Animal Diversity Web

Ge­o­graphic Range

Leop­ard tor­toises (Psam­mo­bates pardalis) are en­demic to Africa, and their ge­o­graphic range ex­tends from Sudan to Ethiopia and from Natal in east­ern Africa to south­ern An­gola and South Africa. They are also dif­fusely dis­trib­uted through­out por­tion of south­west­ern Africa. (Bartlett, et al., 2006; Dou­glas and Rall, 2006; Mc­Mas­ter and Downs, 2009)

• Biogeographic Regions
• ethiopian
  • native

Habi­tat

Leop­ard tor­toises oc­cupy a va­ri­ety of xeric and mesic habi­tats through­out their ge­o­graphic range, rang­ing from dry arid plains to tem­per­ate grass­land ecosys­tems. They are in­tol­er­ant of damp or cold habi­tats. They have also been re­ported in moun­tain­ous ter­rain. These tor­toises spend most of their time in shrub habi­tat with low lying veg­e­ta­tion, which serves as their pri­mary food source. In the sum­mer, they seek shel­ter under var­i­ous low lying plants. (Bartlett, et al., 2006; Dou­glas and Rall, 2006)

• Habitat Regions
• temperate
• terrestrial

• Terrestrial Biomes
• desert or dune
• savanna or grassland
• forest
• mountains

Phys­i­cal De­scrip­tion

Leop­ard tor­toises are the fourth largest tor­toise species in the world. They ex­hibit in­de­ter­mi­nate growth and adults range in mass from 15 to 54 kg, with an av­er­age of 18 kg. Cara­pace length ranges from 30 to 70 cm, with an av­er­age cara­pace length of 45 cm. Fe­males are often larger than males. It also is com­mon for leop­ard tor­toises to have mod­er­ate cara­pa­cial pyra­mid­ing, a shell de­for­mity in which scutes ex­hibit pyra­mi­dal growth. Leop­ard tor­toises dif­fer from other mem­bers of the genus Geoch­e­lone due to their dis­tinct shell mark­ings. Base color of the cara­pace may be tan, yel­low, or some­times shades of dusty brown. The in­ten­sity of shell pat­tern­ing varies. Blotches on the shell are most often black and are typ­i­cally only pre­sent on ju­ve­niles. The head, feet, and tail vary in color but are usu­ally tan to brown. De­spite dif­fer­ences in ap­pear­ance, the diet and habi­tat of tor­toises in the genus Geoch­e­lone are usu­ally sim­i­lar. (Bartlett, et al., 2006; Leu­teritz and Ekbia, 2008; Schmidt, 2006; Simang, et al., 2010)

• Other Physical Features
• ectothermic
• heterothermic
• bilateral symmetry

• Sexual Dimorphism
• female larger

• Range mass

  15 to 54 kg

  33.04 to 118.94 lb

• Average mass

  18 kg

  39.65 lb

• Range length

  30 to 70 cm

  11.81 to 27.56 in

• Average length

  45 cm

  17.72 in

De­vel­op­ment

Leop­ard tor­toises are oviparous, and their eggs are ini­tially leather-like but dry to be­come hard and brit­tle. The egg con­tains a large, nu­tri­ent-rich yolk, which the em­bryo feeds on dur­ing de­vel­op­ment. Sex de­ter­mi­na­tion is tem­per­a­ture de­pen­dent; fe­males de­velop under av­er­age tem­per­a­tures of 30 C or more and males de­velop under av­er­age tem­per­a­tures of less than 30 C. Tem­per­a­ture-de­pen­dent sex de­ter­mi­na­tion sug­gests that the mother has some con­trol over the sex ratio of her off­spring. If she lays her eggs in a warm en­vi­ron­ment the sex ratio fa­vors fe­males, whereas a cool en­vi­ron­ment fa­vors males. Eggs are roughly spher­i­cal and about 57.5 mm in di­am­e­ter. Clutch size ranges from 7 to 20 eggs, and most eggs hatch within 47 to 180 days. After hatch­ing, young tor­toises grow very rapidly dur­ing the first few months of life. In gen­eral, hatch­lings in this genus weigh around 100 grams and grow be­tween 300 and 400 grams dur­ing their first year of life. (Bow­man, 1966; Deem­ing and Fer­gu­son, 1991; Spotila, et al., 1994; Stan­ford, 2010)

