Overview

Brief Summary

Biology

In keeping with its strange appearance, the Patagonian mara displays behavioural traits that are very unusual among the rodents. Active during the day, this species spends long periods basking in the sun, either resting on its haunches or with its front limbs folded under its body like a cat. It remains vigilant for predators, however, and when threatened, may escape by galloping or stotting, a high-speed form of locomotion involving rapid bounces using all four limbs at once. Surprisingly, this diminutive species can reach speeds of 45 kilometres per hour over distances in excess of one kilometre. The Patagonian mara consumes a variety of vegetation, usually feeding in grass or shrubland during the day and sheltering in an abandoned burrow of another species, or amongst dense vegetation at night (2). Perhaps the most unusual feature of this species is its social and reproductive behaviour. Patagonian maras are strictly monogamous (a rare trait in mammals), forming lifelong pairs that generally avoid contact with conspecifics, except during the curious arrangements that form in the austral summer breeding season. The female Patagonian maras dig burrows, in which the young are often reared communally (2) (3). After a gestation period of 90 days (3), the female gives birth to a litter, usually of two young, close to the entrance of a burrow (1) (2). The young, which are born well-developed, with eyes already open, quickly move into the burrow, which may be co-habited by the offspring of several other breeding pairs (2). The young remain in the vicinity of the burrow for the next four months, where they are visited daily by the mother to be suckled. When the breeding pair visits the burrow, the female makes a shrill call, which brings all the infant Patagonian maras in the burrow to the surface. The female then attempts to discern its own offspring within the group by smell, while chasing off the young of other pairs, which try to, and occasionally succeed, in obtaining the female's milk. Once identified, the female leads its offspring a short distance away from the burrow to be fed, while the adult male keeps watch, chasing off other pairs that approach. The other breeding pairs must therefore wait until the preceding pair have departed, before visiting the burrow. This communal behaviour is thought to help reduce predation, as it means that adults are more frequently present around the burrow (3).
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Description

With long hare-like ears and a body resembling a small deer, the Patagonian mara is a distinctly unusual looking rodent (3). The upperparts are brownish grey, with a darker patch on the rump bordered below by a white fringe, while the underparts are whitish (2) (3). The body is well adapted for running, with long, powerful hind limbs, ending in three digits, each bearing a hoof like claw. In contrast, the front limbs are shorter, with four sharp claws that help this species to dig burrows in which the young take refuge (2).
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Distribution

Range Description

This species occurs in Argentina, from Bolson de Pipanaco, Catamarca Province at approx 28 degrees south to 50 degrees south (Woods and Kilpatrick, 2005). Subspecies D. p. centricola occurs in Catamarca Province, the eastern region of La Rioja Province, the northwest of Cordoba Province, and the southwest of Santiago del Estero Province (Cabrera, 1953). Subspecies D. p. patagonum occurs in the central area of Buenos Aires Province, the south of Cordoba Province, San Luis and Mendoza Provinces, and to the south of Argentina to Santa Cruz Province (Cabrera, 1953).
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Geographic Range

Dolichotis patagonum is endemic to the open grasslands and shrubland steppes of Argentina, and is distributed between 28°S and 50°S.

Biogeographic Regions: neotropical (Native )

  • Campos, C., M. Tognelli, R. Ojeda. 2001. Dolichotis patagonum. Mammalian Species, 652: 1-5. Accessed March 15, 2011 at http://www.jstor.org.proxy.lib.umich.edu/stable/3504392.
  • Chillo, V., D. Rodriguez, R. Ojeda. 2010. Niche partitioning and coexistence between two mammalian herbivores in the Dry Chaco of Argentina. Acta Oecologica, 36/6: 611-616.
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Range

The Patagonian mara is endemic to Argentina, where it is widely distributed. Two subspecies are currently recognised: Dolichotis patagonum centricola, which occurs in the north within the provinces of Catamarca, La Rioja, Cordoba and Santiago del Estero; and Dolichotis patagonum patagonum, which occurs from the province of Mendoza, east to Buenos Aires, and south as far as Santa Cruz (1).
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Physical Description

