria and extend to the southern Arabian Peninsula, including Saudi Arabia and Yemen. A 2023 study confirmed the species' presence in northern Algeria, extending into the northern Sahara.^[16]^[23] Historically, the species' range was broader, but it has undergone significant contraction since the early 20th century due to desertification, agricultural expansion, and overgrazing. It is likely extirpated from Djibouti, Togo, and parts of West Africa, including Cameroon and northern Benin, where local populations have vanished amid habitat degradation and human pressures. These declines have fragmented the remaining distribution into isolated pockets within semi-arid savannas and grasslands.^[16]^[18] Fossil evidence from extinct Centrochelys species reveals a more extensive Pleistocene distribution, suggesting the genus once inhabited Mediterranean islands such as Malta and the Canary Islands (e.g., Tenerife and Gran Canaria), as well as Atlantic islands like Cape Verde. Remains of species like C. atlantica in Cape Verde and C. burchardi in the Canary Islands indicate adaptation to insular environments, pointing to a wider historical range before isolation and extinction events.^[24]^[25] Within its contemporary range, C. sulcata displays nomadic movements, traveling considerable distances to follow rainfall and locate temporary water sources and fresh vegetation in arid landscapes.

Habitat preferences

Centrochelys sulcata primarily inhabits arid to semi-arid regions, favoring Sahelo-Sudanian savannas, dry grasslands, and thorn scrub dominated by vegetation such as Acacia spp. and Adansonia digitata. These habitats typically occur at elevations between 200 and 700 m, in areas receiving annual rainfall of 140–600 mm, where the species exploits seasonal vegetation growth during brief wet periods.^[16]^[26] A key aspect of their habitat use involves extensive burrowing, with C. sulcata constructing tunnel systems up to 15 m in length within sandy soils of stabilized dunes or dry riverbeds (known as kori). These burrows serve for thermoregulation, allowing the tortoises to maintain stable body temperatures amid extreme diurnal fluctuations, and for aestivation during the prolonged dry seasons when surface activity ceases.^[16]^[27] In terms of microhabitat selection, C. sulcata shows a strong preference for sites near intermittent water sources like kori streams, which provide moist soils and access to emergent vegetation post-rainfall, while avoiding dense forests, extreme desert dunes, bare sandy expanses (zipele), and areas with high human or livestock disturbance. This selection ensures proximity to foraging opportunities without excessive exposure to predation or aridity.^[26]^[16]^[27]

Behavior

Activity patterns

Centrochelys sulcata exhibits a predominantly diurnal activity pattern, with peak above-ground activity occurring in the early morning and late afternoon, forming a bimodal diel cycle. Individuals emerge from burrows to bask, regulating their body temperature through behavioral thermoregulation before foraging. As ambient temperatures exceed 32°C during midday, they retreat to burrows to avoid overheating, a strategy that also facilitates evaporative cooling via salivation if necessary.^[28]^[29]^[16] Seasonally, activity intensifies during and immediately following the wet season, particularly from August to November, when post-rain vegetation growth supports foraging excursions. In contrast, movement is minimal during the extended dry periods, with individuals entering aestivation in burrows during the hot dry season to conserve energy amid high temperatures and resource scarcity. This pattern aligns with the Sahelian climate, where rainfall triggers bursts of activity.^[28]^[16] Locomotion in C. sulcata is characteristically slow. Males exhibit greater mobility during the breeding period, potentially traveling 10–20 km or more to locate females, contrasting with the more localized movements of females and juveniles. Social interactions, such as aggressive encounters, occasionally occur during these active periods but remain infrequent.^[16] Centrochelys sulcata, the sole extant species in the genus, exhibits a predominantly solitary lifestyle, with adults residing alone in individual burrows and minimizing interactions with conspecifics except during breeding.^[19] This asocial behavior aligns with the species' low population densities in the wild, typically ranging from 0.001 individuals per hectare, reflecting sparse distribution across arid savannas.^[19] Males display strong territoriality, defending areas around their burrows through aggressive actions such as ramming and head-butting conspecifics, where larger males dominate combats to secure resources and mates.^[19]^[30] Females, in contrast, show greater tolerance toward juveniles, permitting multiple young individuals to occasionally share burrows while adults maintain separate residences, often positioned 100–200 meters apart.^[19] No evidence of hierarchical packs exists, though loose aggregations may form near scarce water sources during active wet-season periods.^[19] In captivity, Centrochelys sulcata individuals remain generally docile toward humans but become increasingly combative with maturity, engaging in ramming and dominance displays toward other tortoises, which intensifies territorial conflicts in confined spaces.^[31]^[17]

