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Lifespan, longevity, and ageing

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Maximum longevity: 20.9 years (captivity) Observations: One specimen lived 20.9 years in captivity (Richard Weigl 2005).
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Reproduction

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The mating system is polygynous. Males compete in fighting competition to mate with a group of females.

Mating System: polygynous

Males join the females in December after fierce battles with other males. The winner of the battle obtains the right to breed with group of 10 – 20 females. The gestation period for the ibex is approximately 147 – 180 days. A day after parturition, the young are able to walk on the rock cliffs following their mothers. The young are mature at 8 – 12 months, but don’t breed until 2 or 3 years of age. Ibex typically have one young per year, and more than one is uncommon.

Breeding interval: Alpine ibex breed once yearly.

Breeding season: Mating occurs in late fall.

Range number of offspring: 1 to 2.

Average number of offspring: 1.11.

Range gestation period: 4.9 to 6 months.

Range weaning age: 3 to 12 months.

Range age at sexual or reproductive maturity (female): 10 (low) months.

Range age at sexual or reproductive maturity (male): 10 (low) months.

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

Average birth mass: 2850 g.

Average number of offspring: 1.1.

Females provide milk for their young, as do all mammalian females. The young are precocious, and are able to follow their mothers shortly after birth.

Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); post-independence association with parents

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Untitled

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In the eighteenth century some Europeans believed ibex were magical. Today's equivalent of the magical ibex is the zodiac sign Capricorn.

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Behavior

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Perception Channels: tactile ; chemical

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Conservation Status

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Alpine ibex have sustainable populations due to successful reintroduction programs.

US Federal List: no special status

CITES: no special status

IUCN Red List of Threatened Species: least concern

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Benefits

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Ibex may compete with domestic goats (Capra hircus) for food and water.

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Benefits

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In addition to trophy hunting, there was a market for the parts of ibex believed useful in medicinal purposes.

Positive Impacts: food ; body parts are source of valuable material; source of medicine or drug

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Associations

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As a browser, this ibex probably influences the vegetational community, As a prey species, it is likely that the availablitliy of ibex affects the populations of predators.

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Trophic Strategy

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In the spring the animals migrate back into the mountains to new feeding areas. In the winter when the snow is deep and the weather is severe they migrate down to south facing slopes which have more food and less snow. These browsers and grazers become active in the afternoon and into the evening and feed through out the night in the forest, returning to the rock cliffs in the morning.

Foods commonly eaten include: grasses, forbs, leaves, shoots and bark.

Plant Foods: leaves; wood, bark, or stems

Primary Diet: herbivore (Folivore , Lignivore)

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Distribution

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Alpine ibex, Capra ibex, are found in central Europe south to northern Ethiopia and east to Central China.

Biogeographic Regions: palearctic (Native ); ethiopian (Native )

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Habitat

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Alpine ibex are mountain animals usually living at elevations up to 3,200 meters. Males stay up on the rock cliffs during the day, whereas females stay below in the rolling slopes and brushy areas. At night they will all move down into the forest for the night to feed.

Range elevation: 3200 (high) m.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: chaparral ; forest ; scrub forest ; mountains

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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Life Expectancy

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The lifespan of an alpine ibex in the wild is about 10 – 18 years. In captivity the oldest know individual was 21 years and 3 months.

Range lifespan
Status: captivity:
21.25 (high) years.

Typical lifespan
Status: wild:
10 to 18 years.

Average lifespan
Status: captivity:
22.3 years.

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Morphology

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Alpine ibex are sexually dimorphic. Males range from 65 – 105 cm in height at the shoulder and weigh about 80 - 100 kg. Shoulder heights in females are about 65 – 70 cm and weight varies from 30 – 50 kg. The length of an ibex is about 1.3 – 1.4 m long with a tail length about 120 – 150 cm. Their coats are uniformly brown to gray, with thick beards. The underside of southern alpine ibex is lighter than the northern alpine ibex. Nubian (Capra nubiana) and Walia ibex (Capra walie) are smaller than alpine ibex.

Range mass: 65 to 100 kg.

Range length: 1.3 to 1.4 m.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger; ornamentation

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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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John Sippl, University of Wisconsin-Stevens Point
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Associations

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Ibex are herding animals which are subject to a wide variety predators. Eagles, bears, leopards and humans all play significant roles in regulating the ibex population.

