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The red-footed tortoise (''Chelonoidis carbonarius'') is a species of
tortoise Tortoises ( ) are reptiles of the family Testudinidae of the order Testudines (Latin for "tortoise"). Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like o ...
from northern
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. These medium-sized tortoises generally average as adults, but can reach over . They have dark-colored (nearly black), “loaf”-shaped
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
s (top shell) with a lighter patch in the middle of each
scute A scute () or scutum (Latin: ''scutum''; plural: ''scuta'' "Scutum (shield), shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of Bird anatomy#Scales, birds. The ter ...
(shell segment), and a somewhat lighter-colored
plastron The turtle shell is a shield for the ventral and dorsal parts of turtles (the Order (biology), order Testudines), completely enclosing all the turtle's vital organs and in some cases even the head. It is constructed of modified bony elements such ...
(underbelly). They also have dark limbs dotted with brightly-colored scales, from which they get their name, that range from pale yellow to vivid or dark red. Visible differences are noted between red-footed tortoises from varying eco-regions. They are closely related to the more easterly-distributed
yellow-footed tortoise The yellow-footed tortoise (''Chelonoidis denticulatus''), also known as the Brazilian giant tortoise, is a species of tortoise in the family Testudinidae and is closely related to the red-footed tortoise (''C. carbonarius''). It is found in the ...
(''C. denticulatus'') of the
Amazon Basin The Amazon basin is the part of South America drained by the Amazon River and its tributary, tributaries. The Amazon drainage basin covers an area of about , or about 35.5 percent of the South American continent. It is located in the countries ...
. Their natural habitat ranges from
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
to forest edges around the Amazon Basin. They are
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
tortoises, consuming a wide assortment of plants, grasses, flowers, fruit and (notably)
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
, as well as the occasional
earthworm An earthworm is a soil-dwelling terrestrial invertebrate that belongs to the phylum Annelida. The term is the common name for the largest members of the class (or subclass, depending on the author) Oligochaeta. In classical systems, they we ...
or other
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
. When presented with the opportunity, many tortoises will also scavenge on
carrion Carrion (), also known as a carcass, is the decaying flesh of dead animals. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures ...
. They do not
brumate Dormancy is a period in an organism's life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be cl ...
, due to their equatorial home range, but may
aestivate Aestivation ( (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a lowered me ...
in the hotter, drier months. Red-footed (and yellow-footed) tortoises are popularly kept as
pets A pet, or companion animal, is an animal kept primarily for a person's company or entertainment rather than as a working animal, livestock, or a laboratory animal. Popular pets are often considered to have attractive/ cute appearances, int ...
. Over-collection of eggs, and the capturing of mature animals, from their native habitat has caused them to be listed as “vulnerable” to extinction, despite the relative ease of
captive breeding Captive breeding, also known as captive propagation, is the process of keeping plants or animals in controlled environments, such as wildlife reserves, zoos, Botanical garden, botanic gardens, and other Conservation biology, conservation facilitie ...
and the availability of captive-bred tortoises. Additionally, many remote tribal groups living in South America still consider the tortoises a basic source of meat, and many are unaware of the species’ overall plight; many travelers and activists report seeing live red-footed tortoises for sale in markets where
bushmeat Bushmeat is meat from wildlife species that are Hunting, hunted for human consumption. Bushmeat represents a primary source of animal protein and a cash-earning commodity in poor and rural communities of humid tropical forest regions of the worl ...
and live animals are sold. Eggs, hatchlings, and even mature tortoises are food for many animal predators, including
caiman A caiman ( (also spelled cayman) from Taíno language, Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family (biology), family, the other being alligators. ...
s,
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
s and
giant otter The giant otter or giant river otter (''Pteronura brasiliensis'') is a South American carnivorous mammal. It is the longest member of the weasel family, Mustelidae, a globally successful group of predators, reaching up to . Atypical of mustel ...
s (if near water), as well as
birds of prey Birds of prey or predatory birds, also known as (although not the same as) raptors, are hypercarnivorous bird species that actively predation, hunt and feed on other vertebrates (mainly mammals, reptiles and smaller birds). In addition to speed ...
,
bushdog The bush dog (''Speothos venaticus'') is a canine found in Central and South America. In spite of its extensive range, it is very rare in most areas except in Suriname, Guyana and Peru; it was first described by Peter Wilhelm Lund from fossils ...
s,
jaguar The jaguar (''Panthera onca'') is a large felidae, cat species and the only extant taxon, living member of the genus ''Panthera'' that is native to the Americas. With a body length of up to and a weight of up to , it is the biggest cat spe ...
s,
jaguarundi The jaguarundi (''Herpailurus yagouaroundi''; or ) is a wild felidae, cat native to the Americas. Its range extends from central Argentina in the south to northern Mexico, through Central America, Central and South America east of the Andes. T ...
s,
margay The margay (''Leopardus wiedii'') is a small wild cat native to Mexico, Central and South America. A solitary and nocturnal felid, it lives mainly in primary evergreen and deciduous forest. Until the 1990s, margays were hunted for the wildl ...
s, certain
monkeys Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
,
ocelot The ocelot (''Leopardus pardalis'') is a medium-sized spotted Felidae, wild cat that reaches at the shoulders and weighs between on average. It is native to the southwestern United States, Mexico, Central America, Central and South America, ...
s, pumas and
tayra The tayra (''Eira barbara'') is an omnivorous animal from the mustelid family, native to the Americas. It is the only species in the genus ''Eira''. Tayras are also known as the ''tolomuco'' or ''perico ligero'' in Central America, ''motete'' ...
s. Population density ranges from locally common to very scarce due, in-part, to
habitat destruction Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
and the aforementioned over-collection for food and the pet trade. The species name has often been misspelled as ''carbonaria'', an error introduced in the 1980s when ''Chelonoidis'' was elevated to genus and mistakenly treated as feminine, an error recognized and fixed in 2017.