• Development - Life Cycle
• temperature sex determination

Re­pro­duc­tion

Leop­ard tor­toises are monog­a­mous, and both males and fe­males be­come in­creas­ingly ag­gres­sive when search­ing for a mate. They fight for mates by butting and ram­ming ri­vals. Dur­ing courtship, males fol­low their po­ten­tial mate and re­peat­edly run into them until the fe­males be­come im­mo­bi­lized. Males some­times lift their mate off the ground by ram­ming them. Fe­males be­come de­fen­sive and in some cases try to es­cape. Dur­ing cop­u­la­tion, males mount the fe­male and ex­tend their necks and grunt dur­ing mat­ing. (Ernst and Bar­bour, 1989; Leu­teritz and Ravolanaivo, 2005)

• Mating System
• monogamous

Leop­ard tor­toises breed from May to Oc­to­ber. Males be­come re­pro­duc­tively ma­ture by five years of age, and al­though the spe­cific age of mat­u­ra­tion is un­known, fe­males are thought to be­come re­pro­duc­tively ma­ture later than males. After mat­ing, fe­males dig a hole in the ground, rang­ing from 100 to 300 mm deep, in which to lay her eggs. The frail eggs are white and spher­i­cal. Ges­ta­tion ranges from 9 to 12 months and varies ac­cord­ing to lo­ca­tion, tem­per­a­ture, and pre­cip­i­ta­tion. Fe­males lay 5 to 7 clutches dur­ing a sin­gle breed­ing sea­son, with each clutch sep­a­rated from the pre­vi­ous by about 3 to 4 weeks. Clutches range from 5 to 30 eggs, with larger fe­male hav­ing larger clutches. (Ernst and Bar­bour, 1989; Leu­teritz and Ravolanaivo, 2005)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• fertilization
• oviparous

• Breeding interval

  Leopard tortoises breed once yearly.

• Breeding season

  Leopard tortoises breed from May to October.

• Range number of offspring

  5 to 30

• Range gestation period

  9 to 14 months

• Average gestation period

  12 months

• Range age at sexual or reproductive maturity (male)

  5 to 6 years

• Average age at sexual or reproductive maturity (male)

  5 years

Pa­ter­nal care in leop­ard tor­toises is non-ex­is­tent, as males leave di­rectly after cop­u­la­tion. Fe­males dig a hole in which to lay their eggs, which ranges in depth from 100 to 300 mm. After the eggs are laid, she cov­ers them and leaves. Hatch­lings are im­me­di­ately in­de­pen­dent upon emerg­ing. (Bartlett, et al., 2006; Ernst and Bar­bour, 1989)

• Parental Investment
• no parental involvement

Lifes­pan/Longevity

In the wild, adult leop­ard tor­toises may live for up to 100 years. No records are avail­able re­gard­ing cap­tive in­di­vid­u­als. How­ever, typ­i­cal lifes­pan for other species of Geoch­e­lone tor­toises in cap­tiv­ity is ap­prox­i­mately 50 years. Fac­tors that may limit the lifes­pan of leop­ard tor­toises in­clude human im­pacts through the pet trade and male ag­gres­sion dur­ing courtship and mat­ing. (Bartlett, et al., 2006; Ernst and Bar­bour, 1989; Wim­berger, et al., 2009)

• Range lifespan
  Status: wild

  100 (high) years

• Average lifespan
  Status: captivity

  50 hours

Be­hav­ior

Au­di­tory per­cep­tion is im­por­tant to leop­ard tor­toise mat­ing be­hav­ior, as males vo­cal­ize species-spe­cific calls in an at­tempt to at­tract fe­males. Males also vo­cal­ize dur­ing com­pe­ti­tion for mates. Male vo­cal­iza­tions in­clude a va­ri­ety of grunts and groans. Al­though tor­toises do not au­di­bly re­spond to vo­cal­iza­tions, most are vi­su­ally re­spon­sive. Like most other tor­toises, leop­ard tor­toises are known for their hiss­ing noises. When feel­ing threat­ened, tor­toises quickly pull the head and feet into the shell, re­sult­ing in air being forced from the lungs and thus the dis­tinc­tive hiss­ing noise is pro­duced. Males pos­sess neck glands, which in­form con­specifics of their will­ing­ness to mate or bat­tle an­other male. Ev­i­dence sug­gests that, dur­ing mat­ing sea­son, male tor­toises fol­low fe­males for days or even weeks at a time be­fore ini­ti­at­ing courtship. In order to ini­ti­ate con­tact, males bite the fe­male on the head or feet. So­cial hi­er­ar­chies are es­tab­lished via ag­gres­sion, as dom­i­nance is es­tab­lished through a se­ries of com­pe­ti­tions. So­cial net­works among the tor­toises are fairly weak, with lit­tle in­traspe­cific com­mu­ni­ca­tion oc­cur­ring. (Auf­fen­berg, 1977)