Known Predators:

  • golden eagles (Aquila chrysaetos)
  • bears (Ursinae)
  • wolves (Canis lupus)
  • leopards (Panthera pardus)
  • humans (Homo sapiens)
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Sippl, J. 2003. "Capra ibex" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Capra_ibex.html
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Distribution in Egypt

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Narrow (Sinai, Eastern Desert). 

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Status in Egypt

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Native, resident.

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Brief Summary

provided by EOL authors

The Alpine Ibex (Capra ibex) was at one time considered to include three subspecies: the Alpine Ibex (C. i. ibex), found in Switzerland, southern Germany, Liechtenstein, Austria, northern Italy, and southeastern France and introduced to northwestern Slovenia and Bulgaria; the broadly distributed Siberian Ibex (C. i. sibirica), found mainly in China, Central Asia, and Mongolia; and the Nubian Ibex (C. i. nubiana), found in Egypt east of the Nile River, northeastern Sudan, Israel, western Jordan, Saudi Arabia, southwestern Oman, southeastern Yemen, and possibly Eritrea. These taxa are now generally treated as three full species (C. ibex, C. sibirica, and C. nubiana).

The Alpine Ibex occurs mainly at elevations of 1600 to 3200 m in alpine and subalpine habitats, but can use open forests in rocky terrain associated with ledges, cliffs, and precipitous valleys.

This species was on the verge of extinction in the 1800s, but a protected population remained in northern Italy and animals from this population were used to re-establish the species in the Alps of Austria, Switzerland, Liechtenstein, Germany, and France. There are now over 22,000 animals in free-ranging populations and controlled hunting is allowed in Switzerland, Austria, Liechtenstein, and Slovenia.

(Valdez 2011 and references therein)

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Alpine ibex

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The Alpine ibex (Capra ibex), also known as the steinbock, bouquetin, or simply ibex, is a species of wild goat[2] that lives in the mountains of the European Alps. It is a sexually dimorphic species with larger males that carry larger, curved horns. Their coat colour is typically brownish grey. Alpine ibex tend to live in steep, rough terrain near the snow line. They are also social, although adult males and females segregate for most of the year, coming together only to mate. Four distinct groups exist; adult male groups, female-offspring groups, groups of young individuals, and mixed-sex groups.

During the breeding season, males fight for access to females and use their long horns in agonistic behaviours. After being extirpated from most areas by the 19th century, the Alpine ibex was successfully reintroduced to parts of its historical range. All individuals living today descend from the stock in Gran Paradiso National Park in Aosta Valley [3]. This national park was created to help the ibex to thrive. The ibex is the emblem of both the Gran Paradiso National Park and the Vanoise National Park. The species is currently listed as of least concern by the IUCN, but went through a population bottleneck of less than 100 individuals. This has led to very low genetic diversity across populations.[4]

Taxonomy and phylogeny

The Alpine ibex was first described by Carl Linnaeus in 1758. It is classified in the genus Capra (Latin for "goat") with at least seven other species of wild goats. Both Capra and Ovis (sheep) descended from a goral-like animal from the Miocene and early Pliocene, whose fossils are found in Kenya, China, and Slovenia. The genus Tossunnoria appears in China during the late Miocene and appears to have been intermediate between gorals and goats. Fossils of Alpine ibex date back to the late Pleistocene, when the Spanish ibex and it probably evolved from the extinct Pleistocene species Capra camburgensis.[5] The Nubian (C. nubiana), Walia (C. walie), and Siberian ibex (C. sibirica) were previously considered to be subspecies of the Alpine ibex, giving populations in the Alps the trinomial of C. i. ibex.[6]

Appearance

Video of a herd of Alpine ibex on Augstmatthorn in the Bernese Highlands, Switzerland

Compared with other members of its genus, the Alpine ibex has a short, broad head and a duller coat. It has brownish-grey hair over most of the body, a pale abdomen, and slightly darker markings on the chin and throat and in a stripe along the back. They moult twice a year, firstly in April or May, and then again in September, when they replace the short summer coat with thicker hair and a woolly undercoat.[5]

Males commonly grow to a height of 90 to 101 cm (35 to 40 in) at the withers, with a body length of 149 to 171 cm (59 to 67 in) and weigh from 67 to 117 kg (148 to 258 lb). Females are noticeably smaller, with a shoulder height of 73 to 84 cm (29 to 33 in), a body length of 121 to 141 cm (48 to 56 in), and a weight of 17 to 32 kg (37 to 71 lb). Both male and female Alpine ibex have large, backwards-curving horns with numerous ridges along their length. At 69 to 98 cm (27 to 39 in), those of the males are substantially larger than those of females, which reach only 18 to 35 cm (7.1 to 13.8 in) in length.[5]