Names

Red-footed tortoises have many common names: red-leg, red-legged, or red-foot tortoise (often without the hyphen) and the savanna tortoise, as well as local names, such as ''carumbe'' or ''karumbe'', which means 'slow moving' (Brazil,
Paraguay Paraguay, officially the Republic of Paraguay, is a landlocked country in South America. It is bordered by Argentina to the Argentina–Paraguay border, south and southwest, Brazil to the Brazil–Paraguay border, east and northeast, and Boli ...
), ''wayapopi'' or ''morrocoy'' (
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
,
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...
), and variations of ''jabuti'' such as ''japuta'' and ''jabuti-piranga'' (Brazil,
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
). p. 27-29. In Portuguese, '' jabuti'' (''jaboti'') refers to both the red-footed tortoise and the
yellow-footed tortoise The yellow-footed tortoise (''Chelonoidis denticulatus''), also known as the Brazilian giant tortoise, is a species of tortoise in the family Testudinidae and is closely related to the red-footed tortoise (''C. carbonarius''). It is found in the ...
. It is also called ''kati'' in NatúPompeu Sobrinho, Thomaz. 1958. ''Línguas Tapuias desconhecidas do Nordeste: Alguns vocabulários inéditos''. Boletim de Antropologia (Fortaleza-Ceará) 2. 3-19. and ''sambó'' in Kiriri.


Taxonomy

Red footed tortoises were originally assigned to the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
'' Testudo'' (named by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in 1758) for a short time, but it soon became the term for turtles with high-domed shells, elephantine legs, and completely terrestrial habits—the tortoises. In 1835,
Leopold Fitzinger Leopold Joseph Franz Johann Fitzinger (13 April 1802 – 20 September 1884) was an Austrian zoologist. Fitzinger was born in Vienna and studied botany at the University of Vienna under Nikolaus Joseph von Jacquin. He worked at the Vienna Naturhis ...
used ''
Geochelone ''Geochelone'', from Ancient Greek γῆ (''gê''), meaning "earth", and χελώνη (''khelṓnē''), meaning "turtle", is a genus of tortoises. ''Geochelone'' tortoises, which are also known as typical tortoises or terrestrial turtles, can be ...
'' to differentiate some non-Mediterranean tortoises, apparently based on size and lack of specific identifying characteristics such as the hinged shell in the African hingeback tortoises. He used the term ''Chelonoidis'' as a subgenus for the species from South America. Few people used these terms until they were resurrected by Hewitt in 1933 and Loveridge and Williams in 1957. In 1982, Roger Bour and Charles Crumly each separated ''Geochelone'' into different genera based on anatomic differences, especially in the skulls. That resulted in the formation or restoration of several genera: ''
Aldabrachelys ''Aldabrachelys'' is genus of giant tortoises, including the Aldabra giant tortoise (''Aldabrachelys gigantea'') native to the Seychelles, as well as two extinct species, '' Aldabrachelys abrupta'' and '' Aldabrachelys grandidieri'' known from M ...
'', ''
Astrochelys ''Astrochelys'' is a genus of tortoises in the family Testudinidae. The two species are both found in Madagascar, and both classified as critically endangered on the IUCN Red List. Species A single extinct Extinction is the termination ...
'', ''
Cylindraspis ''Cylindraspis'' is a genus of recently extinct giant tortoises. All of its species lived in the Mascarene Islands (Mauritius, Rodrigues and Réunion) in the Indian Ocean and all are now extinct due to hunting and introduction of non-native pred ...
'', ''
Indotestudo ''Indotestudo'' is a genus of tortoises in the Family (biology), family Testudinidae. The genus is native to South Asia, South and Southeast Asia. The three species in the genus are all Threatened species, threatened. Species ''Indotestudo'' co ...
'', ''
Manouria ''Manouria'' is a genus of tortoises in the family Testudinidae. The genus was erected by John Edward Gray in 1854. Transitional adaptations ''Manouria'' is either the most basal or second most basal member of the family Testudinidae (its phyl ...
'', and ''Chelonoidis''. ''Chelonoidis'' was distinguished from other ''Geochelone'' by their South American location, as well as the absence of the nuchal scute (the marginal centered over the neck) and the presence of a large, undivided supracaudal (the scute or scutes directly over the tail), as well as differences in the skull. Many of these generic names are still debated; for example, no specific definition of ''Geochelone'' is given, and ''Chelonoidis'' is primarily used for geography rather than unique anatomic characteristics. , p. 204. The species name ''carbonarius'' means 'coal-like' referring to a dark coal with glowing patches. It was originally identified by
Johann Baptist von Spix Johann Baptist Ritter von Spix (9 February 1781 – 13 March 1826) was a German natural history, biologist. From his expedition to Brazil, he brought to Germany a large variety of specimens of plants, insects, mammals, birds, amphibians and fish. ...
in 1824. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
was kept in the Zoologischen Sammlung des Bayerischen Staates in
Munich Munich is the capital and most populous city of Bavaria, Germany. As of 30 November 2024, its population was 1,604,384, making it the third-largest city in Germany after Berlin and Hamburg. Munich is the largest city in Germany that is no ...
, Germany, but was lost.
Paulo Vanzolini Paulo Emilio Vanzolini (; April 25, 1924 – April 28, 2013) was a Brazilian scientist and music composer. He was best known for his samba compositions, including the famous ''"Ronda", "Volta por Cima"'', and ''"Boca da Noite"'', and for his sc ...
believes it may have come from near the city of
Manaus Manaus () is the List of capitals of subdivisions of Brazil, capital and largest city of the States of Brazil, Brazilian state of Amazonas (Brazilian state), Amazonas. It is the List of largest cities in Brazil, seventh-largest city in Brazil, w ...
, Brazil, on the Rio Negro. No subspecies of red-footed tortoise are recognized, although many believe the species has five or more variants that may be subspecies or even separate species.


