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Muntiacusmuntjacs

By Petey Maxwell

Diversity

Muntjacs are a genus of deer known for their characteristic bark, which is similar to that of canids. Many muntjacs have a set of elongated canine teeth used for intraspecific combat that protrude below their upper lip. Because of the wide karyotypic range of muntjacs, from 2n=6 (M. muntjak) to 2n=46 (M. reevesi), muntjacs have become an area of great scientific interest. (Atkins, 2004; Li, et al., 2017)

Muntiacus is a genus of family Cervidae and subfamily Cervinae. There is dispute over the exact taxonomy of the genus. Many taxonomic references maintain that there are 11 described species and 14 described subspecies. These species include Muntiacus atherodes, M. crinifrons, M. feae, M. gongshanesis, M. muntjak, M. puhoatensis, M. putaoensis, M. reevesi, M. rooseveltorum, M. truongsonensis, and M. vuquangensis and the subspecies include M. muntjak muntjak, M. muntjak annamensis, M. muntjak aureus, M. muntjak curvostylis, M. muntjak guangdongensis, M. muntjak malabaricus, M. muntjak mengalis, M. muntjak montanus, M. muntjak nigripes, M. muntjak vaginalis, M. muntjak yunnanensis, M. reevesi reevesi, M. reevesi jiangkouensis, and M. reevesi micrurus. The International Union for the Conservation of Nature (IUCN) maintains that M. muntjak montanus and M. muntjak vaginalis are not conspecific with M. muntjak, and should instead be classified as separate species: M. montanus and M. vaginalis. However, the IUCN acknowledges that these specific splits rest on little evidence. (Atkins, 2004; Grubb, 2012; Orrell and Nicolson, 2020; Timmins, et al., 2016a; Timmins, et al., 2016b; Wilson and Reeder, 2005)

Geographic Range

Muntjacs are native to the Eastern, Southeastern, and Southern regions of Asia. Introductions of M. reevesi have resulted in non-native extant ranges in the United Kingdom, Ireland, Belgium, the Netherlands, and Japan. M. reevesi is classified as an invasive species in the United Kingdom, Ireland, Belgium, and Japan and is being monitored for ecosystem risk and economic damage in the Netherlands. M. reevesi was also introduced to France but is now extinct there. (Atkins, 2004; Branquart, et al., 2016; Francis, 2019; Freeman, et al., 2016; Hollander, 2015; "Invasive species of Japan", 2012; Putman, 2019; Timmins and Chan, 2016a)

Habitat

Muntjacs are found in a variety of habitats, including plains, hills, rugged mountains, grasslands, croplands, dense forests, open forests, evergreen areas, and deciduous areas. Fragmented and uncovered canopy sites are preferred for foraging, while covered canopy sites are preferred for bed sites. Muntjacs live from sea level to 3500 m above sea level. M. reevesi is a highly adaptable species, occurring in habitats that range from temperate forests with occasional snowfall to dense forests in the warm subtropical zone, which may explain its propensity to become invasive. (Brodie and Brockelman, 2009; Teng, et al., 2004; Timmins and Chan, 2016b; Timmins and Chan, 2016a; Timmins and Duckworth, 2016a; Timmins and Duckworth, 2016b; Timmins and Duckworth, 2016c; Timmins and Duckworth, 2016d; Timmins and Duckworth, 2016e; Timmins, et al., 2016a; Timmins, et al., 2016c; Timmins, et al., 2016b; Timmins, et al., 2016d; Timmins, et al., 2016e; Timmins, et al., 2016f)

Systematic and Taxonomic History

Muntiacus is a nomen nudum, but it was conserved by Opinion 460 of the International Commission on Zoological Nomenclature. Elaphodus is the only sister genus to Muntiacus within the subfamily Cervinae. Caninmuntiacus, Cervulus, Cervus, Megamuntiacus, Muntjaccus, Muntjacus, Procops, Prox, and Stylocerus are synonyms to Muntiacus. (Atkins, 2004; Bubenik and Bubenik, 1990; "Muntjaccus Rafinesque, 1815 in GBIF Secretariat", 2019; Timmins and Chan, 2016a; Timmins and Duckworth, 2016a; Timmins, et al., 2016b; Wilson and Reeder, 2005)

