Classification: Introduced Species - While the Red-Footed Tortoise has been seen and identified in multiple states, but at this time there are no records of any breeding populations in the wild. Sightings of this species are most likely of released or escaped pets that were imported for the pet trade.

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 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 (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, and rarely any sclera 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 (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 12 to 14 inches with males slightly larger overall. Tortoises up to 18 inches are fairly common and over 20 inches are occasionally discovered. The largest known specimen, from Paraguay, was 24 inches long, and weighed over 62 lbs. 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.

Habitat: The preferred habitat of the red-footed tortoise varies somewhat by region, but generally includes fairly consistent seasonal temperatures near 86℉ that rarely get lower than 68℉ or over 95℉, 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.

Range: Introduced into Florida, Georgia, Texas, & Hawaii with more sightings in Alabama, New Hampshire, and Mississippi. Red-footed tortoises natural range is from southeastern Panama to Venezuela, Guyana, Suriname, and Guiana in the north; south along the Andes to the west in Colombia, Ecuador, Peru, and Bolivia; east to Brazil, and along the southern range in Bolivia, Paraguay, and possibly northern Argentina. They are not evenly distributed within their range.

Found in these States: AL | FL | GA | HI | MS | NH | TX

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 than 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), figs (Ficus), pehen (Acacia aroma), Spondias, Annona, Philodendron, bromeliads, 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.

Reproduction: All turtles and tortoises start as eggs. Red-footed tortoise eggs are roughly spherical and average around 2.0 by 1.7 inches and weigh 1.8 oz. with two to seven eggs in a clutch, although the same females may lay multiple clutches near each other. The incubation period is 105 to 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 1.4 by 2.5 inch 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 7.9 to 9.8 inches, 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. 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, and 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 3.9 by 7.9 inches 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.

Status: The red-foot tortoise is considered vulnerable and is listed in CITES 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 yellow-footed and Chaco tortoises.

Subspecies: None

Taxonomy: