Geographic Range
Manouria emys
is native to southern and southeastern Asia, ranging from extreme eastern India (Assam)
through, Bangladesh, and south and east through Myanmar, Thailand, Malaysia, and onto
the islands of Sumatra and Borneo. Imported specimens have been found in Vietnam.
Habitat
Brown tortoises are found in temperate, moist habitats that are influenced by monsoon
rains. Moderate temperatures of 55 F to 85 F (13 to 29 degrees Celsius) are preferred.
To survive, young
Manouria emys
need the temperature to be above 65 F (18 degrees Celsius). This species also prefers
a humidity of around 60% to 100%. In addition, these tortoises never wander far from
a water source, such as a pond.
M. emys
are typically found in highland tropical forests, and prefers cooler and moister
conditions than other tortoise species.
- Habitat Regions
- temperate
- tropical
- terrestrial
- Terrestrial Biomes
- forest
- rainforest
Physical Description
Manouria emys
is the largest tortoise inhabiting Asia. Several characteristics of its shell distinguish
it from other species. The cervical scute in the shell of the brown tortoise is shorter
and wider when compared to other south Asian tortoises. The brown tortoise also has
divided supracaudal scutes, whereas other tortoises do not. It has many scales on
its hind thighs, a domed carapace, slightly serrated posterior marginals, and varies
in color from brown to black, depending upon the subspecies. There are two recognized
subspecies:
Manouria emys emys
is commonly called the Asian brown tortoise, while
M. e. phayrei
is called the Burmese brown tortoise. The Burmese brown tortoise is larger (it can
grow to 60 centimeters and weigh 37 kilograms) and is darker. The brown tortoise can
grow to 50 centimeters and weigh 20 kilograms. Like all tortoises, Males tend to have
longer, thicker tails than those of females and also have concave plastra (which functions
to facilitate mating).
- Other Physical Features
- ectothermic
- heterothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes shaped differently
Development
We have no information on development in this species.
Reproduction
Courtship in this species is more elaborate than in other tortoises, During courtship, males head-bob to communicate with prospective mates. The female typically does not bob her head in response during courting, though females do bob heads to communicate with each other and with males when not breeding. The two types of head bobbing are horizontal and elevated bobbing. Fixation is another courting behavior performed by males. Males fully extend their head and neck and keep it pointed towards the female as the female moves about. As in head-bobbing, the female does not fixate on the male. Trailing is a maneuver in which the male follows the female very closely behind. If the female slows down or stops, the male usually attempts to mount her. If she does not slow or stop, he resorts to biting her to get her to hold still as he mounts her. Both males and females vocalize while courting. The sounds are low, resembling groans and moans, and most often occur during the head-bobbing phase of courtship. Male vocalizations are typically longer in duration than female vocalizations. As with other tortoises, males vocalized during mating is well.
We don't have information on whether males or females in this species mate more than
once per season, but it's likely that as with other tortoises, both males and females
may have multiple mates, with males more frequently mating multiply.
- Mating System
- polygynandrous (promiscuous)
Reported clutch sizes range from 21 to 53 eggs, but these are all from captive females.
Incubation time ranges from 63-84 days, also in captive situations with artificial
incubation. We have no information on reproduction in this species in the wild.
- Key Reproductive Features
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- oviparous
Unlike most tortoises, females of
Manouria emys
do invest in their offspring after they lay their eggs. For a week prior to laying
her eggs, the female gathers up leaves and ground debris to make a mound into which
she will deposit her clutch. Brown tortoises use their forelimbs to gather debris
while other tortoises use the hind limbs to excavate nest sites. After she finishes
laying, the mother tortoise covers her eggs with vegetation and litter, and remains
at the next site, guarding her eggs and frequently piling more vegetation on them.
If a potential predator approaches her nest, she first attempts to drive it away by
pushing and even biting. If the predator is too quick or persistent, she will passively
defend the eggs by crouching over them. This behavior is unique among chelonians,
no other turtle or tortoise exhibits this high a level of parental care.
- Parental Investment
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
protecting
- female
-
protecting
Lifespan/Longevity
The biggest limiting factor in lifespan seems to be predation, either by humans or
from habitat destruction.
Behavior
Manouria emys moves very slowly, even when threatened. The species is most active during twilight, or during the day if the temperature is not too high. If the climate is too warm, brown tortoises may burrow into damp soil and leaf litter. They also are found soaking in pools and small streams.
Brown tortoises seem to have more complex vocalizations and other communications within
their species than other tortoises do, but they are not known to be particularly social.
Males engage in vocal disputes and shoving matches to discourage rivals from courting
nearby females, and females defend their nest sites from other tortoises, but we have
no additional information about sociality in this species.
- Key Behaviors
- terricolous
- crepuscular
- motile
- sedentary
- solitary
Communication and Perception
This species has more elaborate communication behaviors than other tortoises.
Head-bobbing seems to be a form of social and courtship communication. The two types
of bobbing are horizontal bobbing and elevated bobbing. Bobbing is not specific to
one sex or the other and can be used by an individual towards either sex. However,
during courting by the male, the female does not normally reciprocate head bobbing.
At other times, however, head bobbing was reciprocated. Head-bobbing is often accompanied
with vocalization.
"Fixation" is a sexually oriented behavior in which a male fully extends his head
and neck and keeps it pointed towards the female as they move about. The female does
not ever "fixate" on the male. Fixation seems to be an early part of courtship, whether
or not the male is successful in courting her.
Trailing is another courting maneuver in which the male follows the female from behind.
When the female slows down or stops, the male attempts to mount her. However, if she
does not slow or stop, the male resorts to biting her. He bites her nose or legs in
an effort to get the female to hold still for mounting.
Another manner of communication is vocalization. Males tend to have rhythmic grunts
or bellows. Both males and females have mating calls that they use while head bobbing
occurs. Typically the male has longer and more frequent calls when compared to the
female. Other purposes of vocalization are not entirely known at this point.
- Perception Channels
- visual
- tactile
- acoustic
- vibrations
- chemical
Food Habits
Brown tortoises are herbivores for the most part. Their diet typically consists of
grasses, vegetables, leaves, and fruits. They have been known to eat other animals
on occasion, most of which were small invertebrates and amphibians.
- Animal Foods
- amphibians
- terrestrial worms
- Plant Foods
- leaves
- seeds, grains, and nuts
- fruit
- Other Foods
- fungus
Predation
This species relies on its strong shell for protection from predators. It's preference for twilight hours for activity, and its brownish color of it may also help avoid predators. Female protection and maintenance of the nest may help protect her offspring during their most vulnerable time of life. Though she cannot deter a vigorous egg-eater, simply by distracting predators and creating a disturbance she may discourage or confuse them. Her frequent nest-building may also help hide the scent of the eggs.
Currently humans are by far the most dangerous predator to this species. We have
no specific information on the predators of
M. emys
, but it is likely that large predators, like tigers, canids, or bears, might attack
tortoises. Many smaller predators like foxes and monitor lizards eat tortoise eggs,
and predator birds will attack juvenile tortoises.
Ecosystem Roles
As an herbivore,
M. emys
helps in cleaning up leaves, fungi, and fruit that are on the forest floor. They
do not have any commensal or mutual relationships that are known, and specific predators
of the tortoise were not available for study. Fruit seeds are dispersed through the
tortoise's feces.
- Ecosystem Impact
- disperses seeds
Economic Importance for Humans: Positive
Some people of Southeast Asia use
M. emys
as a food source and in medications. In addition,
M. emys
is a profitable animal in the international pet trade.
- Positive Impacts
- pet trade
- food
- source of medicine or drug
Economic Importance for Humans: Negative
There are no known adverse affects of Manouria emys on humans.
Conservation Status
In several parts of Asia, brown are exploited for food and medicine. Habitat destruction
is also a factor in their 'vulnerable' status. The pet trade is also a problem, as
Brown tortoises are fairly valuable and sought-after animals. Over-consumption and
habitat destruction have caused populations of this species to rapidly decline in
many parts of its range. The species is considered Endangered by the International
Union for the Conservation of Nature is listed in Appendix II of CITES. This is one
of many species terrestrial and freshwater turtles and tortoises in Asia that it threatened
by commerical over-consumption.
Other Comments
Many tortoise experts believe that this species resembles the ancestors of tortoises more than other tortois species, and may be the oldest tortoise species that exists. They base this idea on details of its anatomy, its ecological preference for a humid climate, and other aspects of its behavior.
Additional Links
Contributors
David Armitage (editor), Animal Diversity Web, George Hammond (editor), Animal Diversity Web.
Vijay Virupannavar (author), University of Michigan-Ann Arbor, Phil Myers (editor), Museum of Zoology, University of Michigan-Ann Arbor.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- solitary
-
lives alone
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- vibrations
-
movements of a hard surface that are produced by animals as signals to others
- chemical
-
uses smells or other chemicals to communicate
- pet trade
-
the business of buying and selling animals for people to keep in their homes as pets.
- food
-
A substance that provides both nutrients and energy to a living thing.
- drug
-
a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease
- herbivore
-
An animal that eats mainly plants or parts of plants.
- folivore
-
an animal that mainly eats leaves.
References
Alderton, D. 1988. Turtles & Tortoises of the World . New York, New York: Facts on File Publications.
Jacobsen, G., C. Tabaka. 2004. "Burmese Mountain Tortoise - Manouria emys " (On-line). World Chelonian Trust: Conservation and Care. Accessed September 23, 2004 at http://www.chelonia.org/Articles/burmmtortoise.htm .
McKeown, S., D. Meier, J. Juvik. 1991. The Management and Breeding of the Asian Forest Tortoise ( Manouria emys ) in Captivity. Proceedings of the First International Symposium on Turtles & Tortoises: Conservation and Captive Husbandry: 138-159. Accessed September 23, 2004 at http://www.tortoise.org/archives/manouria.html .
McKeown, S. 1990. Asian Brown Tortoise, Manouria emys . Tortuga Gazette , 33/6: 3-5. Accessed September 23, 2004 at http://www.tortoise.org/archives/manemys2.html .
Moll, E. 1989. Conservation Biology of Tortoises . IUCN - The World Conservation Unit.
Shaffer, C., V. Morgan. 2000. Behavioral observations of captive juvenile Manouria emys phayrei with notes on degrees of intergradation with Manouria emys emys . Turtle and Tortoise Newsletter , 5: 2-6. Accessed September 23, 2004 at http://www.chelonian.org/ttn/archives/ttn5/pp2-6.shtml .