Description

Red-footed tortoises show sex, regional, and individual variations in color, shell shape, and minor anatomical characteristics. Adult carapaces are generally an elongated oval with sides that are nearly parallel, although the sides of males may curve inwards. They are fairly highly domed and smooth with a rather flat back (although the scutes may be raised or 'pyramided' in some individuals, especially captive specimens). Often, a high point over the hips is seen, with a small sloped section over the neck. The vertebral and costal scutes (the scutes along the center and sides of the carapace) are black or dark brown with a pale yellow
areole In botany, areoles are small light- to dark-colored bumps on cactus, cacti out of which grow clusters of Thorns, spines, and prickles, spines. Areoles are important diagnostic features of cactus, cacti, and identify them as a family distinct fr ...
in the center. The marginals (scutes along the edge of the carapace) 'tuck under' along the sides and flare slightly over the limbs. They are dark with the pale aureole along the middle of the lower edge. The nuchal scute (the marginal over the neck) is absent, and the marginals over the tail are joined as one large supracaudal. Growth rings are clearly evident in most individuals, but become worn smooth with age. The
plastron The turtle shell is a shield for the ventral and dorsal parts of turtles (the Order (biology), order Testudines), completely enclosing all the turtle's vital organs and in some cases even the head. It is constructed of modified bony elements such ...
(bottom shell) is large and thick along the edges. The gulars (frontmost pair of plastron scutes) do not protrude much past the front of the carapace. The plastron of a male is deeply indented, and the anal scutes (rearmost pair of plastron scutes) may be used to sex the animal while the color pattern varies by region. The head is relatively small with a squared-off profile and flat on top, longer than it is wide. The eye is large with a brown, almost black
iris Iris most often refers to: *Iris (anatomy), part of the eye * Iris (color), an ambiguous color term *Iris (mythology), a Greek goddess * ''Iris'' (plant), a genus of flowering plants * Iris (given name), a feminine given name, and a list of peopl ...
, and rarely any
sclera The sclera, also known as the white of the eye or, in older literature, as the tunica albuginea oculi, is the opaque, fibrous, protective outer layer of the eye containing mainly collagen and some crucial elastic fiber. In the development of t ...
visible around it. The upper jaw is slightly hooked, and the upper jaw is notched in the front middle. About 15 to 20 'teeth' or fine grooves occur on each side of each jaw. A nearly circular tympanum is located behind and below the eye and is covered with a dark scale. The scales of the head are generally smallish and irregular, becoming small and pebbly on the neck. Many of the scales are colored pale yellow to brick red, especially those on the top of the head, above the tympanum, around the nostrils, on the lower jaw, and on the sides of the neck. Males are usually slightly more colorful than females, and colors vary by region. The limbs are generally cylindrical with five claws on the fore limbs and four on the hind, but no visible toes. The fore limbs are slightly flattened and the front surface is covered with large scales, mostly with the same color as the head. They are not as large or protrusive as they are in more primitive species such as the
African spurred tortoise The African spurred tortoise (''Centrochelys sulcata''), also called the sulcata tortoise, is an endangered species of tortoise inhabiting the southern edge of the Sahara, Sahara Desert, the Sahel, in Africa. It is the largest mainland species o ...
(''Geochelone sulcata''). The tail is muscular, varies in length and overall shape by sex, and lacks any sort of claw on the tip. Average adult sizes vary by region and sex, and 'giants' are often encountered. Red-footed tortoises average with males slightly larger overall. Tortoises up to are fairly common and over are occasionally discovered. The largest known specimen, from Paraguay, was long, and weighed over . It is unknown if the 'giants' represent diet availability, genetic issues, longevity, or other possibilities. Hatchling and young red-footed tortoises have much rounder and flatter carapaces that start off as mostly pale yellow to brown. New growth adds dark rings around the pale center of each scute. The marginals of very young tortoises are serrated, especially over the hind limbs. This probably aids in both camouflage against the leaf litter and in making the small animals harder to eat. Young tortoises are generally more colorful overall. Males are slightly larger and more colorful overall. The carapace of a male from north of the Amazon basin shows a 'wasp waist', or constrictions along the sides. The male's plastron is deeply indented to help with positioning during mating. The male's tail is long and muscular, generally carried along a side while the female's tail is short and conical. The anal scutes vary to allow the male's tail more mobility and allows more protection for the female's hind end. The gap between the points of the anal scales and the marginals is wider and the anal scutes form a broader angle- almost a straight line across- in males to allow the tail to move laterally. The angle is more closed (to about a 90° angle) and the points are closer to the marginals in females.