Muntjacs evolved in the mid or late Pleistocene. Their elongated canines, elongated faces, small antlers, and small body size indicate that they are among the most primitive of living deer. Morphology demonstrates that muntjacs likely evolved from Elaphodus cephalophus (the only extant species of Elaphodus) and that the most primitive muntjacs were most similar to the extant M. atherodes. (Atkins, 2004; Bubenik and Bubenik, 1990; Ma, et al., 1986)

Synapomorphies that diagnose muntjacs are a chromosome number of 46 at most; antlers that are at least 20 mm long; long pedicles that are at least 7 mm thick, extend onto the face towards the orbital rims as thick ribs, and end halfway along the orbital rims; ears that are narrower and more pointed than those of Elaphodus; preorbital fossa that are smaller and less defined than those of Elaphodus; short lateral hoofs that occasionally reach 12 mm; and preorbital fissures that are well-developed. (Bubenik and Bubenik, 1990)

  • Synonyms
    • Caninmuntiacus
    • Cervulus
    • Cervus
    • Megamuntiacus
    • Muntjaccus
    • Muntjacus
    • Procops
    • Prox
    • Stylocerus
  • Synapomorphies
    • 46 chromosomes at most
    • Antlers at least 20 mm long
    • Long pedicles at least 7 mm thick, extending onto face towards orbital rims as thick ribs and ending halfway
    • Narrower and more pointed ears than Elaphodus
    • Preorbital fossa smaller and less defined than Elaphodus
    • Short lateral hoofs occasionally reaching 12 mm
    • Well-developed preorbital fissure

Physical Description

Muntjacs are small deer ranging from 60-160 cm in head and body length, 40-80 cm in shoulder height, 6-24 cm in tail length, and 10-50 kg in weight and are covered in fur that can be a mixture of dark brown, yellowish brown, grayish brown, or reddish brown. Some muntjacs have white or cream markings on their heads or bodies. Both sexes have bony, hair-covered pedicles that range from 2-13 cm in length, and males have short antlers 2.5-5.2 cm in length at the end of their pedicles. Antlers of most muntjacs have a single prong, though some are two-tined, and some species shed their antlers annually. Though more pronounced in males, both sexes have elongated canine teeth protruding below the upper lip that are no longer than a few centimeters. Additionally, muntjacs have a plesiometacarpal ankle structure and marked facial elongation. (Atkins, 2004; Bubenik and Bubenik, 1990; Huffman, 2004a; Huffman, 2004b; "Microlivestock: Little-Known Small Animals with a Promising Economic Future", 1991)

Reproduction

Muntjacs have a polygamous mating system. During courtship, males emit a soft buzzing sound and females whine while lying down flat and weaving their heads. Little other is known about muntjac mating systems. It was previously thought that muntjacs were highly territorial and that males competed for a harem of females, but recent findings showing large home-range overlap and solidarity of females suggest otherwise. (Atkins, 2004; Mccullough, et al., 2000; Odden and Wegge, 2007)

Muntjacs reach sexual maturity at 6-12 months. Gestation period is around seven months, and females bear often one and rarely two fawns. At birth, fawns weigh 0.5-1 kg, and mothers can mate again within days of giving birth. Muntjacs do not have a specific mating season. Females are polyestrous with an estrous cycle of 14–21 days and an estrous period of two days. (Atkins, 2004; Huffman, 2004b; Huffman, 2004a; "Microlivestock: Little-Known Small Animals with a Promising Economic Future", 1991)

After birth, muntjac fawns stay hidden in heavily vegetated areas until they can travel with their mothers. They remain with their mothers for around two months, at which point they are weaned. Males are not involved in the parental care of offspring. (Atkins, 2004; Huffman, 2004a; Huffman, 2004b; "Microlivestock: Little-Known Small Animals with a Promising Economic Future", 1991)

  • Parental Investment
  • altricial
  • female parental care
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

Muntjacs live up to 17 years in the wild and 23.2 years in captivity. (Huffman, 2004b; "Mammals", 2002; Tacutu, et al., 2018)

