鬃毛吼猴：修订间差异

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鬃毛吼猴[1]

保护状况
科學分類
界： 動物界 Animalia
門： 脊索動物門 Chordata
綱： 哺乳動物綱 Mammalia
目： 靈長目 Primates
科： 蜘蛛猴科 Atelidae
屬： 吼猴屬 Alouatta
種： 鬃毛吼猴 A. palliata
二名法
Alouatta palliata
(Gray, 1849)
鬃毛吼猴的分佈[3]
鬃毛吼猴的分佈[3]
亞種
  • A. p. aequatorialis
  • A. p. mexicana
  • A. p. palliata
異名
  • A. albulus
  • A. inclamax
  • A. inconsonans
  • A. matagalpae
  • A. niger
  • A. quichua

鬃毛吼猴（Alouatta palliata），又名長毛吼猴，是中美洲及南美洲的一種吼猴。牠們是中美洲最常見及聽到的猴子之一。牠們的兩側有很長的護毛，故有此名。[4]

鬃毛吼猴是中美洲最大的猴子之一，約有黑掌蜘蛛猴的大小，雄性可以重達9.8公斤。牠們主要吃葉子，有很多構造都適合這種食性。由於葉子很難消化及比其他食物提供較少能量，牠們很多時都是在休息及睡覺。雄性吼猴的舌骨很大，可以擴大叫聲。牠們的吼叫可以幫助互相定位，免卻勞動而浪費能量或面臨不必要的威脅。

鬃毛吼猴的群族可以有超過40個成員。牠們達至性成熟時就會被趕出群族，而群族內的成年吼猴很多時都沒有關係。至上的雄性會享有選擇食物及休息處的權力，且能與大部份的雌性交配。牠們對於雨林的生態甚為重要，因為牠們可以幫助散佈種子及發芽。雖然牠們受到伐林的影響，但仍為適應有限空間的環境。

特徵

正在休息的鬃毛吼猴。

The Mantled Howler's appearance is similar to other howler monkeys of the genus Alouatta except for coloration. The Mantled Howler is primarily black except for a fringe of gold to buff hair on each side that gives it its common name.[5] When the males reach maturity, the scrotum turns white.[5] Females are between 481、632 mm（18.9、24.9英寸） in body length, excluding tail, and males are between 508、675 mm（20.0、26.6英寸）. The prehensile tail is between 545、655 mm（21.5、25.8英寸） long. Adult females generally weigh between 3.1、7.6公斤（6.8、16.8磅）, while males typically weigh between 4.5、9.8公斤（9.9、21.6磅）.[5] Average body weights can vary significantly between monkey populations in different locations.[6] The brain of an adult Mantled Howler is about 55.1 g（1.94 oz）.[5]

The Mantled Howler shares several adaptations with other species of howler monkey that allow it to pursue a folivorous diet, that is, a diet with a large component of leaves. Its molars have high shearing crests, to help it eat the leaves,[7] and males have an enlarged hyoid bone near the vocal chords.[8] This hyoid bone amplifies the male Mantled Howler's calls, allowing it to locate other males without expending much energy.[8]

分佈

The Mantled Howler is native to Colombia, Costa Rica, Ecuador, Guatemala, Honduras, Mexico, Nicaragua and Panama.[3] Its range includes most of Honduras, Nicaragua, Costa Rica and Panama.[3] In Colombia and Ecuador, it is found in a narrow corridor bordered by the Pacific Ocean to the west and the Andes Mountains to the east.[3] In Colombia it is also found in a small area near the Caribbean Sea close to the Panama border.[3] In Guatemala, the Mantled Howler is found through the central part of the country, and into southeastern Mexico south of the Yucatán Peninsula.[3] The Mantled Howler is among the most commonly seen and heard primates in many Central American national parks, including Manuel Antonio, Corcovado, Monteverde and Soberania.[9][10] The Mantled Howler lives in several different types of forest, including secondary forest, semi-deciduous forest and riverine forest but is found in higher densities in older areas of forest and in areas containing evergreen forest.[11][12] The Mantled Howler is sympatric with another howler monkey species, the Guatemalan Black Howler, A. pigra, over a small part of its range, in Guatemala and Mexico near the Yucatan Peninsula.[3]

行為

社會結構

The Mantled Howler lives in groups. Group size usually ranges from 10 to 20 members, generally 1 to 3 adult males and 5 to 10 adult females, but some groups have over 40 members.[10][7] Males outrank females, and younger animals of each gender generally have a higher rank than older animals.[8] Higher ranking animals get preference for food and resting sites, and the alpha male gets primary mating rights.[8] Animals in the group are generally not related to each other because most members of both sexes leave the group before sexual maturity.[7]

Mature male in Alajuela Province, Costa Rica, showing prehensile tail and white scrotum

Grooming activity in the Mantled Howler is infrequent and has been shown to reflect social hierarchy, with dominant individuals grooming subordinates.[13][14] Aggressive interactions between group members is not often observed either.[13] However, studies have shown that aggressive interactions among group members do occur, and are probably not often observed because these interactions tend to be quick and silent.[13]

Mantled Howler groups that have been studied have occupied home ranges of between 10和60公頃（25和148英畝）.[7] Groups do not defend exclusive territories, but rather several groups have overlapping home ranges.[8] However, if two groups meet each group will aggressively attempt to evict the other.[8] On average, groups travel up to about 750米（2,460英尺） each day.[7]

The Mantled Howler has little interaction with other sympatric monkey species but interactions with the White-headed Capuchin sometimes occur. These are most often aggressive, and the smaller capuchins are more often the aggressors.[15] However, affiliative associations between the capuchins and howlers do sometimes occur, mostly involving juveniles playing together.[15] Other times the capuchins and howlers may feed in the same tree, apparently ignoring each other.[15]

食性

The Mantled Howler is the most folivorous species of Central American monkey. It relies on leaves for much of its diet. Leaves make up between almost 50% and 75% of the Mantled Howler's diet.[7][10] The Mantled Howler is selective about the trees it eats from, and it prefers young leaves to mature leaves.[16] This selectivity is likely to reduce the levels of toxins ingested, since certain leaves of different species contain toxin levels. Young leaves generally have less toxins as well as more nutrients than more mature leaves, and are also usually easier to digest.[16][7] Leaves from Ficus trees tend to be preferred over other leaves.[17][8] Although leaves are abundant, they are a low energy food source.[8] The fact that the Mantled Howler relies so heavily on a low energy food source drives much of its behaviour - for example, howling to locate other groups and spending a large portion of the day resting.[8]

Female in a Costa Rican Pacific Dry Forest in a typical resting position

Although leaves tend to make up the majority of the Mantled Howler's diet, fruit can also make up a large portion of the diet. When available, the proportion of fruit in the diet can be as much as 50%, and can sometimes exceed the proportion of leaves.[7] Figs tend to be a preferred fruit.[8] Flowers can also make up a significant portion of the Mantled Howler's diet and are eaten in particularly significant quantities during the dry season.[7][8] The Mantled Howler tends to get the water it needs from its food, but it does sometimes drink from tree holes.[8] It drinks from tree holes more during the wet season than during the dry season.[8] It probably also drinks water trapped in bromeliads.[17]

Like other species of howler monkeys, almost all individual Mantled Howlers have full three color vision.[18] This is different from other types of New World monkeys, in which most individuals have two color vision. The three color vision exhibited by the Mantled Howler is believed to be related to its dietary preferences, allowing it to distinguish young leaves, which tend to be more reddish, from more mature leaves.[18]

運動

The Mantled Howler is diurnal and arboreal.[5] Movement within the rainforest canopy and floor includes quadrupedalism (walking and running on supports), bridging (crossing gaps by stretching), and climbing.[19] It will also sometimes leap to get to another limb.[20]

However, the Mantled Howler is a relatively inactive monkey. It sleeps or rests the entire night and about three quarters of the day. Most of the active period is spent feeding, with only about 4% of the day spent on social interaction.[20] This lethargy is an adaptation to its low energy diet.[8] It uses its prehensile tail to grasp a branch when sleeping, resting or when feeding.[21] It can support its entire body weight with the tail, but more often holds on by the tail and both feet.[21] A study has shown that the Mantled Howler reuses travel routes to known feeding and resting sites, and appears to remember and use particular landmarks to help pick direct routes to its destination.[22]

溝通

Male Mantled Howler howling in the trees

The Mantled Howler gets the name "howler" from the calls made by the males, particularly at dawn and dusk, but also in response to disturbances.[8] These calls are very loud and can be heard for several kilometers.[8] The calls consist of grunts and repeated roars that can last for four to five seconds each.[8] The volume is produced by the hyoid bone — a hollow bone near the vocal chords — amplifying the sound made by the vocal chords. Male Mantled Howlers have hyoid bones that are 25 times larger than similarly-sized spider monkeys, and this allows the bone to act like the body of a drum in amplifying the calls. Females also call but their calls are higher in pitch and not as loud as the males'.[8] The ability to produce these loud roars is likely an energy saving device, consistent with the Mantled Howler's low energy diet. The roars allow the monkey to locate each other without moving around or risking physical confrontations.[8] The Mantled Howler utilises a wide range of other vocalisations. These include barks, grunts, woofs, cackles and screeches.[17] It uses clucking sounds to maintain auditory contact with other members of its group.[23]

Male Mantled Howler breaking a branch

The Mantled Howler also uses non-vocal communication. It engages in "urine rubbing" when in a distressful social situation.[17] This consists of rubbing the hands, feet, tail and/or chest with urine.[17] It marks its scent by rubbing its throat on branches.[5] Lip smacking and tongue movements are signals used by females to indicate a invitation to mate.[17] Genital displays are used to indicate emotional states, and group members shake branches, which is apparently a playful activity.[17]

The Mantled Howler is often indifferent to the presence of humans. However, when it is disturbed by people, it often express its irritation by urinating or defecating on them. It can accurately hit its observers despite being high in the trees.[8]

使用工具

The Mantled Howler has not been observed using tools, and prior to 1997 no howler monkey was known to use tools. However, in 1997 a Venezuelan Red Howler (Alouatta seniculus) was reported to use a stick as a club to hit a Linnaeus's Two-toed Sloth, (Choloepus didactylus), that was resting in its tree.[24] This opens the possibility that other howlers, such as the Mantled Howler, may also use tools in ways that have not yet been observed.

繁殖

Juvenile in a Costa Rican Pacific Dry Forest

The Mantled Howler uses a polygamous mating system in which one male mates with multiple females.[17] Usually, the alpha male monopolises the breeding opportunities, but if the alpha male is distracted, a lower ranking male can get an opportunity to mate.[8] And in some groups, lower ranking males do get regular mating opportunities and do sire offspring.[7][25] Alpha males generally maintain their status for about 2½ to 3 years, during which time they may father 18 or so infants.[5] Adults invite mating by flicking their tongues.[17][8] Females apparently also use chemical signals, since males smell the females' genitals and taste their urine.[17]

The gestational period is 186 days.[17] Births can occur at any time of year.[17] The infant's fur is silver at birth, but turns pale or gold after a few days. After that, the fur starts to darken, and the infant takes on the adult coloration at about 3 months old.[8]

The infant is carried under its mother, clinging to its mother's chest, for the first 2 or 3 weeks of its life.[17] After that, it is carried on its mother's back.[17] At about 3 months the mother will usually start to push the infant off, but will still carry the infant some of the time until it is 4 or 5 months old.[8] After the young can move on its own, the mother will carry it across difficult gaps in the trees.[17] Juveniles play among themselves much of the time.[17] Infants are weaned at 1½ years old at which point maternal care ends.[17] Adult females typically give birth every 19 to 23 months, assuming the prior infant survived to weaning.[7]

Females become sexually mature at 36 months, and males become sexually mature at 42 months.[17] The Mantled Howler differs from other howler monkey species in that the males' testes do not descend until they reach sexual maturity.[26] Upon reaching sexual maturity, the young monkeys are usually evicted from their natal group, although the offspring of a high ranking female may get to stay in its natal group.[8] However, many infants do not reach sexual maturity; high ranking adults sometimes harass or kill the offspring of lower ranking monkeys to eliminate competition for their own offspring.[8] When a male from outside the group ousts the previous alpha male, he normally kills any infants so that the mothers come into estrus quickly and are able to mate with him.[7] Predators such as cats, weasels, snakes and eagles also kill infants.[8] As a result, only about 30% of Mantled Howler infants live more than one year.[4] If it survives infancy, the Mantled Howler is able to live to at least 25 years old.[4]

保育狀況

The Mantled Howler is regarded as "least concern" from a conservation standpoint by IUCN. Nonetheless, its numbers may be adversely impacted by rainforest fragmentation which has caused forced relocation of groups to less habitable regions. However, the Mantled Howler can adapt to forest fragmentation better than other species due to its low energy lifestyle, small home ranges and ability to exploit widely available food sources.[27] The Mantled Howler is important to its ecosystems for a number of reasons, but especially in its capacity as a seed disperser and germinator, since passing through the monkey's digestive tract appears to aid the germination of certain seeds. Dung beetles, which are also seed dispersers as well as nutrient recyclers, also appear to be dependent on the presence of the Mantled Howler.[8] The Mantled Howler is protected from international trade under Appendix I of the Convention on International Trade in Endangered Species (CITES).[28]