• Key Behaviors
• terricolous
• motile
• social

Home Range

There is no in­for­ma­tion avail­able re­gard­ing the av­er­age home range size of leop­ard tor­toises. (Ernst and Bar­bour, 1989)

Com­mu­ni­ca­tion and Per­cep­tion

Com­mu­ni­ca­tion and per­cep­tion within leop­ard tor­toises is pri­mar­ily vi­sual, how­ever, there is lit­tle ev­i­dence to sug­gest that tor­toises are able to dis­tin­guish be­tween dif­fer­ent col­ors. A re­cent study pro­posed ju­ve­nile leop­ard tor­toises most often ap­proach col­ors such as red and dif­fer­ent shades of green. In gen­eral, tor­toises are par­tic­u­larly sen­si­tive to sounds under 1,000 Hz. (Auf­fen­berg, 1977; Simang, et al., 2010)

• Communication Channels
• visual
• tactile
• acoustic
• chemical

• Perception Channels
• visual
• tactile
• acoustic
• chemical

Food Habits

Leop­ard tor­toises are pri­mar­ily her­biv­o­rous, with plant ma­te­r­ial mak­ing up ap­prox­i­mately 98% of their diet. They con­sume berries and other fruits when avail­able. Bone frag­ments and ash may be con­sumed dur­ing times of de­creased re­source abun­dance. Al­though most tor­toises con­sume mostly grasses, leop­ard tor­toises pri­mar­ily con­sume forbs. They feed pri­mar­ily from the ground, par­tic­u­larly from areas within their habi­tat that pro­duce low-ly­ing forbs. (Mc­Mas­ter and Downs, 2008; Schmidt, 2006)

• Primary Diet
• herbivore
  • folivore

• Plant Foods
• leaves
• seeds, grains, and nuts
• flowers

Pre­da­tion

Leop­ard tor­toises are well cam­ou­flaged and are dif­fi­cult to cap­ture given their thick, heavy shells. Tor­toises suf­fer the high­est pre­da­tion rates prior to hatch­ing due to pre­da­tion on eggs. Nearly 80% of hatch­lings may eaten by preda­tors such as foxes, coy­otes, and mon­gooses. Adult tor­toises are preyed upon by hu­mans and are used­ful in cre­at­ing med­i­cine, tools, and are often used as a source of food. (Stan­ford, 2010)

• Known Predators

  • humans (Homo sapiens)
  • foxes (Vulpes vulpes)
  • coyotes (Canis latrans)
  • mongooses (Herpestidae)

Ecosys­tem Roles

Leop­ard tor­toises are im­por­tant seed preda­tors and dis­perse seeds through­out their en­vi­ron­ment. They com­monly for­age on plants that are close to the ground and in­gest a large num­ber of seeds, which are re­dis­trib­uted through­out their ge­o­graphic range via defe­ca­tion. They also con­sume the seeds of berries and other fruits, which are re­dis­trib­uted after con­sump­tion as well. Leop­ard tor­toises are vul­ner­a­ble to tor­toise ticks. There is no other in­for­ma­tion avail­able re­gard­ing par­a­sites of this species. (Horak, et al., 2006; Leu­teritz and Ekbia, 2008; Schmidt, 2006)

• Ecosystem Impact
• disperses seeds

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Leop­ard tor­toises are some­times hunted for their meat and for the pet trade and are used to cre­ate tra­di­tional med­i­cine. As an in­dige­nous food source, they are cooked in their shells, how­ever, this does not occur fre­quently and does not seem to sig­nif­i­cantly af­fect pop­u­la­tion abun­dance. Ac­cord­ing to Sched­ule 2 of the West­ern Cape Na­ture Con­ser­va­tion Laws Amend­ment Act of 2000, leop­ard tor­toises are clas­si­fied as "pro­tected wildlife", which re­stricts trade of whole an­i­mals, dead or alive, and by-prod­ucts of this species. (Leu­teritz and Ekbia, 2008)