Morphology

Physical Description

Dolichotis patagonum is the second largest member of the family Caviidae. One study reported average weights of males and females as 7.73 kg and 8.44 kg, respectively. However, most accounts report that males are larger than females. Length ranges from 610 to 810 mm with an average of 700 mm. They can be distinguished from other members of Caviidae by their large size, long rabbit-like ears (Dolichotis literally means "long ear"), and short, nearly hairless tail which it holds close to the body. Unlike other cavies, which have anal glands anterior to the anus, the anal glands of D. patagonum are located between the anus and the base of the tail. It has short, grizzly gray fur, with a large, conspicuous patch of white on the rump. Midway up the rump a sharply contrasting area of black occurs, which quickly fades to gray. The venter is white, with patches of rusty orange fur on the chin, cheeks, and flanks. Two subspecies are recognized: Dolichotis patagonum centricola and Dolichotis patagonum patagonum, which are distinguished based on geographic location and fur coloration.

Much like ungulates, Dolichotis patagonum has elongated metapodials in its hindlegs as a modification for fast and efficient running. Forelegs are significantly longer than in most other rodents, and both the hind- and forefeet are small with hoof-like claws. Forefeet have four digits while hindfeet have three, and all digits have a claw. The elbow lies relatively high on the forelimb as the radius is longer than the humerus. Dolichotis patagonum has a dental formula of 1/1, 0/0, 1/1, 3/3, and cheek teeth are hypsodont and evergrowing.

Average mass: 8.12 kg.

Range length: 610 to 810 mm.

Average length: 707.2 mm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

Average mass: 12500 g.

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
The species occurs in lowland habitats, including forest and creosote bush (Larrea) flats in northwestern Argentina (Campos et al., 2001). It prefers habitats with shrub cover but will also use overgrazed and bared soils in the Monte Desert biome (Kufner and Chambouleyron, 1991). Maras are herbivores (Kufner and Duranona, 1991). In southern Argentina, breeding is from August to January (Taber and Macdonald, 1992) and in Patagonia, most births occur between September and October (Taber, 1987). In captivity, maras produce three or four litters each year (Dubost and Genest, 1974), but in the wild, most females produce only one litter annually (Taber, 1987). Average litter size is two (range, 1-3) (Campos et al. 2001).

Systems
  • Terrestrial
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Patagonian maras live only in the arid central and southern regions of Argentina. Generally classified as desert, this area exhibits a wide range of distinct microhabitats ranging from sandy plains to thorny shrubland steppes. Rainfall is extremely unpredictable and there are huge shifts in precipitation and floral composition between wet and dry seasons. The area is quite warm, with average temperatures in the summer usually around 20 degrees Celsius and winter temperatures rarely dropping below freezing. Generally, Patagonian maras prefer to build dens in open habitat dominated by grasses and other low-growing plants. Though it dens in areas that are densely vegetated, they visually monitor for predators, which is less effective in more closed habitats. Large settlements of Patagonian maras have been observed on and around sheep ranches, most likely due to their similar habitat preferences.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: desert or dune ; savanna or grassland ; scrub forest

Other Habitat Features: agricultural

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The Patagonian mara inhabits lowland areas of coarse grassland or open scrub desert, but may also occur in forests (1) (2) (3).
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Trophic Strategy

Food Habits

The range of Dolichotis patagonum encompasses a variety of habitats from desert to shrubland steppe. A strict herbivore, D. patagonum shows considerable flexibility in adjusting its diet to different ecosystems. For example, at the southern edge of its range in Sierra las Quijadas National Park north of San Luis, Argentina, D. patagonum experiences considerable variation in rainfall throughout the year and the local environment undergoes considerable changes in respect to floral composition between wet and dry seasons. Regardless of season, however, grasses make up nearly 70% of D. patagonum's diet. Despite the fact that most plant biomass in the region consists of forbs and shrubs, D. patagonum selects relatively rare grasses as its primary forage. Most grasses consumed are in the genus Pappophorum. In total, D. patagonum forages on 24 species of grasses and 22 other species of plants. In addition to grass, a significant portion (11%) of D. patagonum's diet consists of various species of cacti. Generally, cacti is about 75% water by weight and could represent a significant source of water to this species. This may help offset the unpredictability of rainfall throughout the range of D. patagonum>.