Behavior

Muntjacs are solitary deer and are active all hours of the day. They have large, overlapping home ranges with no seasonal variation and are mostly found in the cores of their home range. Though muntjacs are not strictly territorial, they do exhibit site-specific dominance. The social hierarchy is not well understood, but body mass is a good predictor of home range size. Muntjacs secrete pheromones from their preorbital glands, scrape their hooves against the ground, and scrape bark with their lower incisors and antlers to mark their home ranges. (Atkins, 2004; Huffman, 2004a; Huffman, 2004b; Mccullough, et al., 2000; Odden and Wegge, 2007)

Male muntjacs engage in intraspecific sparring and fighting. Sparring often occurs between subordinate or unequal partners and is the safer form of combat. On the other hand, fighting is dangerous and can leave both participants injured. Muntjacs exhibit a dominance display before fighting, in which they stand at least 2 m apart, hold their heads high and tilted towards their opponent, open their preorbital glands, hook their tails upwards and sideways, grind their molar teeth (producing a clicking sound), and take deliberate steps towards their opponent. At this point, either a subordinate male withdraws, or the muntjacs circle briefly before dropping their antlers and lunging forward. The muntjacs meet head-on and try to push their opponent backwards. While engaged, they try to twist the other's neck with the intent of getting them off balance. When an opponent is off balance, the aggressor attempts to use his elongated canines, or tusks, to strike downward on the back of his opponent's neck. After two or three of these tusk blows, the loser runs away and is briefly chased by the winner. The reasons for sparring and fighting are not well understood. (Barrette, 1977)

Maybe the most distinguishing behavior of muntjacs is barking. Muntjacs bark either when they detect a nearby disturbance, or when they are acting in intraspecific subordination. Because some predators of muntjacs are ambush predators, the purpose of barking when disturbances are detected is believed to be an indication to the predator that they have been perceived and that additional stalking is futile. Barking bouts in these contexts often last a few minutes but can last up to several hours. Bouts happen more often and last longer when perceptive ability is reduced by vegetative obstructions or poor light. In subordinate contexts, barking is believed to reduce the potential for intraspecific aggressive encounters. ("Microlivestock: Little-Known Small Animals with a Promising Economic Future", 1991; Yahner, 1980)

Communication and Perception

Muntjacs bark when they sense danger but do not become alert in response to barks of other muntjacs. They also do not respond to chemical markings of predators (i.e. urine, musk, or feces). Muntjacs communicate socially via sex-specific noises during courtship, pheromonal and visual markings on their home ranges, visual dominance displays (in males), and barking in subordinate contexts. (Atkins, 2004; Barrette, 1977; Huffman, 2004b; Yahner, 1980)

Food Habits

Muntjacs are generalist browsers and frugivores and feed mainly on buds, leaves, grasses, shoots, flowers, herbs, forbs, ivy, bushes, bark, twigs, seeds, seed pods, and fruit. They are also concentrate selectors and prefer foods highest in nutrients and lowest in fiber. (Atkins, 2004; "Microlivestock: Little-Known Small Animals with a Promising Economic Future", 1991; Timmins and Chan, 2016b; Timmins and Duckworth, 2016a; Timmins, et al., 2016c; Timmins, et al., 2016d; Timmins, et al., 2016b; Timmins, et al., 2016e)

Predation

Muntjacs are prey to many of the large predators that inhabit Eastern, Southeastern, and Southern Asia, including golden jackals, crocodiles, dholes, leopards, tigers, and pythons. Muntjacs bark to indicate to ambush predators that they have been perceived and that additional stalking is futile. Muntjacs also yap and expose their rump patch while fleeing to surprise predators. (Huffman, 2004b; Yahner, 1980)

Ecosystem Roles

Muntjacs are major seed dispersers and are hosts to Ornithodoros indica. (Rau and Rao, 1971; Timmins, et al., 2016b)

  • Ecosystem Impact
  • disperses seeds
Commensal/Parasitic Species

Economic Importance for Humans: Positive

Muntjacs are widely used for food, clothing, and traditional medicines. In fact, they are one of the most used and desired wild meats in Southeastern and Southern Asia. Muntjacs are sought-out for scientific studies because of the wide karyotypic range across the genus. Because they do so well in captivity, muntjacs are also found in many zoos. (Atkins, 2004; Li, et al., 2017; "Microlivestock: Little-Known Small Animals with a Promising Economic Future", 1991; Timmins and Chan, 2016a; Timmins and Chan, 2016b; Timmins and Duckworth, 2016d; Timmins and Duckworth, 2016c; Timmins and Duckworth, 2016b; Timmins and Duckworth, 2016a; Timmins and Duckworth, 2016e; Timmins, et al., 2016a; Timmins, et al., 2016c; Timmins, et al., 2016d; Timmins, et al., 2016b; Timmins, et al., 2016e; Timmins, et al., 2016f)

