Chelonoidis carbonariaRed-footed Tortoise

Ge­o­graphic Range

Red-footed tor­toises (Ch­e­lonoidis car­bonaria) has a broad ge­o­graphic range east of the Andes that ex­tends from east­ern Co­lum­bia through the Guianas, con­tin­u­ing south through east­ern Brazil to Rio de Janeiro, and west to Bo­livia, Paraguay and north­ern Ar­gentina. They ap­pear to be ab­sent from all but the east­ern­most Ama­zon Basin. Red-footed tor­toises are also na­tive to Panama and west­ern Columbian on main­land South Amer­ica, and the is­land of Trinidad in the Caribbean. They have re­cently been in­tro­duced to other is­lands in the Caribbean. (Ernst, et al., 1997; Legler, 1963; Spiess, 1997)

Habi­tat

Ch­e­lonoidis car­bonaria can be found in rain­forests, dry thorny forests, tem­per­ate forests, and in sa­vanna areas. It prefers heav­ily forested, humid habi­tats but avoids muddy areas due to low bur­row­ing ca­pac­ity of these habi­tats. This species is less com­mon in for­est clear­ings as­so­ci­ated with palms, man­goes, and other veg­e­ta­tion. (Ernst, et al., 1997; Legler, 1963)

Phys­i­cal De­scrip­tion

The cara­pace of red-footed tor­toises ranges from dark-brown to black­ish brown. Each scute con­tains a pale cen­ter called the are­o­lae. Outer re­gions of each scute are golden-brown, and the edges tend to be dark-brown. The plas­tron can range from pale to a dark yel­low, and the scales on the limbs and tail are en­larged and can range from yel­low to dark red­dish or­ange. Scales on the dor­sal sur­face of the head have dark mar­gins and are bright in color. Pari­etal scales are the bright­est on the head, and frontal scales are typ­i­cally more pale in com­par­i­son. Males are larger than fe­males and have a more ob­tuse pos­te­rior anal notch. Fe­males have an acute or rounded pos­te­rior anal notch and their xiphiplas­tron (the pos­te­rior-most scute) is not as heav­ily de­vel­oped when com­pared to males. In males, the pos­te­rior edge of the xiphiplas­tron tends to form a thick­ened, nearly ver­ti­cal free mar­gin, whereas in fe­males, it tends to be beveled in­ward and up­ward. Adults can reach 510 mm in cara­pace length. (Ernst, et al., 1997; Legler, 1963)

  • Sexual Dimorphism
  • male larger
  • Range length
    510 (high) mm
    20.08 (high) in

De­vel­op­ment

At re­pro­duc­tive ma­tu­rity, Ch­e­lonoidis car­bonaria ex­hibits a cen­tral nar­row­ing of the shell. In older in­di­vid­u­als, the sides of the cara­pace ex­pand, es­pe­cially in fe­males, and the ab­domi­nofemoral re­gion be­comes in­dented and slightly sad­dle-backed. The tail is longer and more dis­tinct in males when com­pared to fe­males. Males have a small open­ing be­tween the cara­pace and plas­tron, which al­lows for move­ment of their heav­ier tail. Off­spring sex is de­ter­mined via am­bi­ent tem­per­a­ture dur­ing de­vel­op­ment. (Legler, 1963)

  • Development - Life Cycle
  • temperature sex determination

Re­pro­duc­tion

Ch­e­lonoidis car­bonaria is polyg­y­nous, and males pro­duce sounds and calls as­so­ci­ated with dis­tinct gular mo­tions that are meant to at­tract po­ten­tial mates and ward off com­peti­tors. Calls con­sist of a se­ries of “clucks”, sim­i­lar to those pro­duced by chick­ens. Males com­pete for mates, and typ­i­cally move their heads in a bob­bing mo­tion prior to wrestling. He who flips his com­peti­tor on his back gains ac­cess to the fe­male and an op­por­tu­nity to mate. (Camp­bell and Evans, 1967; Spiess, 1997)

Most red-footed tor­toises be­come sex­u­ally ma­ture by 200 to 250 mm in length, which typ­i­cally oc­curs around 5 years of age. They mate through­out the year in cap­tiv­ity, but nest­ing oc­curs from June through Sep­tem­ber. Fe­males may nest sev­eral times dur­ing a sea­son, de­posit­ing 2 to 15 eggs per clutch. In­cu­ba­tion times range from 117 to 158 days. Hatch­lings have a cara­pace length rang­ing from 39 to 45 mm and weigh 22 to 30 g. (Ernst, et al., 1997; Legler, 1963; Spiess, 1997)