Distribution and habitat

Red-footed tortoises range from southeastern Panama to Venezuela,
Guyana Guyana, officially the Co-operative Republic of Guyana, is a country on the northern coast of South America, part of the historic British West Indies. entry "Guyana" Georgetown, Guyana, Georgetown is the capital of Guyana and is also the co ...
,
Suriname Suriname, officially the Republic of Suriname, is a country in northern South America, also considered as part of the Caribbean and the West Indies. It is a developing country with a Human Development Index, high level of human development; i ...
, and
Guiana The Guianas, also spelled Guyanas or Guayanas, are a geographical region in north-eastern South America. Strictly, the term refers to the three Guianas: Guyana, Suriname, and French Guiana, formerly British, Dutch, and French Guiana respectiv ...
in the north; south along the
Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...
to the west in Colombia,
Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...
,
Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...
, and
Bolivia Bolivia, officially the Plurinational State of Bolivia, is a landlocked country located in central South America. The country features diverse geography, including vast Amazonian plains, tropical lowlands, mountains, the Gran Chaco Province, w ...
; east to Brazil, and along the southern range in Bolivia, Paraguay, and possibly northern
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. In Panama they live from the
Darien Gap Darien may refer to: Places Central America * Darién Gap, break in the Pan-American Highway between Colombia and Panama * Darién National Park * Darién Province * Gulf of Darién * "... a peak in Darien", phrase in Keats's poem " On First ...
to the canal. They are not evenly distributed within their range. For example, they are not often found in central Brazil or in heavily forested areas in general, and have only documented in Peru since 1985. Accurate range information is complicated by the sheer size of the range, political and geographic barriers, and confusion about where many specimens were collected. They are also found on several
Caribbean Islands Most of the Caribbean countries are islands in the Caribbean Sea, with only a few in inland lakes. The largest islands include Cuba, Hispaniola, Jamaica and Puerto Rico. Some of the smaller islands are referred to as a ''rock'' or ''reef.'' ''I ...
, although it is not always clear if they are native or brought by humans. Many of the colonies seem to have been established in the 17th century as food supplies or as pets. They are found on the former
Netherlands Antilles The Netherlands Antilles (, ; ), also known as the Dutch Antilles, was a constituent Caribbean country of the Kingdom of the Netherlands consisting of the islands of Saba (island), Saba, Sint Eustatius, and Sint Maarten in the Lesser Antilles, ...
, Trinidad, Tobago, Grenada, Barbados, St. Vincent, the Grenadines, Santa Lucia, Martinique, Dominica, Guadalupe, the Leeward Islands, the Virgin Islands, Barbuda living in sink-holes forests and Puerto Rico. The preferred habitat of the red-footed tortoise varies somewhat by region, but generally includes fairly consistent seasonal temperatures near that rarely get lower than or over , generally with high humidity and plenty of rainfall, although some of the areas can get quite dry. Most of the range experiences cooler wet seasons (April to August) and warmer dry seasons (September to March), but some parts of the southern range have occasional cold snaps. Red-footed tortoises are often found in or near transitional areas between forest and savannah, such as forest clearings, wood edges, or along waterways.


Regional variations

Various authors have divided the red-footed tortoise into different groups by anatomy and geography.
Peter Pritchard Peter Charles Howard Pritchard (June 26, 1943 – February 25, 2020) was a leading turtle zoologist. Pritchard was educated at Oxford University and the University of Florida, where he received a Ph.D. and specialized in Zoology. He was most com ...
recognized seven types, but DNA research has identified five genotypes. The most obvious differences are between the groups found north or south of the Amazon basin. The 'northern' variants all look very much like the holotype and are distinguished primarily by shell, head, and limb coloration. The variants south of the Amazon are generally both larger and smaller than the holotype, have a very different plastral pattern, and have an enlarged scale or 'spur' on the inside of the fore limb elbow.


Northeastern variant

This is the holotype of the species. Head and limb colors are generally light orange to red. Plastrons are mostly pale yellow. They range in the
Guiana Shield The Guiana Shield (; ; ; ) is one of the three cratons of the South American Plate. It is a 1.7 billion-year-old Precambrian geological formation in northeast South America that forms a portion of the northern coast. The higher elevations on ...
- Venezuela, Guyana, Suriname, Guiana, and northern Brazil.


Northwestern variant

They are similar to the northeastern variant, but their carapace base color is grey, dark brown, or coffee rather than black. Their pale plastrons have central dark areas resembling an exclamation point. Their heads and limbs are generally pale yellow to orange. The average size is slightly smaller than usual- 30–35 cm. They are found in southeast Panama and Colombia.


Northern variant

These also are similar to the northeastern variant, with head and limb colors generally pale yellow to light orange, rarely red, and their heads and limbs are often slightly different colors. The average size is slightly smaller than usual- 30–35 cm. They are found in Colombia, Ecuador, and Peru. pp. 7-10.


Southern variant

The southern variants' carapaces are often not quite black to dark brown, sometimes with light grey or whitish between the scutes. Their plastrons are mostly dark in a symmetrical mottled pattern. Size tends to be larger on average than northeastern variants, with the largest individuals found in this area. Fore limbs feature a slightly enlarged scale on the side of the 'elbow'. Adult males do not have the constricted waist, and females average a bit larger than the males. They are seen in the
Gran Chaco The Gran Chaco or simply Chaco is a sparsely populated, hot and semiarid lowland tropical dry broadleaf forest natural region of the Río de la Plata basin, divided among eastern Bolivia, western Paraguay, northern Argentina, and a portion o ...
- Bolivia, Paraguay, and northern Argentina.


Eastern variant

Carapaces of the eastern variants are often light grey or whitish between the scutes. Their plastrons are mostly dark in a symmetrical mottled pattern. Size tends to be smaller on average than northeastern variants, also reaching sexual maturity at a smaller size. Fore limbs feature a slightly enlarged scale on the side of the 'elbow'. Their heads and limbs are either yellowish or red, ranging to brilliant cherry-red. They are located in east to southeast Brazil. The red-headed type of this variant is often called a 'cherry-head' in the pet trade.