Near the central part of its range, evidence suggests that the grass genera Poa and Panicum make up the bulk of Dolichotis patagonum's diet, followed by Stipa and Bromus. Open-shrubland inhabitants also forage on Doli chotis patagonum and grassland inhabitants consume Lycium. Dolichotis patagonum in sandy grasslands and lithosol shrublands prefer Prosopis. These habitat specific differences display the dietary flexibility of this herbivore. During droughts, Dolichotis patagonum adjusts its diet to include more moisture-rich plants.

Patagonian maras are hindgut fermenters. Their diet is very high in fiber and cellulose, which is broken down by bacterial fermentation in a pouch called the cecum attached to the large intestine. They produce a special feces, which is ingested and re-digested. Wild Patagonian maras have been observed near sheep ranches consuming sheep dung.

Plant Foods: leaves; seeds, grains, and nuts; fruit; flowers

Other Foods: dung

Primary Diet: herbivore (Folivore ); coprophage

  • Sombra, M., A. Mangione. 2005. Obsessed with grasses? The case of mara Dolichotis patagonum (Caviidae : Rodentia). Revista Chilena de Historia Natural, 78/3: 401-408.
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Associations

Ecosystem Roles

Patagonian maras are herbivores and affect their environment by grazing on plants. They also eat fruit and distribute seeds through their feces. Patagonian mara pups form an important prey base for many species of birds, canids, and felids. They are also host a number of parasitic roundworms, including Wellcomia dolichotis, Trichostrongylus retortaeformis, and Graphidioides affinis.

Ecosystem Impact: disperses seeds

Commensal/Parasitic Species:

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Predation

Dolichotis patagonum has evolved a predator response system very similar to that of ungulates. It relies on its well-developed senses of hearing, vision and smell for early predator detection in open habitats. In the the event of a chase, D. patagonum is able to run very fast (up to 45 mph). This species exhibits stotting behavior identical to that seen in some ungulates. Its brown coloration helps camouflage it from potential predators. The main predators of the mara are Suran's foxes, South American grey foxes and pumas; however, due to human activities populations of both of these species have declined sharply. Humans now present the main threat to D. patagonum, both through habitat alteration and poaching. Other predators include lesser grison and variable hawks.

Known Predators:

Anti-predator Adaptations: cryptic

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Life History and Behavior

Behavior

Communication and Perception

Dolichotis patagonum prefers open habitats. Similar to ungulates, D. patagonum relies on early detection of predators to allow adequate time for escape. Also similar to some ungulates, Patagonian maras exhibit stotting behavior, which is characterized by a bounding gait that advertises strength and speed and discourages a long and costly chase by the predator.

Like many rodents, Dolichotis patagonum has a pair of anal scent glands. Males are frequently observed anal dragging, which results in a unique looking scent mark. Anal gland secretions play a major role in the mobile territory that males form around their mate. It is not know whether D. patagonum uses auditory signals to communicate with conspecifics.

Communication Channels: visual ; chemical

Other Communication Modes: pheromones ; scent marks

Perception Channels: visual ; acoustic

  • Puig, S., M. Cona, F. Videla, E. Mendez. 2010. Diet of the mara (Dolichotis patagonum), food availability and effects of an extended drought in Northern Patagonia (Mendoza, Argentina). Mammalian Biology, 75/5: 389-398.
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Life Expectancy

Lifespan/Longevity

Little information could be found on the lifespan of wild or captive mara.