Distribution and ecology

 src=
Ibex standing on cliff in winter

The Alpine ibex was, at one point, restricted only to the Gran Paradiso National Park in northern Italy, and in the Maurienne Valley in the French Alps [7] but it was reintroduced to most of the European Alps. Reintroductions started in 1906 into Switzerland. Alpine ibex are now found in most or all of the Italian and French alpine ranges, southern Germany, Switzerland[8] and Austria. It has also been introduced to Bulgaria and Slovenia.[1]

An excellent climber, its preferred habitat is the rocky region along the snow line above alpine forests, where it occupies steep, rough terrain at elevations of 1,800 to 3,300 m (5,900 to 10,800 ft).[9] Alpine ibex are typically absent from woodland areas,[5] although adult males in densely populated areas may stay in larch and mixed larch-spruce woodland if no snow has fallen.[10] Males spend the winter in coniferous forests.[5] For most of the year, males and females occupy different habitats.[11] Females rely on steep terrain more so than males.[12] Males use lowland meadows during the spring, which is when snow melts and green grass appears.[12] They then climb to alpine meadows during the summer.[10] When winter arrives, both sexes move to steep, rocky slopes that amass little snow.[13] They prefer slopes of 30–45° and use small caves and overhangs for shelter.[14] Home ranges are highly variable, depending on the availability of resources, and vary in size throughout the year. Figures from 180 to 2,800 ha (0.69 to 10.81 sq mi) have been recorded.[5][13] Home ranges tend to be largest during summer and autumn, smallest in winter, and intermediate in spring.[5] Female home ranges are usually smaller than those of males. Alpine ibex appear to have a low rate of predation and in Gran Paradiso typically die of age, starvation, or disease.[5]

Foraging

Alpine ibex are strictly herbivorous, with over half of their diet consisting of grasses, and the remainder being a mixture of mosses, flowers, leaves, and twigs.[5] If leaves and shoots are out of reach, they often stand on their rear legs to reach this food. Grass genera that are the most commonly eaten are Agrostis, Avena, Calamagrostis, Festuca, Phleum, Poa, Sesleria, and Trisetum.[5] The climbing ability of the Alpine ibex is such that it has been observed standing on the sheer face of the Cingino Dam in Piedmont, Italy, where it licks the stonework to obtain mineral salts.[15]

Life history

Although the Alpine ibex is a social species, they segregate sexually and spatially depending on the season.[12] Four types of groups exist: Adult male groups, female-offspring groups, groups of young individuals 2–3 years old, and mixed-sex groups.[5][16] Young groups are numerous at the beginning of summer, but are expelled by females at the end of their gestation period. Female and offspring groups occur year-round, at least in an area of the French Alps.[16] Mixed sex groups of adult males and females occur during breeding, which lasts from December to January. By April and May, the adults separate.[16] The largest aggregations of either sex occur during June and July. Gatherings of males begin to decrease during October and November, and are lowest from the rut from December to March.[16] The males then leave their separate wintering areas and gather again.[17]

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Male ibex locking horns

A linear dominance hierarchy exists among males. In small populations, which are more cohesive, males know their place in the hierarchy based on memories of past encounters,[5] while in mobile and large groups, where encounters with strangers are common, rank is based on horn size.[18] Antagonistic behavior in males can come in the form of "direct" or "indirect" aggression. With direct aggression, one male bumps another with his horns or places himself in front of his opponent. He stands on his hind legs and comes down on his opponent with his horns. This may signal that he is ready to clash or may be attempting a real clash.[5] Indirect aggression is mostly intimidation displays.[5]

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Mother with young

Reproduction and growth

The breeding season starts in December, and typically lasts around six weeks. During this time, male herds break up into smaller groups that search for females. The rut takes place in two phases. In the first phase, the male groups interact with the females that are all in oestrous. The higher the male's rank, the closer he can get to a female.[5] Males perform courtship displays. In the second phase of the rut, one male separates from his group to follow an individual female. He displays to her and guards her from other males. Before copulation, the female moves her tail and courtship becomes more intense. They copulate and then he rejoins his group and reverts to the first phase.[5] Gestation lasts around 167 days, and results in the birth of one or two kids, with twins making up about 20% of births.[19]