Population features

Little information is available about population density or sex ratios. Many specimens are recorded from near research stations and cities, but that is almost certainly more due to the ease of finding them there than higher localized populations. They are considered one of the most common turtle species in many localities. When a dam was being built in the Edo region of Venezuela, several hundreds of the red- and yellow-footed tortoises were captured for relocation. Large numbers are also found in markets, confiscated at airports, etc. However, very few records exist from Peru, Ecuador, Argentina, and central Brazil. In the Guiana Shield region, islands of higher populations and stretches where few have been located are seen. Counting tortoises that are often well-hidden, dug in, deep in burrows, etc. is difficult. Trained dogs discover many that otherwise might not be found. One study showed a nearly balanced mix of 1.32 males per female, and 1.05 red-footed tortoises per
hectare The hectare (; SI symbol: ha) is a non-SI metric unit of area equal to a square with 100-metre sides (1 hm2), that is, square metres (), and is primarily used in the measurement of land. There are 100 hectares in one square kilometre. ...
(2.4 acres) on the island, which may be an underestimate due to the difficulty in finding the tortoises. p. 41-42


Evolutionary history and fossil record

The genus ''Chelonoidis'' has two main subcategories based on appearance and habitat- the ''C. carbonarius'' and the ''C. chilensis'' groups. The ''C. carbonarius'' group has the closely related red- and yellow-footed tortoises that clearly share a common ancestor. The ''C. chilensis'' group features the
Chaco tortoise The Chaco tortoise (''Chelonoidis chilensis''), also known commonly as the Argentine tortoise, the Patagonian tortoise, or the southern wood tortoise, is a species of tortoise in the family Testudinidae. The species is endemic to South America. ...
(''C. chilensis'') and
Galápagos tortoise The Galápagos tortoise or Galápagos giant tortoise (''Chelonoidis niger'') is a very large species of tortoise in the genus ''Chelonoidis'' (which also contains three smaller species from mainland South America). The species comprises 15 subsp ...
(''C. niger''), which share similar habitats and a basic appearance, but otherwise do not seem to be closely related. The relationship between the groups is unclear. Several theories are offered to explain the relatively small number of tortoise species in South America and the relationship between them, but the fossil record is not very complete. One long-held theory is that they came from Asia using the land bridge, then spread down through North America and shared ancestors with the gopher tortoises (''
Gopherus ''Gopherus'' is a genus of fossorial tortoises commonly referred to as gopher tortoises. The gopher tortoise is grouped with land tortoises that originated 60 million years ago, in North America. A genetic study has shown that their closest relat ...
'' species). Another holds ''Geochelone'' ancestors floated over from central Africa, taking advantage of their ability to float, resist salt water, and go without food for extended periods. DNA studies suggest that the ''carbonarius'' group may be related to the African hingeback tortoises (''Kinixys'' species). This suggests that they might have come from
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
before it separated into Africa and South America some 130 million years ago. One ancestral form from about 5mya, ''Chelonoidis hesterna'' (Auffenberg 1971), is thought to have lived in wet forests and split into two species in the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
with the yellow-footed tortoises remaining in the deep forest and the red-footeds colonizing the edges of the forests and the emerging savannahs. As the climate and topography changed, groups of red-footed tortoises became physically separated and genetically isolated.


Ecology and behavior

The red-footed tortoise's climate in the northern part of the range changes little day to day and rarely gets too hot for them, so the tortoises do not need to practice any form of
dormancy Dormancy is a period in an organism's Biological life cycle, life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolism, metabolic activity and therefore helps an organism to conserv ...
and can often forage all day long. The tortoises in Moskovitz's study area were most active after 3:00pm while many species from warmer climes would be most active in the morning and evening. Tortoises from the southern parts of the range experience much hotter, colder, and drier conditions than most of the range and aestivate when food becomes scarce. They may aestivate or brumate when the temperatures are low enough. Most species of tortoise spend much of their day inactive, and red-footed tortoises generally spend over 50% of the daylight hours at rest. They may rest for even longer after a large meal, with five- to ten-day stretches being common. One large specimen seems to have stayed in the same position for over a month. Resting tortoises barely move, allowing leaf litter to accumulate on them, and
termites Termites are a group of detritophagous eusocial cockroaches which consume a variety of decaying plant material, generally in the form of wood, leaf litter, and soil humus. They are distinguished by their moniliform antennae and the sof ...
have built tunnels on the carapaces of resting red-footed tortoises. They seek shelter in places that offer
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
and protection from predators. Treefalls are a favored site, as are debris piles, burrows (especially those of the
armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...
(''Dasypodidae'' family) and
agouti The agouti (, ) or common agouti is any of several rodent species of the genus ''Dasyprocta''. They are native to Central America, northern and central South America, and the southern Lesser Antilles. Some species have also been introduced else ...
(''Dasyprocta'' species)), hollow logs, holes, and heavy vegetation cover. They generally try to find tight-fitting resting places, and will occasionally 'wedge' themselves between roots and trunks but remain otherwise exposed. Burrows and holes are often flooded and the tortoises will rest in the water and mud with just their nostrils and eyes exposed. In warmer weather, they press up against moister, cooler surfaces in shelter areas. The tortoises show personal preferences with many individuals always seeking out a specific type of shelter. Shelters are often communal with as many tortoises as can fit in the space. Good shelters are so important, and visibility is so poor that the tortoises will leave scent trails that they or others can follow. Some shelters are so heavily used by tortoises that clear trails in the dirt lead to them, though individual tortoises may not return to a given shelter regularly. Besides communal shelters and scent marking, red-footed tortoises also show other signs of semisocial behavior such as lack of aggression at feeding sites, not protecting territory, and group feeding at fruit falls and carrion (although it is not unusual for one tortoise to block access or even try to make off with some of the food). Red-footed tortoises often follow each other, usually a smaller one following a larger and quite often males following males, but all combinations are seen. They have also been observed following apparent scent trails laid by an individual a day or two earlier.