Average lifespan

Status: captivity:
14.0 years.

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Lifespan, longevity, and ageing

Maximum longevity: 14.4 years (captivity) Observations: One captive specimen was still alive at 14.4 years of age (Richard Weigl 2005).
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Reproduction

Dolichotis patagonum is strongly monogamous, and male-female pairs usually bond for life. The pair-bond is maintained mainly by the efforts of the male, who follows and guards the female wherever she goes. To mark her as his territory, he urinates on her, spreads anal gland secretions around her and fiercely defends her from rival males. Males have been observed fighting, but there has never been a documented case of mate stealing. Male typically defend a 20 m^2 mobile territory centered around his female. Because estrus occurs once every 3 to 4 months and lasts for just half an hour, monogamy is advantageous, as spending all his time with one female assures a male that he does not miss his chance to breed. Monogamy may have also arisen as a response to the patchy and sparsely distributed food resources of the region. Since this sort of resource distribution is likely to result in a very scattered distribution of females, a male has the highest chance of successful breeding by finding one female and remaining with her. Another benefit to pair-bonds is that females are able to invest additional time and attention to caring for her young, relying on the male to watch for predators. Monogamy increases the males reproductive success both by lowering the death rate of his own offspring and increasing the longevity of his mate, allowing him more chances to breed.

Mating System: monogamous

Female Patagonian maras are sexually mature by 8 months of age. Estrus occurs once every 3 or 4 months and lasts only about half an hour. This extremely short estrous cycle is very unusual and most likely played a part in the evolution of monogamy in this species. Females in captivity often conceive shortly after parturition and can give birth to 3 or 4 litters per year, however, in the wild only one litter per year is produced. Litters range in size from 1 to 3 pups, with average litters containing 2. Rarely, a few females may have a second litter around January, but the majority of pups are born between mid August and late December. There is a large pulse of births between mid September and late October, with almost two thirds of pups being born during this time period. Gestation lasts an average of 100 days in the wild.

Females have 6 or 8 teats and nurse 1 or 2 pups at a time. On rare occasions females have been seen nursing up to 4 pups, suggesting occasional milk stealing by unrelated pups. It is thought that females recognize their young mainly by size, so pups of similar size to a female's own pups may have greater milk stealing success. Females also recognize their young by sound and scent cues. Occasionally, orphaned pups may be "adopted" by another female and allowed to suckle. However, females usually aggressively reject interloping pups by lunging, chasing, biting and shaking, or throwing them away from her. Some pups make frequent attempts to steal milk and many have tattered and damaged ears from these fierce rejections. Nursing bouts last for about half an hour and adult visits to the den usually last about an hour. Pups are usually nursed for about 75 to 78 days, which is unusually long compared to other rodents.

Like many ungulates, Dolichotis patagonum exhibits the "hider" strategy through the first part of their lives, staying close to the burrow and hiding at the signal from an adult sentry. As pups mature, they go through a "follower" phase where they trail behind their parents on foraging expeditions farther away from the burrow. Most young disperse at the time of weaning, however some stay with their parents until the next breeding season.

Though members of this species spend most of the year associating strictly as male-female pairs, when pups are born Dolichotis patagonum gathers in large groups around large warren "settlements" and raise young in communal creches. Each communal den has on average 4.26 breeding adults and 4.46 pups; however, up to 29 pairs have been observed sharing a single den. Since males aggressively guard their mates, only one pair at a time is present at the den.

Breeding interval: Patagonian maras breed year-round.

Breeding season: Patagonian maras primarily breed from mid August through late December.

Range number of offspring: 1 to 3.

Range gestation period: 91 to 111 days.

Average gestation period: 100 days.

Range weaning age: 75 to 78 days.

Range time to independence: 75 to 78 days.

Average age at sexual or reproductive maturity (female): 8 months.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous

Average birth mass: 561.94 g.