• Positive Impacts
• pet trade
• food
• source of medicine or drug

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Psam­mo­bates pardalis is re­garded as an agri­cul­tural pest (pump­kins, beans, and cow­peas) through­out its ge­o­graphic range. (Kabigu­mila, 1998)

• Negative Impacts
• crop pest

Con­ser­va­tion Sta­tus

Al­though many species of tor­toise are be­com­ing in­creas­ingly threat­ened, leop­ard tor­toises have not been eval­u­ated by the In­ter­na­tional Union for the Con­ser­va­tion of Na­ture (IUCN). They are en­demic to Africa and are cur­rently listed under CITES' Ap­pen­dix II. Al­though Tan­za­nia has the largest recorded leop­ard tor­toise pop­u­la­tion, with an es­ti­mated 5,990 in­di­vid­u­als, it also has the high­est mor­tal­ity rate of any coun­try con­tain­ing leop­ard tor­toises. Ethiopia is sec­ond, with only 500 in­di­vid­u­als. Leop­ard tor­toises are re­garded as agri­cul­tural pests through­out their ge­o­graphic range, and as a re­sult, re­tal­ia­tory killings are not un­com­mon. (Kabigu­mila, 1998; Leu­teritz and Ekbia, 2008)

• IUCN Red List

  Not Evaluated No special status

• US Federal List

  No special status

• CITES

  Appendix II

• State of Michigan List

  No special status

Con­trib­u­tors

Hillary H. Baker (au­thor), Rad­ford Uni­ver­sity, Jor­dan N. Grubb (au­thor), Rad­ford Uni­ver­sity, Chris­tine Small (ed­i­tor), Rad­ford Uni­ver­sity, John Berini (ed­i­tor), An­i­mal Di­ver­sity Web Staff.

Ref­er­ences

Auf­fen­berg, W. 1977. Dis­play Be­hav­ior in Tor­toises. Ox­ford, United King­dom: Ox­ford Uni­ver­sity Press.

Bartlett, R., P. Bartlett, M. Earle-Bridges. 2006. Tur­tles and Tor­toises. Haup­pauge, New York: Bar­ron's Ed­u­ca­tional Se­ries, Inc.

Bow­man, R. 1966. The Gala­pa­gos. Lon­don, Eng­land: Cam­bridge Uni­ver­sity Press.

Deem­ing, D., M. Fer­gu­son. 1991. Egg In­cu­ba­tion: Its Ef­fects on Em­bry­onic De­vel­op­ment in Birds and Rep­tiles. Cam­bridge, United King­dom: Cam­bridge Uni­ver­sity Press.

Dou­glas, R., M. Rall. 2006. Sea­sonal shel­ter se­lec­tion by leop­ard tor­toises Geoch­e­lone pardalis in the Franklin Na­ture Re­serve, Free State, South Africa. Ch­e­lon­ian Con­ser­va­tion & Bi­ol­ogy, 5/1: 121-129.

Edqvist, U. 2008. "Tor­toise Trust" (On-line). Ac­cessed March 15, 2011 at http://​www.​tortoisetrust.​org/​articles/​elegans.​html.

Ernst, C., R. Bar­bour. 1989. Tur­tles of the World. Wash­ing­ton, D.C., and Lon­don: Smith­son­ian In­sti­tu­tion Press.

Fer­gus, C. 2007. Tur­tles. Me­chan­ics­burg, PA: Stack­pole Books.

Fledelius, B., G. Jørgensen, H. Jensen, L. Brimer. 2005. In­flu­ence of the cal­cium con­tent of the diet of­fered to leop­ard tor­toises Geoch­e­lone pardalis. Vet­eri­nary Record: Jour­nal of the British Vet­eri­nary As­so­ci­a­tion, 156/26: 831-835.

Guzmán, A., P. Steven­son. 2008. Seed dis­per­sal, habi­tat se­lec­tion and move­ment pat­terns in the Ama­zon­ian tor­toise, Geoch­e­lone den­tic­u­lata. Am­phibia-Rep­tilia, 29/4: 463-472.

Horak, I., I. McKay, H. Heyne, A. Spick­ett. 2006. Hosts, sea­son­al­ity and ge­o­graphic dis­tri­b­u­tion of the South African tor­toise tick, Am­bly­omma mar­moreum. On­der­stepoort Jour­nal of Vet­eri­nary Re­search, 73/1: 1-3.