Predators

Hatchling and young tortoises are at high risk of predation. Tegu lizards (''
Tupinambis ''Tupinambis'' is a lizard genus which belongs to the Family (biology), family Teiidae and contains eight described species. These large lizards are commonly referred to as tegus (''teiús'' in Portuguese language, Portuguese). ''Tupinambis meria ...
'' species), ring-tailed
coatis Coatis (from Tupí), also known as coatimundis (), are members of the family Procyonidae in the genera '' Nasua'' and '' Nasuella'' (comprising the subtribe Nasuina). They are diurnal mammals native to South America, Central America, Mexico, ...
(''Nasua nasua''), and introduced rats and mongooses attack nests and eggs. Many predators take the young tortoises, including large lizards, snakes, crocodilians, and even large turtles; predatory birds such as the curassow (''
Crax ''Crax'' is a genus of curassows in the order Galliformes, a clade of large, heavy-bodied, ground-feeding birds. They are known from tropical South America with one species, the great curassow, ranging northwards through Central America as far a ...
'' species), guan (''
Penelope Penelope ( ; Ancient Greek: Πηνελόπεια, ''Pēnelópeia'', or , ''Pēnelópē'') is a character in Homer's ''Odyssey.'' She was the queen of Homer's Ithaca, Ithaca and was the daughter of Spartan king Icarius (Spartan), Icarius and ...
'' species),
rails Rail or rails may refer to: Rail transport *Rail transport and related matters * Railway track or railway lines, the running surface of a railway Arts and media Film * ''Rails'' (film), a 1929 Italian film by Mario Camerini * ''Rail'' (1967 fi ...
,
cuckoo Cuckoos are birds in the Cuculidae ( ) family, the sole taxon in the order Cuculiformes ( ). The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals, and anis. The coucals and anis are somet ...
s, and
falcon Falcons () are birds of prey in the genus ''Falco'', which includes about 40 species. Some small species of falcons with long, narrow wings are called hobbies, and some that hover while hunting are called kestrels. Falcons are widely distrib ...
s; and mammals such as cats (
Felidae Felidae ( ) is the Family (biology), family of mammals in the Order (biology), order Carnivora colloquially referred to as cats. A member of this family is also called a felid ( ). The 41 extant taxon, extant Felidae species exhibit the gre ...
species),
opossum Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 126 species in 18 genera. Opossums originated in South America and entered North A ...
s,
fox Foxes are small-to-medium-sized omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull; upright, triangular ears; a pointed, slightly upturned snout; and a long, bushy tail ("brush"). Twelve species ...
es,
peccaries Peccaries (also javelinas or skunk pigs) are pig-like ungulates of the family Tayassuidae (New World pigs). They are found throughout Central and South America, Trinidad in the Caribbean, and in the southwestern area of North America. Peccari ...
, and feral dogs. Other than humans, the main predators of the adult tortoises are
jaguars The jaguar (''Panthera onca'') is a large cat species and the only living member of the genus ''Panthera'' that is native to the Americas. With a body length of up to and a weight of up to , it is the biggest cat species in the Americas an ...
(''Panthera onca''). Red-footed and yellow-footed tortoises seem to be a significant food source for jaguars in some parts of their respective ranges, such as
Manú National Park Manu National Park () is a national park and biosphere reserve located in the regions of Madre de Dios and Cusco in Peru. It protects a diverse number of ecosystems including lowland rainforests, cloud forests and Andean grasslands. History M ...
in Peru. Jaguars bite at the carapace and work at cracking or prying it apart to extract the soft tissues. Many tortoises show toothmarks from attacks that they survived, often on the hind end when they were otherwise protected in a burrow or shelter.


Diet

Discussing the diet of the red-footed tortoise is difficult due to the wide variety of foods it eats, the variety and seasonal availability of plants available across the extensive range, interpreting what field reports are trying to describe, and because studying fecal pellets gives very different results from what the animals are observed eating. Forest-dwelling tortoises in the ''Chelonoidis'', ''Indotestudo'', ''Manouria'', and ''Kinixys'' genera are omnivores with upper and lower intestines about the same length, while herbivorous genera such as ''Gopherus'' and ''Testudo'' have longer large intestines to digest fibrous grasses. Most omnivorous tortoises have no other specialized digestive structures, reflecting their generalized, flexible diet. The bulk of the diet is some sort of fruit or seed pod. Common fruits come from cacti (''
Opuntia ''Opuntia'', commonly called the prickly pear cactus, is a genus of flowering plants in the cactus family Cactaceae, many known for their flavorful fruit and showy flowers. Cacti are native to the Americas, and are well adapted to arid clima ...
''), figs (''
Ficus ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family (biology), family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few spe ...
''), pehen ('' Acacia aroma''), ''
Spondias ''Spondias'' is a genus of flowering plants in the cashew family, Anacardiaceae. The genus consists of 17 described species, 7 of which are native to the Neotropics and about 10 are native to tropical Asia. They are commonly named hog plums, Sp ...
'', ''
Annona ''Annona'' or Anona (from Taíno ''annon'') is a genus of flowering plants in the pawpaw/ sugar apple family, Annonaceae. It is the second largest genus in the family after '' Guatteria'', containing approximately 166
'', ''
Philodendron ''Philodendron'' is a large genus of flowering plants in the family Araceae. , the Plants of the World Online accepted 621 species; other sources accept different numbers. Regardless of number of species, the genus is the second-largest member o ...
'',
bromeliad The Bromeliaceae (the bromeliads) are a Family (biology), family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the Tropics, tropical Americas, with several species found in the American subtropics and on ...
s, and more. Up to five different kinds of fruits are often found in fecal pellets. The entire fruit is eaten, and the seeds are passed and can germinate, giving red- and yellow-footed tortoises a significant role in seed dispersal. Red-footed tortoises have been observed at the base of fruit trees, apparently waiting for fruit to fall. The rest of the diet includes grasses, leaves, flowers, roots, and shoots from a wide assortment of plants as well as
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
, live
invertebrates Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...
(such as ants, termites, beetles, butterflies, snails, and worms), carrion, and feces (especially from foxes). Tortoises are often found at carcasses feeding until gorged, and occasionally eat small live animals such as snakes and rodents. Pebbles and sand are also often found in fecal pellets. The diet changes based on season and availability. In the wet season it may be roughly 70% fruit, 25% fresh leaves and shoots, and the rest being fungi and animal food. The dry season sees 40% fruits, 23% flowers, 16% fresh leaves and shoots, and the rest fungi, moss, and animal food. Omnivorous tortoises seek out foods high in calcium even if other foods are more readily available, and even eat mineral-rich soil if they cannot get adequate calcium in the diet.


Movement

Red-foot tortoises forage over areas ranging from usually making a 'spiderweb' pattern centered on a good hide or a recent fruit fall. They generally move methodically around 5 to 20 m/hr (5.5-22 yd/hr), but can raise up on their long legs and move up to 100 m/hr (109 yd/hr) when they want. While they generally forage in zig-zag or looping patterns, they sometimes move or more in fairly straight lines, often at a rapid pace. They show a marked preference for moving under medium to dense vegetation cover.


Lifecycle

All turtles and tortoises start as eggs. Red-footed tortoise eggs are roughly spherical and average around and weigh with two to seven eggs in a clutch, although the same females may lay multiple clutches near each other. The incubation period is 105–202 days, with 150 being typical. Hatchlings use an egg tooth to open the egg. They stay in the egg or nest for several days. Hatchling shells are bent almost in half in the egg and take some time to straighten out. The hatchling's carapace is flat, somewhat creased from being folded in the egg, and has serrated sides. Little is known of the daily activities or diet of hatchling wild tortoises. Subadult tortoises grow quickly to reach breeding sizes- roughly , depending on the average adult size of the regional variant. The peak time for courtship and reproduction is the early wet season in April and May, although it can happen at any time. Courtship noises and possibly scent cues seem to attract other tortoises to 'courting sites' under fruiting trees such as ''
Genipa ''Genipa'' is a genus of trees in the family Rubiaceae. This genus is native to the American tropical forests. Description Tall trees, without any spines, prickles or thorns; with large opposite leaves of almost leathery texture, smooth or hai ...
''. When two tortoises meet about a meter apart, they engage in some specific behaviors to identify the other. The first trigger is head and limb color; the bright red, orange, yellow, or white colors on the dark skin identify the other animal as the proper species. Next, the larger tortoise makes jerky side-to-side head movements for two to four seconds. If both tortoises are males, one will either withdraw and retreat, or they may try to ram each other, trying to get their gular scutes under the other one, then pushing them several meters away as quickly as possible. The defeated tortoise is sometimes flipped onto his back in the process. The defeated tortoise will leave the area afterwards. Neither head bobbing nor ritual combat have been observed in tortoises south of the Amazon Basin, possibly due to the lack of yellow-footed tortoises in the area. Males mounting other males, and even females mounting either sex have been witnessed and are thought to show dominance. If the other tortoise is a female, she will move away and the male will follow, touching her carapace and occasionally sniffing at her cloaca. If the female stops, the male may either wait for her to resume moving or leave. Males make loud 'clucking' sounds during the chase. After trailing, the male mounts the female, his feet planted on the costals of her carapace, rams his anal scutes against her supracaudal, and makes a loud raspy 'bark'. If the female resumes walking, he may fall off and resume trailing. Females sometimes seem to intentionally use low limbs to knock males off. A receptive female extends her hind legs and lifts her plastron as the male plants himself on his own extended hind legs as he works to align their cloacae for insertion. The tail, scutes, an
penis
of the tortoise are designed to work around the awkwardness of the shell. The male often leans his head over her head and holds his jaws wide open making calls that get louder. He may bite her, as well, sometimes quite aggressively. The shells can make loud clacking noises during the forceful thrusts. The female walks away after copulation, sometimes knocking the male off her. The female begins nesting five to six weeks after mating. Digging the nests is often difficult in hard soil. The female may urinate to soften the soil before using her hind legs to dig a chamber about in about three and a half hours. Inexperienced females often dig several partial nests, and even experienced females may abandon a nest they are working on and start another. When the nest is ready, she lowers her tail as deep into the nest as she can and deposits an egg every 30 to 120 seconds. She recovers the nest and tamps the soil down. Females get better at digging, covering, and camouflaging nests over time. After it is covered and hidden, she often gets a long drink of water, then finds a shelter and rest. Very rarely, a red-footed tortoise lays eggs on the surface, or within a patch of cacti. As with other tortoises, red-footed tortoises can reproduce most of their lives, although the number of eggs laid and the ratio of successful hatchlings improves as the tortoise matures, then drops off again as the tortoise ages. Because of the difficulty in determining the age of a wild tortoise, few data on longevity exist, although many live for 30 years or more in captivity.


