The Pancake Tortoise

David T. Kirkpatrick, Ph.D

Animals and their environments interact on various levels. Individual animals will seek out the habitat for which they are best suited, while on a larger scale populations of animals evolve to becoming increasingly well-adapted to the environment in which they live. This continuous back-and-forth interplay can have profound effects on the form and lifestyle of an animal, drastically altering its shape and habits in comparison to its ancestors and distant relatives. The Pancake Tortoise, Malacochersus tornieri, provides a perfect example of this process.

Rocky outcroppings with their many nooks, crannies and crevices provide protection for a number of species of animals. One tortoise, the Pancake tortoise, has evolved to take advantage of this cover. Unlike most tortoises, Pancakes do not possess domed shells. Instead, they have flat carapaces. Although they can reach a length of approximately seven inches, they rarely exceed an inch to an inch and a half in height. With this flat, pancake-like profile, Malacochersus tornieri is able to fit into narrow crevices and cracks in its rocky home, and thus exploit an environment that no other tortoise is capable of using. The shell of a Pancake tortoise is very flexible and can be bent and depressed with only mild pressure. This pliability lets the tortoise squeeze into very tight openings. The bones that underlie most tortoises' shells are solid, with very little space between the individual bones. However, in the Pancake tortoise these bones have many openings, or fenestrations, in them, which gives the shell its flexibility. In fact, early reports of the Pancake tortoise often refer to it as the "Softshell" tortoise. In other tortoise species, hatchlings and young juveniles possess similar fenestrations in the bones under the carapace and plastron, but, unlike in the Pancake tortoise these gaps fill in as the tortoise grows. Basically, M. tornieri has sacrificed its armor plating to gain a safe hiding spot.

Pancake tortoises are incredibly well adapted for the rocky outcrops, called kopjes, of south-eastern Africa where they are found. Other adaptations in addition to their flexible, flattened profile aid the tortoise in its home. Pancakes have enlarged scales on their legs, with points that project both downward and outward. When a tortoise is in its favorite crevice, it rotates its legs outward, and in this position is almost impossible to remove. At one time it was thought that they also inflated their body while wedged in cracks, in order to fit even more tightly. However, L. C. Ireland and Carl Gans demonstrated in 1972 that this inflation did not occur. Unfortunately, cracks and crevices only provide protection if you can reach them. Luckily for the Pancake tortoise, less bone means a lighter animal, so Pancakes can move more quickly than a tortoise with a thick shell. It is quite possible that they are the fastest tortoises in the world. When threatened, a Pancake will not crouch down and withdraw into its shell. Instead, it will make a dash for the nearest rocky shelter. Pancake tortoises tend not to stray very far from protection, and they also seem to be able to locate their favorite hideaways quickly when displaced. Finally, the Pancake's flat shell and extreme agility allow it to flip itself over quickly when it lands on its back, a handy ability to possess for a tortoise that spends most of its time climbing around on boulders and rocky hills.

Pancake tortoises have a relatively limited distribution in Kenya and Tanzania. As previously mentioned, they live in dry areas of scrub brush on rocky hills or outcroppings. The kopjes Pancakes inhabit can be located from 100 to 6000 feet in altitude. This variation in altitude means that Pancake tortoises have to be able to adapt to a range of temperatures. Average temperatures tend to fall around 24 to 30 degrees C., but can fall as low as 12 degrees C. Because Pancakes tend not to venture very far from protection, and favorable habitats can be separated by large distances, colonies of Pancake tortoises generally are isolated from each other. Within one habitat, however, they coexist peacefully, with many individuals sharing favorite crevices or basking sites. The only exception to this occurs during the January to February mating season, when males may fight among themselves. In the wild, individual animals can be very hard to spot. Although they leave their cracks and fissures in the morning to bask in the sun and to eat, the brown or horn color of their carapace, combined with a random pattern of radiating lines on each scute, allows Pancake tortoises to blend into the background, making them difficult to see.

Dry grasses and vegetation make up the bulk of a Pancake tortoise's diet in the wild. In captivity they eat much the same things as in the wild, but will also take food such as lettuces, squash, carrot, broccoli, green beans, cabbage, some melons, and occasionally other fruit as well. Vitamin supplements are not essential if a varied diet is provided, although a small amount can be given infrequently. Pancakes seem to get most of their water from the foods they eat, a survival trait in their natural environment. In captivity, a water bowl can be provided, although they rarely use it. Alternately, they can be misted heavily with a house plant mister once or twice a week. A once or twice monthly soak in warm water can also be beneficial.

Their small size and unusual appearance make Pancake tortoises desirable animals for both individual and institution collections. Fortunately they make hardy captives, if their needs are properly addressed. Although they are small compared to some tortoises, they still require a large area due to their active natures. Their enclosure should have a number of hiding spots, at least equal to the number of Pancakes kept, and preferably more. The area should be furnished with a variety of surfaces, including rocks, for the tortoises to climb around and over. A sand pit four or five inches deep should also be available, especially if breeding is desired. When constructing an enclosure, keep in mind that Pancake tortoises are excellent climbers - they will climb over any wall that is too low and disappear under the refrigerator, stereo cabinet, or similar heavy item of furniture if indoors, or disappear entirely, probably forever, if outdoors. In the early morning Pancakes will venture out from their favorite hiding spot, bask for a while to warm up, and then climb around the cage searching for food or an escape route. However, by mid-day they generally retreat back into their favorite crevice.

Pancake tortoises become somewhat more active during the breeding season, especially the males. If two or more males are kept together, they will sometimes fight over the females, who tend to ignore the males unless the male is actively pursuing or circling her. One male in our collection consistently waited until the females were eating in the morning before sneaking up and mounting them from behind. The females would often continue eating for quite a while before noticing the presence of the male. If you wish to breed Pancake tortoises, at least a pair is necessary, of course, but some sources recommend keeping a group of tortoises together, including three or four females and at least a pair of males. Combat between males prior to breeding has led to better reproductive successes in a number of tortoise species, including Malacochersus tornieri. Sexing adult Pancake tortoises is relatively easy. Adult males can be distinguished from females by their larger and longer tails. Additionally, males tend to be smaller than the females, and the rayed pattern on their scutes less distinct.

Mating among Pancake tortoises in the wild occurs near the beginning of the year, with nesting in July or August. In captivity animals will breed year-round, but breeding success can be increased by providing the Pancakes with a yearly light cycle in which the length of the "day" varies over the course of the year, similar to that provided for other turtles, snakes, and lizards. Females that have successfully mated will usually lay a single egg, burying it under three to four inches of loose, sandy dirt, although some will deposit eggs in out-of-the-way places in the cage if they find the nesting area unsuitable. One female can lay multiple eggs over the course of a single season, with eggs appearing every six weeks to two months. Each egg is about two inches long, but only half this in width. Eggs can be incubated at approximately 30 degrees Celsius; incubation seems to last for 140 to 190 days although incubation lengths of over eight months have been reported. Females that are laying eggs should have their diet supplemented with calcium, to aid in egg development.

Surprisingly, hatchling Pancake tortoises look like most other hatchling tortoises. Typically, hatchlings are dark yellow, with brown or black markings on the scutes on both their plastron and carapace. Their carapace is even slightly domed, unlike that of their parents! Upon hatching, juvenile Pancakes are about one and a half inches long. Hatchlings and juveniles eat the same diet that adults eat, although growing animals should be given calcium and vitamin supplements in addition, to insure proper growth. Hatchlings should not be overfed, as this may adversely affect their growth.

Pancake tortoises can be hardy and long-lived in captivity. Frank Slavens' Reptiles and Amphibians in Captivity, 1991 edition, notes that the National Zoo in Washington D.C. has a female that has been in captivity for over a quarter of a century, as it was acquired in 1965. Captive breeding programs have been successful in zoos and private collections around the country and few problems have been reported in rearing captive born animals. These successes in breeding and long-term care are heartening, as the wild populations of Pancake tortoises are not extensive. Moreover, most of the individual animals in the reptile trade still originate from wild populations. Pancake tortoises are perennially rumoured as about to be placed on CITES Appendix II, which would limit imports of wild-caught individuals. As breeding occurs readily in captivity given the proper set-up and care, anyone keeping Pancakes should strongly consider establishing a breeding group. If captive born individuals are available, the necessity for wild-caught animals will be greatly reduced.

Malacochersus tornieri, a strange and fascinating tortoise, has given up its hard, domed shell to take advantage of an unexploited environment - the dry, rocky kopjes of south-eastern Africa. Evolutionary forces have adapted the tortoise over the years to take full advantage of this environment. However, this adaptation is not without its price. Because one kopje is separated from another by open stretches, the Pancake tortoise, no longer protected by a hard shell, cannot readily cross this gulf, and so populations have become isolated. In return for increased protection in a hostile environment, the ancestors of the Pancake tortoise gave up their freedom to roam.

This article copyright 1993 by David T. Kirkpatrick. Originally published in Reptile & Amphibian Magazine, May/June 1993, pages 2-9.

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