Average number of offspring: 2.

Average age at sexual or reproductive maturity (male)

Sex: male:
183 days.

Average age at sexual or reproductive maturity (female)

Sex: female:
158 days.

Males contribute very little in terms of direct parental care. They rarely interact with small pups, and interactions with large pups is limited to sitting or foraging nearby. However, males spend the majority of their time watching for predators, which significantly lowers predation risk faced by his offspring and mate.

Parental Investment: male parental care ; female parental care ; pre-fertilization (Provisioning, Protecting: Male); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male); pre-independence (Provisioning: Female, Protecting: Male)

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2008

Assessor/s
Ojeda, R. & Pardinas, U.

Reviewer/s
Amori, G. (Small Nonvolant Mammal Red List Authority) & Schipper, J. (Global Mammal Assessment Team)

Contributor/s

Justification
Listed as Near Threatened because, although it is widespread, its populations are declining due to loss of habitat, competition with large introduced herbivores and hunting, probably at a rate close to 30% over ten years. Almost qualifies as threatened under criterion A2cde.
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Dolichotis patagonum is classified as near threatened on the IUCN's Red List of Threatened Species. Habitat change, most likely caused by the introduction of domestic sheep, is a major problem facing this species. Overgrazing by sheep causes a shift from large grassy patches to a landscape characterized by smaller patches of woody plants. Reproductive success of D. patagonum populations in open grassland habitats tends to be greater than those in closed habitats. Dolichotis patagonum are also vulnerable to hunting, as it is killed for its meat and skin. Adults are killed with guns or wire snares, while pups are captured in nets placed over the entrances of burrows. Lepus europaeus (European hares), which is not native to South American, has introduced Jonhe' disease and toxoplasmosis to D. patagonum. In many areas, competition with L. europaeus is so intense that D. patagonum has been become locally extinct.

US Federal List: no special status

CITES: no special status

IUCN Red List of Threatened Species: near threatened

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Status

Classified as Near Threatened (NT) on the IUCN Red List (1).
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Population

Population
The population is decreasing and the species is not easily found (Ojeda, R. pers. comm.).

Population Trend
Decreasing
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Threats

Major Threats
Habitat destruction for agriculture, and hunting for their skins have resulted in localized extinctions such as Buenos Aires Province (Cabrera, 1953). The species competes with large, introduced herbivores such as sheep and the European hare (Jayat, J. pers. comm.).
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As a result of habitat loss, hunting and competition with large introduced herbivores such as sheep and the European hare, the Patagonian mara population is undergoing a worrying decline. In Buenos Aires Province, hunting for this species' skin and clearance of its habitat for agriculture has resulted in localised population extinctions. Unless measures are taken to mitigate the threats to this species, it is likely that it will soon qualify for threatened status (1).
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Management

Conservation Actions

Conservation Actions
The species occurs in at least 12 protected areas. It is categorized as Vulnerable in the Red Book of Mammals of Argentina (Diaz and Ojeda, 2000).
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Conservation

At present, the Patagonian mara is found in at least 12 protected areas within its range (1), where it received varying degrees of protection against habitat loss and hunting (4).
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

There are no known adverse effects of Dolichotis patagonum on humans.

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Economic Importance for Humans: Positive

Patagonian maras breed easily in captivity and make excellent zoo specimens. They can live with other representative species of the region and make for an exciting walk-through exhibit. They are very charismatic and are used as an ambassador species to educate the general public about the rapidly declining Pampas, a large area of South American lowlands. Patagonian maras can be kept as pets and can be trained to walk on a leash. They are hunted for their skins, which are used to make bedspreads and rugs in Argentina, and their meat.

Positive Impacts: pet trade ; food ; ecotourism

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Wikipedia

Patagonian Mara

The Patagonian Mara, Dolichotis patagonum, is a relatively large rodent in the mara genus (Dolichotis). It is also known as the "Patagonian cavy" or "Patagonian hare".

