REPTILIA 43 descendant of an extinct European of southern through to Sumatra, and the Tertiary. A fossil specimen found in the Üetliberg Borneo. However, attempts by herpetologists to find region (near Zurich, Switzerland), and now at the the species in the wild have been mostly unsuccessful. Zoological Museum in Zurich, is Although it has sometimes been larger but otherwise morphologi- possible to gain limited access to cally almost identical to M. . the region, an actual expedition is The Tertiary is a period of mod- almost impossible for political ern geological time characterized and military reasons. Also, the by the formation of the large fold dense vegetation in the distribu- mountains and the sunken area of tion area does not exactly permit the Mediterranean basin. pleasure-hiking. Manouria emys inhabits ever- Description green tropical rainforest and Manouria emys phayrei grows mixed deciduous transitional for- larger than Manouria emys emys, est. Temperatures during the and there are clear morphological coldest months average about differences between the two sub- 18°C (64°F). Climate studies of species. The shells differ in gen- Bhamo (northeastern Upper eral form, apparent from the rear Burma) can serve as an indica- view, and in M. e. phayrei, the tion of appropriate environmen- pectoral scutes meet along the tal conditions for the species. plastral midline, whereas in M. e. This information is important for emys they do not (see Figure 1). Distribution of Manouria emys according to Iverson setting up a terrarium. Manouria emys phayrei is the largest Asian . It grows to 60 centimeters in cara- Captive housing pace length and weighs up to 37 kilograms. The plain Our three sexually mature specimens are provided brown carapace is highly domed. The extremities are with a summer outdoor enclosure of about 10 x 11 broad and plump. The anterior and posterior marginal meters. Through the middle runs a large hill about 8 scutes are upturned. Large pointed tubercles on the meters long, 4 meters wide, and 2 meters high. At one thighs are responsible for the Thai name tao hook dum, side there is a round built-in bath of 3 square meters and which means something like “brown six-footed .” 50 centimeters in depth. The indoor enclosure is a small free-standing house that Adult specimen eating the grass that grows in the garden Distribution and habitat the can enter or exit at will during the day. It has Manouria emys phayrei is supposedly found from the a floor area of 3 x 4 meters and a sloped roof from 2.5 to In the wild, the Asian or Burmese brown tor- Manouria (Indochinese tortoises), species M. emys Assam region of India through Burma to northern and 3.4 meters high. The floor, sides, and roof are all insulat- toise, Manouria emys phayrei (Blyth, 1853), (Burmese brown tortoise), subspecies M. e. phayrei. west-central Thailand; Manouria emys emys, from ed. There are windows in the south- and east-facing walls. lives in highly inaccessible areas, and reports The species was originally described as emys of field observations are therefore practically by Schlegel and Müller in 1844. The subspecies phayrei Figure 1. Shell differences between Manouria emys emys and Manouria emys phayrei. unavailable. Hence, an understanding of this was first described by Blyth in 1853. Reiman later species must be formed mainly on the basis of its anato- described the same taxon as Manouria emys nutapundi Shaded areas show how the pectoral scutes of M. e. emys do not meet my and physiology in conjunction with climate studies. in his 1978 book, which was greatly influenced by Thai at the plastral midline, whereas those of M. e. phayrei do. That is the aim of this article. zoologist Wirot Nutaphand. However, the older name In German, this tortoise was once referred to by the for the subspecies should still be valid. impressive common name Thailändische Riesenschild- The definitive names of the two subspecies now gen- kröte (Thai giant tortoise). However, borrowing from erally recognized are Manouria emys emys (Schlegel the English common name, the German name has been and Müller, 1844) and Manouria emys phayrei (Blyth, Rear view of the shell of Manouria emys emys simplified to the rather nondescript Braune Land- 1853). Because the tortoises were originally assigned schildkröte (brown tortoise). In Thailand it is called to the Testudo, the names of the authors are tao hook dum — say these syllables out loud, and it written in parentheses. In any case, the subspecies sounds something like the tortoise working its way name phayrei should not appear with the authors through the brush. Reimann or Wirot. A second species of Manouria has also been described: the , Manouria impressa The full scientific classification of this taxon is (Günther, 1882). as follows: Testudines ( and tortoises), Plastron of Manouria emys Plastron of Manouria emys Origin emys phayrei suborder (turtles capable of retracting their Rear view of the shell of Manouria emys phayrei heads into their shells by means of a vertical S-shaped Manouria are considered probably the oldest of bend), Testudinidae (true tortoises), genus known living tortoises. Manouria emys is a direct

REPTILIA 44 REPTILIA 45 Figure 2. Comparison of average maximum temperatures in they must rely on the continuous ºC Bhamo (Burma) and Zurich (Switzerland) availability of water. Zoo specimens 40 in Thailand lie in shallow water for 35 Bhamo days, and we have observed that in 30 our captive setup the tortoises use 25 20 their baths fully, especially when 15 temperatures are above 24°C (75°F). 10 Zurich Our tortoises often defecate in 5 their indoor bath, but feces are usu- 0 ally not found in the outdoor bath. The water container must be July May April June cleaned daily, and the them- March August January selves are also regularly scrubbed October February December November under running water. They like to September Month bury themselves in the straw, which appears to also have a cleaning effect — the tortoises always look The floor of the house is covered with They like to sit for long periods of clean and polished. bark mulch, on which a 30-centimeter time in piles of branches or under layer of straw is strewn. In one cor- shrubs. Indoors they bury them- Temperature and light ner there is a bath 60 centimeters selves in the straw with only their Tortoises of this species do not square and 7 centimeters deep. In nostrils or eyes just barely showing. really bask, and in the beginning another corner, a shelf is mounted 60 our freshly imported specimens centimeters above the floor, under Water even avoided bright light for sever- which the tortoises sleep. An electric The rainy climate of their natural al years, retreating to the cooler oil-filled radiator is set on top of the range corresponds with a very inter- shade. In recent years they tend to Female M. e. phayrei preparing to bask shelf to control the ambient tempera- esting physiological characteristic of spend more time lying in the sun- ture within the house, and a heat mat these tortoises. Manouria species do light that comes in through the win- intestinal structure (including a well-developed cecum) In winter, when they stay indoors, the tortoises are is set on the floor under the shelf on not excrete uric acid — the white dow of their house, but they have of this species indicate that it is herbivorous. The beak is given salad greens such as endive, lettuce, and chicory. the coldest days. insoluble form of nitrogenous waste never been seen basking outdoors. worn down sufficiently only when tough, stemmy fodder This is given whole, and always covered with a good The adult tortoises are active in excreted by other tortoises (e.g., Juveniles like to lie under a halogen is provided. The intestinal flora — including flagellate handful of hay. If the food is a little late, the tortoises the morning, take a siesta during the Testudo hermanni). Instead, these lamp, or by the window in the sun. and ciliate protozoa, which help in the breakdown of cel- start eating the straw that is strewn as a floor covering. heat of day, and become active again species excrete nitrogen as urea, In winter, the ambient tempera- lulose — also indicates a vegetarian diet. Our specimens in the evening. On balmy summer requiring more water, and reflecting ture in the tortoise house should be graze like cows, and no vegetation is too tough for them. Social and mating behavior evenings they may graze long into the fact that these tortoises do not kept at 20–28°C (68–82°F). On very With this diet, the tortoises defecate large quantities. A special characteristic of these tortoises is the way the twilight. experience a dry season. In captivity cold days the floor temperature Their feces is compact, sausage-shaped, and dark olive- they greet each other with head bobbing. This behav- could drop too low, and an addition- green in color. We strongly recommend against adding ior can also mean the beginning of combat. Such head al infrared lamp may be needed. fruit or products to the diet. We notice that large bobbing, by both animals, can go on for several min- The tortoises can go outside when numbers of slugs crawl utes. The male also bobs the temperature is above 18°C freely through the enclosure its head at the female in (65°F), but even in summer they without the tortoises show- courtship. always spend the night inside their ing any interest in them. The female can emit noises house. In summer the tortoises that sound like a combina- eat what grows in their en- tion of muffled growling Diet closure — mostly well and groaning. During cop- Sharing experiences with other established grass. From ulation, the male emits Manouria keepers has shown that April to October, no other long (2–3 seconds) rhyth- diet is a tricky topic. Our feeding food is given; they “hunt” mic muffled groans, start- scheme is based on morphological for their own food. In the ing with a deep pitch and considerations, and supported by 20 morning they leave their then rising to a higher years of experience keeping these shelter and stop at the first pitch over a period of a few tortoises. green thing they come to. minutes. Thai biologist Wirot Nutaphand Then, not being at all picky, Males initiate copulation in believes that Manouria emys follows they simply eat everything all seasons. Although it has mountain streams, from which it within reach of their necks not been observed when eats vegetation such as lotus blos- in all directions, and work temperatures are very hot

3-week-old hatchling soms. The chewing apparatus and their way gradually forward. M. e. phayrei 1-year-old juvenile in the summer, copulation

REPTILIA 46 REPTILIA 47 suddenly go out, always during the first week of May, we learned to remove the shelf in mid April. Since then our coffee machine has worked uninterrupted through the spring.

Eggs During the first week of May, practically to the exact week, egg- laying takes place, usually secretly. The clutch consists of 45–60 soft- shelled spherical eggs. They are often dented on the sides where they touch each other. As soon as they are discovered, usually on the same day they are laid, the eggs are transferred to an incubator. The female continues to guard the nest for 2 weeks at the Female M. e. phayrei laying eggs most, and then interest is lost. This year we did not remove the eggs, and the nest was guarded for at least 6 weeks. One of the females was constantly sniffing and rearranging and adding substrate to the mound. But finally interest again was lost. These eggs died because the nest substrate was not at all suitable. Close-up of a juvenile M. e. phayrei We recommend quickly transfer- ring the eggs to very moist substrate otherwise takes place year-round, especially before mid- female is driven away. The strength of these tortoises for artificial incubation. The dented day. The male first sniffs the front legs of the female, is incredible, and if they are to be separated, it is quite sides usually return to their spherical and then, after this initial contact, moves to her rear. difficult to loosen their anchor grip on the ground. shape in a very short time. If they The whole procedure is very peaceful; females seem to The winning female then begins to meticulously build are not quickly reburied and kept accept it calmly. Aggressive biting or chasing through her nest. For hours she uses her front legs to move sub- moist enough, however, the eggs will the enclosure, as seen in Testudo hermanni, has not strate back into a pile. In the tortoise house, we keep become permanently dented. been observed in Manouria our specimens on a substrate of We incubate eggs in vermiculite or emys. bark mulch and straw, and no expanded clay granules. Because of Once a year, our male is stem remains unused. Finally the our inexperience with incubating Male observing the eggs taken away and put with a female climbs on top of the soft-shelled eggs, we had to pay the female of a different group for mound and waits. Our tortoises price of the learning process. We several weeks. When he are usually kept together in a were so impatient that we some- returns home, head bobbing is group, and we have often seen the times checked the eggs daily. But it fervent, and not 5 minutes go male come merrily along and dig was not until after 3 weeks that the by before he mounts a grazing into the nest mound, oblivious to first blood vessels actually became female, which she does not try all the hard work of the female — visible. During the first successful to prevent. typical male (sorry!). incubation, we sorted the eggs by For several years, we repeatedly smell. Many eggs developed embry- Nest building experienced an embarrassing os, but for some reason these died Manouria emys has a particu- mishap. The covered sleeping cor- and the eggs putrified. larly characteristic behavior ner of the tortoise house — with One day before hatching, the when preparing to lay eggs. the electric radiator mounted on eggs begin to sweat and the shell About 2–3 weeks before laying, the shelf above, and the heat mat develops hairline cracks. In our Recently hatched neonate the female becomes very terri- on the floor — was regularly cho- first successful incubation, eclosion torial. Our two females sometimes engage in serious sen for the nest site. With the added height of the nest, started after 67 days, and the last combat. With retracted heads, they move back to get a the carapaces of the females would push up on the shelf egg hatched after 74 days. Hatch- running start, push off with their hind legs, and ram each and move the radiator, causing a short circuit. After 3 lings emerged with shockingly large other with the fronts of their carapaces. The weaker years of having our coffee machine and the kitchen light yolk sacs, and were immediately Eclosion has begun!

REPTILIA 48 REPTILIA 49 Group of M. e. phayrei juveniles transferred to small plastic contain- quite compact. In 6 months, weight their food should be tough and ers lined with rings of moistened increased by 50–100 percent. After stemmy enough to prevent the beak gauze. Here the hatchlings were left 3 years, the tortoises weigh six times from growing too long. Otherwise to rest and absorb their yolk sacs, their birth weight. In size, I believe we have had no health problems which surprisingly were gone in 3 the tortoises are growing too fast, with our tortoises. days. At birth, our hatchlings have but compared to specimens kept by weighed 41–59.1 grams. other keepers, ours are are among Conclusion the smallest. The Burmese brown tortoise is a Hatchlings Like the adults, our juvenile tor- quiet and fascinating animal that can Manouria emys hatchlings have a toises are fed a vegetarian diet. apparently be kept in captivity with- uniform warm brown coloration, They eat eagerly and are not picky. out any problem. Like other tortois- and strongly serrated margins. The We offer them whatever “weeds” es, it reacts with consternation to egg tooth is well developed. The we get from the garden. In the win- being moved to a new enclosure, but extremities are covered with hard ter months, we offer endive, lettuce, within its usual home environment it scales. The babies sometimes start and similar leafy greens. Hay is is very curious, and shows no aggres- hissing even before they are fully turned down, and the woodlice and sion toward conspecifics except dur- out of their shells. They panic when snails that get into the tortoise ing egg-laying. they are picked up, and hide imme- enclosure are fully ignored. For proper captive care of this tor- diately when they are released. A daily bath is important for the toise, the keeper must consider the We keep hatchlings on very moist young tortoises, and they are often significant size and weight of the bark mulch. Outdoors, great care seen soaking in the shallow trough. species. Manouria emys should not must be taken to ensure that the The water must be changed several be kept without an enclosure of at humidity is kept high enough to pre- times a day because bathing evi- least 100 square meters, summer vent eye inflammations. Being dently stimulates defecation. and winter, otherwise it will feel transferred from indoor to outdoor So far we have not observed any cramped. Our winter enclosure is enclosures is especially stressful for rivalry among juveniles. Our speci- too small, so we no longer keep our the baby tortoises, and can cause mens are now in their fourth year. specimens . panic reactions. We are therefore Manouria emys require an envi- The tropical climate to which this using the outdoor enclosure less. ronment with rough surfaces and species is native must also be taken The tortoises from our first suc- plenty of space to move around in into account. Manouria emys needs cessful incubation show surprisingly so that they can sufficiently wear a very warm and wet environment, harmonious growth. They have down their claws and the horny with warm showers. The diet should smooth domed carapaces, and look scales on their extremities. Also, be entirely vegetarian.

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