分類

Mantled Howler in a wildlife sanctuary, Gulf of Dulce, Costa Rica

The Mantled Howler belongs to the New World monkey family Atelidae, the family that contains the howler monkeys, spider monkeys, woolly monkeys and muriquis. It is a member of the subfamily Alouattinae and genus Alouatta, the subfamily and genus containing all the howler monkeys.[1][3]

Three subspecies are recognized:[3]

Two additional subspecies of the Mantled Howler are sometimes recognised, but these are more generally recognised as subspecies of the Coiba Island Howler, Allouatta coibensis. However, mitochondrial DNA testing of their status has been inconclusive:[3]

參考

  1. ^ 1.0 1.1 Groves, C.P. Wilson, D.E. & Reeder, D.M. , 编. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore, Maryland: Johns Hopkins University Press. 2005: 149. ISBN 978-0-8018-8221-0. LCCN 2005001870. OCLC 62265494. OL 3392515M.  NLC 001238428.
  2. ^ Cuarón, A.D., Shedden, A., Rodríguez-Luna, E., de Grammont, P.C., Link, A., Palacions, E. & Morales, A. Alouatta palliata. The IUCN Red List of Threatened Species 2008. [7 October 2008]. 
  3. ^ 3.00 3.01 3.02 3.03 3.04 3.05 3.06 3.07 3.08 3.09 Rylands, A., Groves, C., Mittermeier, R., Cortes-Ortiz, L., & Hines, J. Taxonomy and Distributions of Mesoamerican Primates. Estrada, A., Garber, P., Pavelka, M. & Luecke, L. (编). New Perspectives in the Study of Mesoamerican Primates. New York: Springer. 2006: 47–55. ISBN 978-0-387-25854-6. 
  4. ^ 4.0 4.1 4.2 Henderson, C. Field Guide to the Wildlife of Costa Rica. Austin, Tex.: Univ. of Texas Press. 2000: 450–452. ISBN 0-292-73459-X. 
  5. ^ 5.0 5.1 5.2 5.3 5.4 5.5 5.6 Rowe, N. The Pictorial Guide to the Living Primates. East Hampton, N.Y.: Pogonias Press. 1996: 109. ISBN 0-9648825-0-7. 
  6. ^ Glander, K. Average Body Weight for Mantled Howling Monkeys (Alouatta palliata). Estrada, A.; Garber, P.A.; Pavelka, M.S.M.; Luecke, L. (编). New Perspectives in the Study of Mesoamerican Primates. New York: Springer. 2006: 247–259. ISBN 978-0-387-25854-6. 
  7. ^ 7.00 7.01 7.02 7.03 7.04 7.05 7.06 7.07 7.08 7.09 7.10 7.11 Di Fiore, A. and Campbell, C. The Atelines. Campbell, C., Fuentes, A., MacKinnon, K., Panger, M., & Bearder, S. (编). Primates in Perspective. New York: Oxford University Press. 2007: 155–177. ISBN 978-0-19-517133-4. 
  8. ^ 8.00 8.01 8.02 8.03 8.04 8.05 8.06 8.07 8.08 8.09 8.10 8.11 8.12 8.13 8.14 8.15 8.16 8.17 8.18 8.19 8.20 8.21 8.22 8.23 8.24 8.25 8.26 8.27 Wainwright, M. The Natural History of Costa Rican Mammals. Miami, FL: Zona Tropical. 2002: 139–145. ISBN 0-9705678-1-2. 
  9. ^ Hunter, L. & Andrew, D. Watching Wildlife Central America. Footscray, Vic.: Lonely Planet Publications. 2002: 97, 100, 102, 130. ISBN 1-86450-034-4. 
  10. ^ 10.0 10.1 10.2 Reid, F. A Field Guide to the Mammals of Central America and Southeast Mexico. Oxford University Press, Inc. 1998: 178–179. ISBN 0195064011. 
  11. ^ Emmons, L. Neotropical Rainforest Mammals A Field Guide Second Edition. Chicago, Ill. ;London: Univ. of Chicago Pr. 1997: 130–131. ISBN 0-226-20721-8. 
  12. ^ DeGama, H. and Fedigan, L. The Effects of Forest Fragment Age, Isolation, Size, Habitat Type, and Water Availability on Monkey Density in a Tropical Dry Forest. Estrada, A.; Garber, P.A.; Pavelka, M.S.M.; Luecke, L. (编). New Perspectives in the Study of Mesoamerican Primates. New York: Springer. 2006: 165–186. ISBN 978-0-387-25854-6. 