Contents

Description

Closeup of mara head

The Patagonian mara resembles a jackrabbit.[2] It has distinctive long ears and long limbs. It’s hind limbs are longer and more muscular than it’s forelimbs and it has a longer radius than humerus.[3] The feet are compressed, making them hoof-like. The forefeet have 4 digits while the hind feet have 3 digits. Its tail is short, depressed and almost hairless.[3] It has a gray dorsal pelage with a white patch on the rump separated from the dorsal fur by a black area.[4] In addition, the mara has a white underside with a somewhat orange flank and chin.[3] The mara as a head and body length of 690-750 mm with a tail of 40-50mm. It weighs 8-16 kg. Unlike most other cavids, the anal glands of the mara are between the anus and the base of the tail rather than being anterior to the anus.[3]

Ecology and activates

Mara foraging

The Patagonian mara is found only in Argentina. It is distributed 28ºS to 50ºS. Maras prefer to live in habitats with lots of shrub cover. However they also inhabit overgrazed and barren soils in the Monte Desert biome.[3] In northwestern Argentina the mara primarily inhabits lowland habitats like forest and creosote bush or larrea. Maras prefer sandy and low shrub habitat in Peninsula Valdes.[3] It well adapted to a cursorial lifestyle on the open plains and steppe, with its long legs, reduced clavicle and well-developed sensory organs making it capable of running and communicating in these open habitats. When running, maras have been compared to deer and antelope.[2] Maras are largely herbivorous. They feed primarily on green vegetation and fruit. In the Monte Desert, monocots make up 70% of its diet while dicots make up 30%.[5] Preferred grass species eaten are those of the genus Chloris, Pappophorum and Trichloris while dicots that are eaten are Atriplex lampa, Lyicum and Prosopis.[5]

Maras are primarily diurnal and spend around 46% of its daily activities are made of feeding.[6] The temporal activity rhythms of maras are related to environmental factors.[7] Light, precipitation and temperate have a positive effect on annual activities while darkness and relative humidity have a negative effect. The daily activity of the mara is unimodal in winter and biomodal in other season.[7] The preferred temperature is 20°C.[7] Females, due to the demands of gestation and lactation spend more time feeding than the males. Males spend most of the day sitting, being vigilant for predators.[6] Predators of maras, particularly the young, are felids, grisons, foxes and birds of prey.[6] Mara are also hosts for parasites like Wellcomia dolichotis.[3]

Social behavior and reproduction

Mara pair with young

The social organization has a unique combination of monogamy and communal breeding.[8] Being monogamous, pairs of maras stay together for life with replacement of partners only occurring after its death.[9] The males has almost the sole responsibility in maintaining the pair by following the female where ever she goes.[6] A male will mark his female with urine and mark the ground around her with secretions from his glands and with feces, making the grounds around the female a mobile territory.[9] Pairs will breed together alone or with other pairs in warrens shared by up to 29 pairs.[8] Maras breed, at least in southern Argentina, from August to January. Gestation lasts 100 days in wild. [6] Most births, at least in Patagonia, occur between September and October which is before the summer dry season and after the winter rains. [6] Female produce one litter each year in the wild, [6] but can produce as many as four litters a year in captivity.[9] Young can walk almost immediately postpartum.[3]

Young mara and older one sniffing each other. From the zoological garden of the Jardin des Plantes in Paris.
Mara at Barbados Wildlife Reserve, Barbados

Dens are dug during the breeding season for the young to be raised.[6] Litters from 1-22 pairs are grouped together in these dens. Communal living provides protection from predators with the survival rate for young being higher in larger groups than in smaller groups.[6] One pair visits the den at a time for around one hour and the other parents will circle around the den.[10] 1-2 pups are nursed at a time by a female.[6] A female may sometimes nurse a young from another pair. While a female may prevent young other than her own from nursing her, some young are able to steal milk.[9][8] There is no active cooperation raising of the young by mothers.[9][8] For the first three weeks, young remain near the den. At this time the relationship among pups is characterized by low inter-individual distance, frequent body contact, huddling, allogrooming and extended play.[3] After this, the young are able to leave the den and graze with their parents. Young are weaned after 13 weeks.