Economic Importance for Humans: Negative

Muntjacs damage trees by tearing off bark and, especially in countries where they have been introduced, damage crops. (Atkins, 2004; Cooke and Farrell, 2001; "Invasive species of Japan", 2012)

  • Negative Impacts
  • crop pest

Conservation Status

All muntjac species with associated data have decreasing populations, though M. vuquangensis is the only species listed as critically endangered by the IUCN. Habitat destruction and hunting are the main threats to muntjac populations. (Timmins and Chan, 2016b; Timmins and Chan, 2016a; Timmins and Duckworth, 2016d; Timmins and Duckworth, 2016c; Timmins and Duckworth, 2016b; Timmins and Duckworth, 2016a; Timmins and Duckworth, 2016e; Timmins, et al., 2016a; Timmins, et al., 2016c; Timmins, et al., 2016b; Timmins, et al., 2016d; Timmins, et al., 2016e; Timmins, et al., 2016f)

Fortunately, conservation sites have been identified for all muntjac species. The two most threatened species, M. vuquangensis and M. crinifrons, are internationally managed and their trade is controlled. (Timmins and Chan, 2016b; Timmins and Chan, 2016a; Timmins and Duckworth, 2016d; Timmins and Duckworth, 2016c; Timmins and Duckworth, 2016b; Timmins and Duckworth, 2016a; Timmins and Duckworth, 2016e; Timmins, et al., 2016a; Timmins, et al., 2016c; Timmins, et al., 2016b; Timmins, et al., 2016d; Timmins, et al., 2016e; Timmins, et al., 2016f)

  • IUCN Red List [Link]
    Not Evaluated

Other Comments

The name "Muntiacus" comes from "muntjak," the native name for small deer in the Sunda language. Interestingly, M. muntjak (2n=6) and M. reevesi (2n=46) can produce viable but sterile offspring, despite the strong karyotypic differences. (Huffman, 2004a; Huffman, 2004b; "Microlivestock: Little-Known Small Animals with a Promising Economic Future", 1991)

Contributors

Petey Maxwell (author), Colorado State University, Genevieve Barnett (editor), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

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.

chemical

uses smells or other chemicals to communicate

crepuscular

active at dawn and dusk

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

folivore

an animal that mainly eats leaves.

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

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.

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

scrub forest

scrub forests develop in areas that experience dry seasons.

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sexual ornamentation

one of the sexes (usually males) has special physical structures used in courting the other sex or fighting the same sex. For example: antlers, elongated tails, special spurs.

solitary

lives alone

tactile

uses touch to communicate

taiga

Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.

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).

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

References

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Timmins, R., J. Duckworth, W. Robichaud, B. Long, T. Gray, A. Tilker. 2016. "Muntiacus vuquangensis" (On-line). The IUCN Red List of Threatened Species 2016. Accessed May 11, 2020 at https://www.iucnredlist.org/species/44703/22153828#habitat-ecology.

Timmins, R., R. Steinmetz, N. Samba Kumar, M. Anwarul Islam, H. Sagar Baral. 2016. "Muntiacus vaginalis" (On-line). The IUCN Red List of Threatened Species 2016. Accessed May 10, 2020 at https://www.iucnredlist.org/species/136551/22165292.

Timmins, R., R. Steinmetz, W. Chutipong. 2016. "Muntiacus feae" (On-line). The IUCN Red List of Threatened Species 2016. Accessed May 11, 2020 at https://www.iucnredlist.org/species/13927/22160266#habitat-ecology.

Wilson, D., D. Reeder. 2005. "GENUS Muntiacus" (On-line). Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed). Accessed May 10, 2020 at https://www.departments.bucknell.edu/biology/resources/msw3/browse.asp?s=y&id=14200400.

Yahner, R. 1980. Barking in a Primitive Ungulate, Muntiacus reevesi: Function and Adaptiveness. The American Naturalist, 116/2: 157-177.

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