Conservation and relations with humans

The red-foot tortoise is considered a
vulnerable species A vulnerable species is a species which has been Conservation status, categorized by the International Union for Conservation of Nature as being threatened species, threatened with extinction unless the circumstances that are threatened species, ...
and is listed in
CITES CITES (shorter acronym for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of inte ...
Appendix II, restricting international trade- although this does not offer protection within a country and smuggling still occurs in large numbers. Conservation parks and refuges, captive-breeding farms in natural conditions, and increased captive breeding in other countries has helped, but they are still exported in large numbers (35,565 from 2000 to 2005), mostly as pets and food. The recorded exports do not include smuggling or other losses, which some estimate to be well over twice that number. They are considered especially at risk in Argentina and Colombia, and are considered more at risk than both the
yellow-footed tortoise The yellow-footed tortoise (''Chelonoidis denticulatus''), also known as the Brazilian giant tortoise, is a species of tortoise in the family Testudinidae and is closely related to the red-footed tortoise (''C. carbonarius''). It is found in the ...
, and
Chaco tortoise The Chaco tortoise (''Chelonoidis chilensis''), also known commonly as the Argentine tortoise, the Patagonian tortoise, or the southern wood tortoise, is a species of tortoise in the family Testudinidae. The species is endemic to South America. ...
. They are widely used as food throughout their ranges, especially where other meats are limited. Their ability to go a long time without eating makes it easy to catch and keeps them fresh for extended periods. The Roman
Catholic Church The Catholic Church (), also known as the Roman Catholic Church, is the List of Christian denominations by number of members, largest Christian church, with 1.27 to 1.41 billion baptized Catholics Catholic Church by country, worldwid ...
allows tortoises to be eaten on fasting days when most meats are forbidden, such as
Lent Lent (, 'Fortieth') is the solemn Christianity, Christian religious moveable feast#Lent, observance in the liturgical year in preparation for Easter. It echoes the 40 days Jesus spent fasting in the desert and enduring Temptation of Christ, t ...
. Tortoise pie (''pastel de morrocoy'', served in a tortoise shell) is a favorite food for those times, and large numbers of tortoises are exported just for that purpose. Even people living in countries with an abundance of available livestock enjoy wild game, such as tortoises, when possible. Hunting for food is so extensive that Colombia and some other countries import tortoises from neighbors.
Habitat destruction Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
is another significant threat to the red-footed tortoise, and so many other species. They are also widely collected as local pets and their shells are sold as souvenirs.


Captive care

Red-footed tortoises are popular pet tortoises around the world. They are relatively inexpensive, are a manageable size, have interesting personalities, and are colorful. Pet tortoises should be purchased as captive-bred from a reliable breeder when possible to both help protect wild populations and to avoid internal parasites. Any reptile can carry ''
Salmonella ''Salmonella'' is a genus of bacillus (shape), rod-shaped, (bacillus) Gram-negative bacteria of the family Enterobacteriaceae. The two known species of ''Salmonella'' are ''Salmonella enterica'' and ''Salmonella bongori''. ''S. enterica'' ...
'' species, so keepers should practice proper hygiene, such as washing their hands after handling the animals or their wastes. Dogs, even well-behaved dogs, often attack or chew on tortoises, so great care must be taken if they are around.


Housing

All tortoises should be housed outdoors when conditions allow. Outdoor pens should allow space for exercise, secure walls that are at least 1.5 times taller than the tortoise is long, and security from both predators and escape. Shelter and water should always be provided. Indoor housing is generally scaled to the size of the tortoise and must be secure and waterproof for this high-humidity species, as well as offering adequate space. Aquaria and plastic tubs are often used for younger tortoises, while a "tortoise table" (similar to shelf-less bookcase lying on its back), indoor minigreenhouse, or large indoor pen can be used for larger tortoises or groups of tortoises. An indoor habitat should be lined with a substrate, such as hardwood mulches (like cypress), coconut coir, soil and sand mixes, or some combination. A hide and water dish are necessary, and live or silk plants also help.


Environment

Heat, light, and humidity must be kept within proper guidelines for healthy tortoises. Red-footed tortoises are most active at temperatures from . A warmer area of is recommended, and night temperatures can drop a few degrees lower. High levels of humidity should be available in some part of the habitat. Lighting should be low and diffuse or plenty of shade should be offered. Lights that emit
UVB Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of the ...
wavelengths are recommended to help the tortoise metabolize calcium correctly and help regulate the
pineal gland The pineal gland (also known as the pineal body or epiphysis cerebri) is a small endocrine gland in the brain of most vertebrates. It produces melatonin, a serotonin-derived hormone, which modulates sleep, sleep patterns following the diurnal c ...
if the tortoise will be indoors for extended periods.


Captive diet

Pet tortoises should be allowed to forage in a well-planted outdoor pen whenever possible. The basic diet should consist of a variety of plants, vegetables, and fruits with occasional meat, and be high in calcium and fiber, and low in sugars and fats. Fruits should be kept as whole as possible. Commercial tortoise pellets can be offered along with fresh foods. The bulk of the diet should be leafy greens, such as turnip or collard greens, dandelion, leafy or curly lettuces, endive, kale, cabbage, edible tree or plant leaves, such as mulberry or hibiscus, lettuce mixes, parsley, grape leaves, etc. Vegetables and fruits offer variety and other nutrients, but are a smaller part of the captive diet. Good choices would include: cactus pads and fruits, papaya, figs, mango, mushrooms, pumpkin, squash, melon, pineapple, strawberry, cucumber, hays and grasses (such as wheatgrass), corn, peas and beans, carrot, apple, pear, plums, and okra. Meats form a very small part of the captive diet and can include live bugs or invertebrates (especially slugs), baby mice or rats, chicken, egg, organ meat, lean beef, tuna or other 'oily' fish, or cat or dog food. Calcium should be supplemented frequently in very small amounts. Other vitamins can be offered in very small amounts as well, but are best provided through a balanced, varied diet.Pingleton, p. 77-80.


References


External links


See video of red-footed tortoises at the Naples Zoo

Red Foot Tortoise
gov.vi

{{DEFAULTSORT:Red-Footed Tortoise Chelonoidis Turtles of South America Turtles of Brazil Reptiles of the Caribbean Reptiles of Saint Barthélemy Reptiles of Saint Martin (island) Reptiles of Guyana Vulnerable animals Vulnerable biota of South America Reptiles described in 1824 Articles containing video clips