Maras will make a number of vocalizations during grazing or slow locomotion.[9] When seeking contact, a mara will emit an inflected wheet while a low repetitive grunt is made when following a conspecific.[11] Mara tooth chatter and emit low grunts when threatened. They also produce" a series of short grunt when grooming.[11] Scent marking is used by maras for complex and intense social interactions.[10] Maras will starch and stiff the soil before sitting upright with an arched back and the anogenital area flattened to the ground, [12] a process known as anal digging.[3] In addition a male will stand his hind legs and urinate on a female’s rump to which the female will respond by spraying a jet of urine backwards into the face of the male.[9] The male’s urination is met to repel other males from his partner while the female’s urination is a rejection of any approaching male when she is not respective.[9] Both anal digging and urination are more frequent during the breeding season and are more commonly done by males.[12]

Status

The Patagonian mara is considered to be a near threatened species.[1] Historically, maras have ranged from north-central Argentina south almost to Tierra de Fuego.[13] Nevertheless, maras have been greatly affected by hunting and habitat alteration and have become extinct in some areas including Buenos Aires Province.[3] The skins of maras have been used for bedspreads and rugs.[3]

References

  1. ^ a b Ojeda, R. & Pardinas, U. (2008). Dolichotis patagonum. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 5 January 2009.
  2. ^ a b Smythe. N., (1970) "On the existence of pursuit invitation signals in mammals". American Naturalist 104(938): 491-94.
  3. ^ a b c d e f g h i j k l Campos. C. M., Tognelli. M. F., Ojeda. R. A., (2001) Dolichotis patagonu, Mammalian Species, 625:1-5
  4. ^ Redford, K. H., and J. F. Eisenberg, (1992) Mammals of the Neotropics: The Southern Cone, University of Chicago Press.
  5. ^ a b Claudia M. Campos and Ricardo A. Ojeda, (1997) "Dispersal and germination of Prosopis flexuosa (Fabaceae) seeds by desert mammals in Argentina", Journal of Arid Environments, 35(4):707-14.
  6. ^ a b c d e f g h i j TABER AB (1987) "The Behavioural Ecology of the Mara". Dolichotis patagonum. Ph. D. Thesis, Belliol College, University of Oxford, Oxford, United Kingdom.
  7. ^ a b c Kufner. M. B. (1995) "Temporal activity of the mara (Dolichotis patagonum) in the Monte Desert, Argentina". Studies on Neotropical Fauna and Environment 30:37–43.
  8. ^ a b c d Taber, A. B. and Macdonald, D. W. (1992), "Communal breeding in the mara, Dolichotis patagonum ". Journal of Zoology, 227: 439–452.
  9. ^ a b c d e f g h Genest, H. & Dubost, G. (1974) "Pair living in the mara ( Dolichotis paragonum Z ) " Mammalia 38: 155-162.
  10. ^ a b U. GANSLOSSER, S. WEHNELT (1997) "Juvenile development as part of the extraordinary social system of the Dolichotis patagonum (Rodentia: Caviidae) ", Mammalia 61:3-15.
  11. ^ a b Eisenberg, J. F. (1974). "The function and motivational basis of hystricomorph vocalizations". Symp. Zool. Soc. London 34, 211-248.
  12. ^ a b TABER, B. E., AND D. W. MACDONALD. (1984) "Scent dispersing papillae and associated behaviour in the mara, Dolichotis patagonum (Rodentia: Caviomorpha) ". Journal of Zoology 203:298-301.
  13. ^ Rood. J. P. (1972) Ecological and behavioral comparisons of three genera of Argentine cavies, Animal Behavior Monographs 5:1-83.
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