Ecology

Diet

Centrochelys sulcata is primarily herbivorous, with its diet consisting mainly of grasses such as Dactyloctenium aegyptium, Digitaria ciliaris, Eleusine indica, and Eragrostis tenella, succulents like Portulaca oleracea, and shrubs including Leptadenia hastata.^[16] It opportunistically consumes flowers, fruits, and other herbaceous plants, particularly following rainfall when vegetation is more abundant in its arid Sahelian habitat. While primarily herbivorous, it occasionally consumes carrion or garbage near human settlements.^[16] The species employs a grazing foraging strategy, browsing on low-lying vegetation in open savannahs and dune areas where food is available.^[16] Its high-fiber intake necessitates specialized digestion, including geophagy to obtain calcium and minerals from soils.^[32] Seasonal variations influence feeding patterns, with active foraging on fresh plants during the wet season (typically August) when activity peaks in the early morning; during the dry season, individuals rely on stored body fat and consume drier, withered vegetation while aestivating in burrows.^[16] Juveniles incorporate more protein-rich plant items, such as tender succulents and herbs, to support growth.^[33] Digestive adaptations include a slow metabolism suited to sparse resources and hindgut fermentation in the gastrointestinal tract, enabling efficient breakdown of fibrous cellulose from plant material.^[16]

Reproduction

Reproduction in Centrochelys sulcata is seasonal and tied to environmental cues, particularly the post-rainy period. Mating typically occurs from September to November, following the rainy season, when males actively search for females.^[16] Courtship involves males circling and mounting females, often accompanied by aggressive ramming of rivals to assert dominance based on size and strength; females appear to select mates exhibiting these dominant traits.^[17] Actual copulation lasts about 15 minutes and may be repeated several times daily over a week.^[34] Approximately 60 days after mating, gravid females roam to locate suitable nesting sites, excavating multiple test pits—up to four or five—over 5–15 days before depositing eggs in a final flask-shaped chamber.^[17] The nest is a shallow pit roughly 15–20 cm deep and 60 cm wide, dug primarily with the hind legs.^[20] From September to May, females produce 2–3 clutches annually, each containing 14–40 eggs (average 18–25), which are white, spherical, and measure about 52 × 44 mm with a mass of 55 g.^[18] Clutch success and overall fecundity (around 45 eggs per year) are influenced by rainfall patterns, as adequate precipitation supports vegetation and nesting conditions; droughts can reduce breeding frequency and survival.^[16] Eggs incubate in the soil for 85–120 days at temperatures around 30°C and 70–80% humidity, with hatching often synchronized over a few days. As in many turtles, sex is determined by incubation temperature, with warmer temperatures producing females and cooler ones males.^[18] Hatchlings emerge fully independent, measuring 4–7 cm in straight carapace length and weighing 20–46 g, with yellow-tan coloration and serrated shells for burrowing.^[17] They must immediately seek shelter to evade predators such as monitor lizards and birds, facing high mortality rates of 50–70% in the early stages due to predation and environmental stressors.^[16] Sexual maturity is reached at 10–15 years of age, with females maturing slightly earlier (10–12 years) than males (13–15 years), typically at a carapace length of 35–50 cm and body mass of 15–20 kg.^[35]