Alpine ibex reach sexual maturity at 18 months, but females do not reach their maximum body size for five to six years, and males not for 9-11 years. The horns grow throughout life, growing most rapidly during the second year of life, and thereafter by about 8 cm (3.1 in) a year, eventually slowing to half that rate once the animal reaches 10 years of age. Alpine ibex live for up to 19 years in the wild.[20]

Conservation status

 src=
Young Alpine ibex on a cliff

The Alpine ibex historically ranged through France, Italy, Switzerland, Liechtenstein, Bavaria, Austria and Slovenia. Starting in the early 16th century and with firearms becoming common, the overall population declined due to overexploitation and poaching. The ibex became extinct in Switzerland and Germany by the 18th century, and was extinct in Austria and northeastern Italy by the 19th century. They remained only in and around the adjacent Gran Paradiso and Vanoise Massifs, then both part of the Kingdom of Sardinia. Located in the western Italian Alps and the Maurienne valley in the north eastern French alps, bordering the Vanoise and Gran Paradiso Massif, the park was declared a royal hunting reserve in 1854 with the name of "Royal hunting reserve of Gran Paradiso" by Victor Emmanuel II, the first king of a united Italy .[21]

Ibex were protected from poaching and their numbers increased, reaching 3,020 in 1914. The ibex enjoyed further protection when Gran Paradiso was made into a national park in 1922. Animals from this population naturally dispersed into surrounding regions. However, reintroductions have been the predominant source of new populations. Today, the total population of Alpine ibex is over 30,000 and is considered to be of Least Concern by the IUCN. However, Alpine ibex do have low genetic diversity putting them at risk of inbreeding depression.[4]

In popular culture

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Zlatorog or Goldhorn beer is one of the most popular brands in Slovenia.

The Alpine ibex is a mountain icon. It is represented in many official emblems throughout the Alpine range, from France to Austria, as in the coat of arms of the Canton of Grisons in Switzerland or logos such as Pro Natura.

The animal is at the center of the Goldhorn legend. This, in turn, underlies the emblem of the Laško Brewery, the largest brewery in Slovenia. In addition, the first Slovene-language, full-length film, recorded in 1931 by Janko Ravnik, was titled In the Kingdom of the Goldhorn.

In 2009, DNA analysis of the stomach contents of the Alpine iceman known as "Ötzi" confirmed that his last meal included ibex meat.[22]