  • Breeding season
    Red-footed tortoise mate throughout the year but nest from June to September.
  • Range number of offspring
    2 to 15
  • Average age at sexual or reproductive maturity (female)
    5 years
  • Average age at sexual or reproductive maturity (male)
    5 years

In order to en­sure suc­cess­ful egg pro­duc­tion, fe­male Ch­e­lonoidis car­bonaria store sub­stan­tial en­ergy in the form of fat and se­quester min­er­als in their bones for the for­ma­tion of the egg-shell. Dur­ing egg de­vel­op­ment, they feed on a nu­tri­ent-rich diet in order to main­tain min­eral de­posits. Ch­e­lonoidis car­bonaria lay their eggs in flask shaped nests that are about 20 cm deep in the ground. Prior to ovipo­si­tion, fe­males be­come rest­less and search for a good nest­ing area. Once oviposited, eggs are cov­ered and there is no parental care. Hatch­lings dig their way out of the nest and are im­me­di­ately in­de­pen­dent. (Ernst, et al., 1997; Moskovits and Ki­ester, 1987; Spiess, 1997; Wilkin­son, 2010)

  • Parental Investment
  • pre-hatching/birth
    • provisioning
      • female

Lifes­pan/Longevity

There is no in­for­ma­tion avail­able re­gard­ing the av­er­age lifes­pan of Ch­e­lonoidis car­bonaria.

Be­hav­ior

Red-footed tor­toises "gaze fol­low" con­specifics, which con­sists of gaz­ing in the same di­rec­tion as con­specifics in the im­me­di­ate area. They are di­ur­nal and ap­pear to avoid mov­ing great dis­tances. Out­side of their re­pro­duc­tive be­hav­ior, lit­tle else is known of the gen­eral be­hav­ior of this species. (Wilkin­son, 2010)

Home Range

There is no in­for­ma­tion avail­able re­gard­ing the home range of red-footed tor­toises.

Com­mu­ni­ca­tion and Per­cep­tion

Ch­e­lonoidis car­bonaria pro­duces a se­ries of clucks, which sound sim­i­lar to those pro­duced by do­mes­tic chick­ens. Clucks are pri­mar­ily pro­duced by males dur­ing court­ing or mat­ing. Ju­ve­niles may also cluck or chirp and pro­duce these sounds on a reg­u­lar basis while for­ag­ing. Prior to cluck­ing, ju­ve­niles stop for­ag­ing and bob their head while pro­duc­ing the sound. Lit­tle else is known of com­mu­ni­ca­tion and per­cep­tion in this species. (Camp­bell and Evans, 1972)

Food Habits

Ch­e­lonoidis car­bonaria feeds pri­mar­ily on fruit dur­ing the wet sea­son and flow­ers dur­ing the dry sea­son. Through­out the year, they also con­sume dead and liv­ing fo­liage, soil, fungi, stems, sand, peb­bles, and car­rion. Males con­sume greater amounts of fruit com­pared to fe­males. Geo­cehlone car­bonaria oc­ca­sion­ally in­gests sand, which has been hy­poth­e­sized to act as an abra­sive agent en­hanc­ing di­ges­tion of plants ma­te­ri­als. (Moskovits and Bjorn­dal, 1990)

  • Plant Foods
  • leaves
  • wood, bark, or stems
  • fruit
  • flowers

Pre­da­tion

Other than hu­mans, there is no in­for­ma­tion avail­able con­cern­ing preda­tors spe­cific to Ch­e­lonoidis car­bonaria. (Spiess, 1997)

Ecosys­tem Roles

As fru­gi­vores, red-footed tor­toises may be im­por­tant dis­persers of seeds of trop­i­cal plants such as figs and bromeli­ads. One study found that vi­able seeds of both types of plants were found in the feces of sev­eral red-footed tor­toises in Brazil. (Strong and Fragoso, 2006)

  • Ecosystem Impact
  • disperses seeds

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Red-footed tor­toises are hunted for their meat through­out their ge­o­graphic range. (Legler, 1963)

  • Positive Impacts
  • food

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There are no known ad­verse ef­fects of G. car­bonaria on hu­mans. (Spiess, 1997)

Con­ser­va­tion Sta­tus

Ch­e­lonoidis car­bonaria has not been eval­u­ated by the In­ter­na­tional Union for Con­ser­va­tion of Na­ture (IUCN), how­ever, many is­land pop­u­la­tions are likely in se­ri­ous de­cline as habi­tat loss and over hunt­ing pose a se­ri­ous threat through­out its ge­o­graphic range. Ch­e­lonoidis car­bonaria oc­curs in nu­mer­ous wildlife re­serves and na­tional parks through­out its ge­o­graphic range. With­out cur­rent pop­u­la­tion es­ti­mates, it is dif­fi­cult to pre­dict the po­ten­tial con­ser­va­tion and man­age­ment needs for this species. The (Spiess, 1997)