  13. ^ 13.0 13.1 13.2 Sussman, R. Primate Ecology and Social Structure Volume 2: New World Monkeys Revised First Edition. Boston, MA: Pearson Custom Publ. 2003: 142–146. ISBN 0-536-74364-9. 
  14. ^ Jones, C. Grooming in the Mantled Howler Monkey, Alouatta palliata (Gray). Primates. 1979., 20 (2): 289–292. doi:10.1007/BF02373380. 
  15. ^ 15.0 15.1 15.2 Rose, L., Perry, S., Panger, M., Jack, K., Manson, J., Gros-Louis, J., and Mackinnin, K. Interspecific Interactions between Cebus capucinus and other Species: Data from Three Costa Rican Sites (PDF). International Journal of Primatology. 2003, 24 (4): 780–785 [2009-02-15].  已忽略未知参数|month=（建议使用|date=） (帮助)
  16. ^ 16.0 16.1 Glander, K. Poison in a Monkey's Garden of Eden. The Primate Anthology. Upper Saddle River, N.J.: Prentice Hall. 1977: 146–152. ISBN 0-13-613845-4. 
  17. ^ 17.00 17.01 17.02 17.03 17.04 17.05 17.06 17.07 17.08 17.09 17.10 17.11 17.12 17.13 17.14 17.15 17.16 17.17 Defler, T. Primates of Colombia. Bogotá, D.C., Colombia: Conservation International. 2004: 370–384. ISBN 1-881-17383-6. 
  18. ^ 18.0 18.1 Stoner, K., Riba-Hernandez, P., & Lucas, P. Comparitive Use of Color Vision for Frugivory by Sympatric Species of Platyrrhines (PDF). American Journal of Primatology. 2005, 67: 399–409 [2009-02-15]. doi:10.1002/ajp. 
  19. ^ Gebo, D. Locomotor and Postural Behavior in Alouatta palliata and Cebus capucinus. American Journal of Primatology. 1992, 26: 277–290. doi:10.1002/ajp.1350260405. 
  20. ^ 20.0 20.1 Nowak, R. Walker's Primates of the World. Baltimore: Johns Hopkins University Press. 1999: 103–105. ISBN 0-8018-6251-5. 
  21. ^ 21.0 21.1 Kavanagh, M. A Complete Guide to Monkeys, Apes and Other Primates. London: Cape. 1983: 95–98. ISBN 0-224-02168-0. 
  22. ^ Garber, P. and Jelink, P. Travel Patterns and Spatial Mapping. Estrada, A.; Garber, P.A.; Pavelka, M.S.M.; Luecke, L. (编). New Perspectives in the Study of Mesoamerican Primates. New York: Springer. 2006: 287–306. ISBN 978-0-387-25854-6. 
  23. ^ da Cunha, R.G.T. & Byrne, R. The Use of Vocal Communication in Keeping the Spatial Cohesion of Groups: Intentionality and Specific Functions. Garber, P., Estrada, A. Bicca-Marques, J.C., Heymann, E., Strier, K (编). South American Primates. Springer. 2009: 344–345. ISBN 978-0-387-78704-6. 
  24. ^ Richard-Hansen, C., Bello, N., & Vie, J. Tool use by a red howler monkey (Alouatta seniculus) towards a two-toed sloth (Choloepus didactylus). Primates. 1998, 39 (4): 545–548 [2009-02-15]. doi:10.1007/BF02557575.  已忽略未知参数|month=（建议使用|date=） (帮助)
  25. ^ Di Fiore, A. Genetic Approaches to the Study of Dispersal and Kinship in New World Primates. Garber, P., Estrada, A., Bicca-Marques, J.C., Heymann, E. & Strier, K. (编). South American Primates: Comparative Perspectives in the Study of Behavior, Ecology and Conservation. Springer. 2009: 223–225. ISBN 978-0-387-78704-6. 
  26. ^ Kinzey, W. Alouatta. Kinzey, W. (编). New World Primates Ecology, Evolution and Behavior. New York: Aldine de Gruyter. 1997: 184. ISBN 0-202-01186-0. 
  27. ^ Garber, P., Estrada, A., and Pavelka, M. Concluding Comments and Conservation Priorities. Estrada, A., Garber, P., Pavelka, M. & Luecke, L. (编). New Perspectives in the Study of Mesoamerican Primates. New York: Springer. 2006: 570–571. ISBN 0-387-25854-X. 
  28. ^ Alphabetical Primate CITES Appendix I. AESOP-Project. [2009-02-10]. 

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