Kabigu­mila, J. 2000. Growth and cara­pa­cial colour vari­a­tion of the leop­ard tor­toise, Geoch­e­lone pardalis bab­cocki , in north­ern Tan­za­nia. African Jour­nal of Ecol­ogy, 38/3: 217-223.

Kabigu­mila, J. 2001. Sight­ing fre­quency and food habits of the leop­ard tor­toise, Geoch­e­lone pardalis, in north­ern Tan­za­nia. African Jour­nal of Ecol­ogy, 39/3: 276-285.

Kabigu­mila, J. 2001. Size com­po­si­tion and sex ra­tion of the leop­ard tor­toise Geoch­e­lone pardalis in north­ern Tan­za­nia. African Jour­nal of Ecol­ogy, 39/4: 393.

Kabigu­mila, J. 1998. Com­mu­nity at­ti­tudes to tor­toises Geoch­e­lone pardalis bab­cocki and their con­ser­va­tion in North­ern Tan­za­nia. African Study Mono­graphs, 19/4: 201-216.

Leu­teritz, T., R. Ravolanaivo. 2005. Re­pro­duc­tive ecol­ogy and egg pro­duc­tion of the ra­di­ated tor­toise Geoch­e­lone ra­di­ata in south­ern Mada­gas­car. African Zo­ol­ogy, 40/2: 233-242.

Leu­teritz, T., H. Ekbia. 2008. Not all roads lead to re­silience: a com­plex sys­tems ap­proach to the com­par­a­tive analy­sis of tor­toises in arid ecosys­tems. Ecol­ogy and So­ci­ety, 13/1: 1-14.

Mc­Mas­ter, M., C. Downs. 2008. Di­ges­tive pa­ra­me­ters and water turnover of the leop­ard tor­toise. Com­par­a­tive Bio­chem­istry & Phys­i­ol­ogy Part A: Mol­e­c­u­lar & In­te­gra­tive Phys­i­ol­ogy, 151/1: 114-125.

Mc­Mas­ter, M., C. Downs. 2009. Home range and daily move­ment of leop­ard tor­toises Stig­mochelys pardalis in the Nama-Ka­roo, South Africa. Jour­nal of Her­petol­ogy, 43/4: 561-569.

Mc­Mas­ter, M., C. Downs. 2006. Pop­u­la­tion struc­ture and den­sity of leop­ard tor­toises Geoch­e­lone pardalis on farm­land in the Nama-Ka­roo. Jour­nal of Her­petol­ogy, 40/4: 495-502.

Schmidt, W. 2006. Rep­tiles & Am­phib­ians of South­ern Africa. Cape Town: Struik Pub­lish­ers.

Simang, A., P. Cun­ning­ham, B. Henen. 2010. Color se­lec­tion by ju­ve­nile leop­ard tor­toises Stig­mochelys pardalis in Namibia. Jour­nal of Her­petol­ogy, 44/2: 327-331.

Spotila, J., L. Zim­mer­man, C. Binck­ley, J. Grum­bles, D. Ros­tal, A. List, E. Beyer, K. Phillips, S. Kemp. 1994. Ef­fects of in­cu­ba­tion con­di­tions on sex de­ter­mi­na­tion, hatch­ing suc­cess, and growth of hatch­ling desert tor­toises Go­pherus agas­sizii. Her­peto­log­i­cal Mono­graphs, 8/17: 103-116.

Stan­ford, C. 2010. The Last Tor­toise: A Tale of Ex­tinc­tion in Our Life­time. United States: Har­vard Uni­ver­sity Press.

Tu­berville, T., T. Nor­ton, B. Waffa, C. Hagen, T. Glenn. 2011. Mat­ing sys­tem in a go­pher tor­toise pop­u­la­tion es­tab­lished through mul­ti­ple translo­ca­tions: ap­par­ent ad­van­tage of prior res­i­dence. Bi­o­log­i­cal Con­ser­va­tion, 144/1: 175-183.

Wilkin­son, A., Hui-Minn Chan, G. Hall. 2007. Spa­tial learn­ing and mem­ory in the tor­toise Geoch­e­lone car­bonaria. Jour­nal of Com­par­a­tive Psy­chol­ogy, 121/4: 412-418.

Wim­berger, K., A. Arm­strong, C. Downs. 2009. Can re­ha­bil­i­tated leop­ard tor­toises, Stig­mochelys pardalis, be suc­cess­fully re­leased into the wild. Ch­e­lon­ian Con­ser­va­tion & Bi­ol­ogy, 8/2: 173-184.