References

  1. ^ a b Aulagnier, S.; Kranz, A.; Lovari, S.; Jdeidi, T.; Masseti, M.; Nader, I.; de Smet, K. & Cuzin, F. (2008). "Capra ibex". IUCN Red List of Threatened Species. IUCN. 2008: e.T42397A10695445. doi:10.2305/IUCN.UK.2008.RLTS.T42397A10695445.en. Database entry includes a brief justification of why this species is of least concern.
  2. ^ Lovari, Sandro. Ibex. Encyclopaedia Britannica
  3. ^ Biebach, I (2009). "Inbreeding in reintroduced populations: the effects of early reintroduction history and contemporary processes". Conservation Genetics.
  4. ^ a b Biebach, I.; Keller, L. F. (2009). "A strong genetic footprint of the re-introduction history of Alpine ibex (Capra ibex ibex)". Molecular Ecology. 18 (24): 5046–58. doi:10.1111/j.1365-294X.2009.04420.x. PMID 19912536.
  5. ^ a b c d e f g h i j k l m n o p Parrini, F.; Cain III, J. W.; Krausman, P. R. (2009). "Capra ibex (Artiodactyla: Bovidae)". Mammalian Species. 830: 1–12. doi:10.1644/830.1.
  6. ^ Shackleton, D. W. (1997). Wild Sheep and Goats and Their Relatives: Status Survey and Action Plan for Caprinae. International Union for Conservation of Nature and Natural Resources. Species Survival Commission. Caprinae Specialist Group. p. 12. ISBN 2831703530.
  7. ^ Girard, Irène (2000). "Dynamique des populations et expansion geographique du bouquetin des alpes (Capra ibex ibex) dans le parc national de la vanoise". PhD thesis, Université Savoie Mont Blanc. Archived from the original on 2013-12-28. Retrieved 2013-09-07.
  8. ^ (in French) Heinz Staffelbach, Manuel des Alpes suisses. Flore, faune, roches et météorologie, Rossolis, 2009 (ISBN 978-2-940365-30-2). Also available in German: Heinz Staffelbach, Handbuch Schweizer Alpen. Pflanzen, Tiere, Gesteine und Wetter, Haupt Verlag, 2008 (ISBN 978-3-258-07638-6).
  9. ^ Parrini, F.; et al. (2003). "Spatial behaviour of adult male Alpine ibex Capra ibex ibex in the Gran Paradiso National Park, Italy". Acta Theriologica. 48 (3): 411–423. doi:10.1007/BF03194179.
  10. ^ a b Grignolio, S.; Parrini, F.; Bassano, B.; Luccarini, S.; Apollonio, M. (2003). "Habitat selection in adult males of Alpine ibex, Capra ibex ibex" (PDF). Folia Zoologica. 52 (2): 113–20. Archived from the original (PDF) on 2013-05-22. Retrieved 2012-12-26.
  11. ^ ToÏgo C.; Gaillard J. M.; Michallet J. (1997). "Adult survival pattern of the sexually dimorphic Alpine ibex (Capra ibex ibex)". Canadian Journal of Zoology. 75: 75–79. doi:10.1139/z97-009.
  12. ^ a b c Francisci, F., S. Focardi, and L. Boitani. (1985) "Male and female Alpine ibex: phenology of space use and herd size", pp. 124–133. in The biology and management of mountain ungulates. Lovari, S. Croom Helm. London, United Kingdom. ISBN 978-94-015-7346-7
  13. ^ a b Grignolio, S.; et al. (2004). "Seasonal variations of spatial behaviour in female Alpine ibex (Capra ibex ibex) in relation to climatic conditions and age". Ethology Ecology and Evolution. 16 (3): 255–264. doi:10.1080/08927014.2004.9522636.
  14. ^ Wiersema, G. (1984). "Seasonal use and quality assessment of ibex habitat". Acta Zoologica Fennica. 172: 89–90.
  15. ^ Nutkins, Terry (3 November 2010). "The goats with a head for heights". Guardian.co.uk. Guardian News and Media Limited. Retrieved 4 November 2010.
  16. ^ a b c d Villaret, J. C.; Bon, R. (1995). "Social and spatial segregation in Alpine ibex (Capra ibex) in Bargy, French Alps". Ethology. 101 (4): 291–300. doi:10.1111/j.1439-0310.1995.tb00366.x.
  17. ^ Parrini, F.; Grignolio, S.; Luccarini, S.; Bassano, B.; Apollonio, M. (2003). "Spatial behaviour of adult male Alpine ibex Capra ibex ibex in the Gran Paradiso National Park, Italy". Acta Theriologica. 48 (3): 411–23. doi:10.1007/BF03194179.
  18. ^ Schaller, G. B. (1977) Mountain monarchs: wild sheep and goats of the Himalaya. University of Chicago Press. Chicago, Illinois.
  19. ^ Stüwe, M.; Grodinsky, C. (1987). "Reproductive biology of captive Alpine ibex (Capra i. ibex)". Zoo Biology. 6 (4): 331–339. doi:10.1002/zoo.1430060407.
  20. ^ ToÏgo, C.; et al. (2007). "Sex- and age-specific survival of the highly dimorphic Alpine ibex: evidence for a conservative life-history tactic". Journal of Animal Ecology. 76 (4): 679–686. doi:10.1111/j.1365-2656.2007.01254.x. PMID 17584373.
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Alpine ibex: Brief Summary

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The Alpine ibex (Capra ibex), also known as the steinbock, bouquetin, or simply ibex, is a species of wild goat that lives in the mountains of the European Alps. It is a sexually dimorphic species with larger males that carry larger, curved horns. Their coat colour is typically brownish grey. Alpine ibex tend to live in steep, rough terrain near the snow line. They are also social, although adult males and females segregate for most of the year, coming together only to mate. Four distinct groups exist; adult male groups, female-offspring groups, groups of young individuals, and mixed-sex groups.

During the breeding season, males fight for access to females and use their long horns in agonistic behaviours. After being extirpated from most areas by the 19th century, the Alpine ibex was successfully reintroduced to parts of its historical range. All individuals living today descend from the stock in Gran Paradiso National Park in Aosta Valley . This national park was created to help the ibex to thrive. The ibex is the emblem of both the Gran Paradiso National Park and the Vanoise National Park. The species is currently listed as of least concern by the IUCN, but went through a population bottleneck of less than 100 individuals. This has led to very low genetic diversity across populations.

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