Conservation

Status and threats

The African spurred tortoise (Centrochelys sulcata), the only extant species in the genus Centrochelys, is classified as Endangered on the IUCN Red List under criteria A4bcd, based on an assessment conducted in 2020 and published in 2021.^[3] This status reflects an estimated population decline of 50–75% over approximately 90 years (encompassing the past 60 years and the next 30 years), driven by ongoing habitat degradation and exploitation.^[35] The species now has a highly fragmented distribution into isolated subpopulations across the Sahel and southern Sahara regions.^[16] Population estimates are imprecise due to the vast and remote habitat, but late 1990s surveys suggested 18,000–20,000 individuals total (including immature), with fewer than 50,000 mature individuals inferred from density data (e.g., 0.0021–0.167 individuals per hectare in surveyed areas) and ongoing declines; current numbers are uncertain but the overall trend is decreasing.^[35]^[16] The primary threat to C. sulcata is habitat loss and degradation, primarily from agricultural expansion, overgrazing by livestock, desertification, and wildfires, which collectively account for the majority of impacts (estimated at around 60% of the decline's drivers based on regional threat modeling).^[35]^[36] These pressures exacerbate range contraction, as high livestock densities (e.g., >10 cattle per km²) correlate strongly with tortoise absence, while fires amplify this effect by destroying vegetation cover essential for foraging and shelter.^[36] Additional threats include direct exploitation through the illegal pet trade, with at least 9,132 wild individuals reported in international trade between 1990 and 2010 (averaging ~435 per year, predominantly of unknown origin), alongside collection for meat, eggs, and traditional medicine.^[36] Roadkill from increasing vehicle traffic and competition with livestock for resources further compound mortality, particularly in fragmented habitats near human settlements.^[35] Climate change intensifies desertification, reducing suitable arid savannah and grassland patches.^[35] In 2023, the species' range was extended to include Algeria.^[37] Historical extinctions within the genus provide context for current vulnerabilities, as human activities have driven range contractions with parallels to modern threats. For instance, Centrochelys burchardi, a giant tortoise endemic to Tenerife in the Canary Islands, went extinct during the Middle Pleistocene due to environmental changes and possible early human impacts, mirroring the exploitation and land-use changes now affecting C. sulcata.^[12] Such patterns underscore the genus's sensitivity to anthropogenic pressures, with C. sulcata's fragmented subpopulations at high risk of local extirpations if threats persist.^[35]

Protection efforts

Centrochelys sulcata is listed under Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) since 1977, as part of the family Testudinidae listing, which regulates international trade to prevent overexploitation. A zero annual export quota has been established for wild specimens traded for primarily commercial purposes, further restricting legal trade. Ongoing CITES reviews as of 2025 continue to monitor trade impacts.^[38]^[39] In range countries, the species receives protection within designated areas, such as Senegal's Ferlo Nord and Ferlo Sud Faunal Reserves managed by the National Parks Department, and Chad's Zakouma National Park.^[34] Conservation initiatives are led by organizations like the IUCN SSC Tortoise and Freshwater Turtle Specialist Group (TFTSG), which has produced detailed action plans, including habitat protection and population assessments across the Sahel.^[16] Reintroduction efforts have occurred in Senegal's North Ferlo region, with tortoises from captive programs released to bolster wild populations between 2013 and 2014 in collaboration with local authorities.^[40] Anti-poaching patrols in Chad's Zakouma National Park actively deter illegal collection and habitat encroachment, protecting resident C. sulcata through ranger enforcement and community partnerships. Captive breeding programs in zoos worldwide have proven successful, producing over 100 hatchlings annually on average, with institutions like Linton Zoo in the UK achieving record clutches of 45 in a single year as part of long-term efforts.^[41] These programs support head-start initiatives, where juveniles are reared in controlled environments to increase survival rates before release into protected habitats, such as those in Senegal.^[42] Ongoing research highlights gaps, including the need for genetic studies to assess haplotype diversity across fragmented populations and enhanced monitoring using camera traps to track densities in the Sahel.^[16] Community education campaigns in Sahel countries, led by groups like the African Chelonian Institute, focus on reducing illegal trade by raising awareness among locals about the species' vulnerability and promoting sustainable alternatives to collection.^[42]

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[40]
Exploring the main threats to the threatened African spurred tortoise ...
Jan 23, 2017 · The African spurred tortoise Centrochelys sulcata is the second largest terrestrial turtle, with a scattered distribution across the West African Sahel.Missing: etymology | Show results with:etymology<|control11|><|separator|>
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Centrochelys sulcata - CITES
Centrochelys sulcata ; Kingdom. Fauna ; Phylum. Chordata ; Class. Reptilia ; Order. Testudines ; Family. Testudinidae.<|separator|>
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Program for the conservation of the African spurred tortoise and its ...
The first to benefit from the program were two women's groups in Ranérou (GPF Baamtaré and GIE Kaoral). ... The African spurred tortoise (Centrochelys sulcata).<|control11|><|separator|>
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Tortoise hatches 'record' 45 babies at Linton Zoo - BBC News
Dec 21, 2011 · Linton Zoo has run an African spurred tortoise breeding programme for 15 years, and director Kim Simmons said this was its most successful herd.
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Projects - African Chelonian Institute
Sulcata Preserve Project (SSP) Since 2022, ACI has identified Senegal's last wild population of Centrochelys sulcata. The project now focuses on securing ...