Other Com­ments

Ch­e­lonoidis car­bonaria is also re­ferred to as Che­le­noidis car­bonaria and Tes­tudo car­bonaria. (IUCN, 1982)

Con­trib­u­tors

Khush­want Kaur (au­thor), In­di­ana Uni­ver­sity-Pur­due Uni­ver­sity Fort Wayne, Mark Jor­dan (ed­i­tor), In­di­ana Uni­ver­sity-Pur­due Uni­ver­sity Fort Wayne, John Berini (ed­i­tor), An­i­mal Di­ver­sity Web Staff.

Glossary

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

acoustic

uses sound to communicate

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carrion

flesh of dead animals.

detritus

particles of organic material from dead and decomposing organisms. Detritus is the result of the activity of decomposers (organisms that decompose organic material).

diurnal
  1. active during the day, 2. lasting for one day.
ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

folivore

an animal that mainly eats leaves.

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

marsh

marshes are wetland areas often dominated by grasses and reeds.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

polygynous

having more than one female as a mate at one time

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

year-round breeding

breeding takes place throughout the year

Ref­er­ences

Camp­bell, H., W. Evans. 1972. Ob­ser­va­tions on the Vocal Be­hav­ior of Ch­e­lo­ni­ans. Her­peto­log­ica, 28/3: 277-280.

Camp­bell, H., W. Evans. 1967. Sound pro­duc­tion in two species of tor­toises. Her­peto­log­ica, 23/3: 204-209.

Ernst, C., R. Al­tenburg, R. Bar­bour. 1997. "Geoch­e­lone car­bonaria" (On-line). Tur­tles of the World. Ac­cessed May 26, 2011 at http://​nlbif.​eti.​uva.​nl/​bis/​turtles.​php?​selected=beschrijving&​menuentry=soorten&​id=380.

IUCN, 1982. The IUCN Am­phibia-Rep­tilia Red Data Book Part 1: Tes­tudines, Croc­o­dylia, Rhyn­chocaphalia. Gland, Switzer­land: IUCN. Ac­cessed June 15, 2011 at http://​books.​google.​com/​books?​id=Nw8KKyu32v8C&​pg=PA61&​lpg=PA61&​dq=Chelonoidis+carbonaria+IUCN&​source=bl&​ots=hBYuKrj0n8&​sig=IDrMJWyAsL-dXO2VAoVJuRedu1I&​hl=en&​ei=hCD5TfqhCsrVgQeq46T7Cw&​sa=X&​oi=book_​result&​ct=result&​resnum=1&​ved=0CBoQ6AEwAA#​v=onepage&​q=Chelonoidis%20carbonaria&​f=false.

Legler, J. 1963. Tor­toises (Geoch­e­lone car­bonaria) in Panama: dis­tri­b­u­tion and vari­a­tion. Amer­i­can Mid­land Nat­u­ral­ist, 70/2: 490-503.

Moskovits, D., A. Ki­ester. 1987. Ac­tiv­ity lev­els and rang­ing be­hav­ior of the two Ama­zon­ian tor­toises, Geoch­e­lone car­bonaria and Geoch­e­lone den­tic­u­lata, in north-west­ern Brazil. Func­tional Ecol­ogy, 1/3: 203-214.

Moskovits, D., K. Bjorn­dal. 1990. Diet and food pref­er­ences of the tor­toises Geoch­e­lone car­bonaria and G. den­tic­u­lata in north­west­ern Brazil. Her­peto­log­ica, 46/2: 207-218.

Spiess, P. 1997. "The Red-Footed Tor­toise (Geoch­e­lone car­bonaria), a South Amer­i­can Trea­sure" (On-line). Ac­cessed March 27, 2011 at http://​www.​kingsnake.​com/​rockymountain/​RMHPages/​RMHredfoot.​htm.

Strong, J., J. Fragoso. 2006. Seed dis­per­sal by Geoch­e­lone car­bonaria and Geoch­e­lone den­tic­u­lata in north­west­ern Brazil. Biotrop­ica, 38/5: 683-686.

Wilkin­son, A. 2010. Gaze fol­low­ing in the red-footed tor­toise (Geoch­e­lone car­bonaria). An­i­mal Cog­ni­tion, 13/5: 765-769.