The Alpine ibex (''Capra ibex''), also known as the steinbock, bouquetin, or simply ibex, is a species of

wild goat The wild goat (''Capra aegagrus'') is a wild goat species, inhabiting forests, shrublands and rocky areas ranging from Turkey and the Caucasus in the west to Turkmenistan, Afghanistan and Pakistan in the east. It has been listed as near thre ...

that lives in the

mountain A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually higher ...

s of the

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

Alps The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, Sw ...

. It is a sexually dimorphic species: males are larger and carry longer, curved horns than females. Its coat colour is typically brownish grey. Alpine ibex tend to live in steep, rough terrain near the

snow line The climatic snow line is the boundary between a snow-covered and snow-free surface. The actual snow line may adjust seasonally, and be either significantly higher in elevation, or lower. The permanent snow line is the level above which snow wil ...

. 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 behaviour. 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 and

Piemonte it, Piemontese , population_note = , population_blank1_title = , population_blank1 = , demographics_type1 = , demographics1_footnotes = , demographics1_title1 = , demographics1_info1 = , demographics1_title2 ...

(

Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical ...

), a national park created to help the ibex thrive. The ibex is the emblem of both the Gran Paradiso National Park, and the contiguous Vanoise National Park across the French border. The species is currently listed as of least concern by the

IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...

, but went through a

population bottleneck A population bottleneck or genetic bottleneck is a sharp reduction in the size of a population due to environmental events such as famines, earthquakes, floods, fires, disease, and droughts; or human activities such as specicide, widespread violen ...

of fewer than 100 individuals during its near-extinction event. This has led to very low genetic diversity across populations.

Taxonomy and phylogeny

The Alpine ibex was first described by

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...

in 1758. It is classified in the

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

'' 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 The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

and early

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Tossunnoria'' 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 The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...

, when the Spanish ibex and it probably evolved from the extinct Pleistocene species '' Capra camburgensis''. The

Nubian Nubian may refer to: *Something of, from, or related to Nubia, a region along the Nile river in Southern Egypt and northern Sudan. *Nubian people *Nubian languages *Anglo-Nubian goat, a breed of goat * Nubian ibex * , several ships of the Britis ...

(''C. nubiana''), walia (''C. walie''), and Siberian ibexes (''C. sibirica'') were previously considered to be subspecies of the Alpine ibex, giving populations in the Alps the trinomial of ''C. i. ibex''.

Appearance

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. Males commonly grow to a height of at the

withers The withers is the ridge between the shoulder blades of an animal, typically a quadruped. In many species, it is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, cattle a ...

, with a body length of and weigh from . Females are noticeably smaller, with a shoulder height of , a body length of , and a weight of . Both male and female Alpine ibex have large, backwards-curving horns with numerous ridges along their length. At , those of the males are substantially larger than those of females, which reach only in length.

Distribution and ecology

During the Last Glacial Period the species range was reaching as far south as

Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders wi ...

, where it disappeared in the

Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...

, after 5,600 BC due to hunting, with the last known remains being found on mountains surrounding

Thessaly Thessaly ( el, Θεσσαλία, translit=Thessalía, ; ancient Thessalian: , ) is a traditional geographic and modern administrative region of Greece, comprising most of the ancient region of the same name. Before the Greek Dark Ages, Thes ...

. The Alpine ibex was, at one point, restricted only to the Gran Paradiso National Park in northern

, and the contiguous

Maurienne Maurienne ( frp, Môrièna) is one of the provinces of Savoy, corresponding to the arrondissement of Saint-Jean-de-Maurienne in France. It is also the original name of the capital of the province, now Saint-Jean-de-Maurienne. Location The Mau ...

Valley in the French Alps but it was reintroduced to most of the

. Reintroductions started in 1906 into Switzerland. Alpine ibex are now found in most or all of the Italian and French alpine ranges, southern

Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...

Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...

Heinz Staffelbach, ''Manuel des Alpes suisses. Flore, faune, roches et météorologie'', Rossolis, 2009 (). Also available in German: Heinz Staffelbach, ''Handbuch Schweizer Alpen. Pflanzen, Tiere, Gesteine und Wetter'', Haupt Verlag, 2008 (). and

Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...

. It has also been introduced to

Bulgaria Bulgaria (; bg, България, Bǎlgariya), officially the Republic of Bulgaria,, ) is a country in Southeast Europe. It is situated on the eastern flank of the Balkans, and is bordered by Romania to the north, Serbia and North Macedo ...

and

Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, and ...

. An excellent climber, its preferred

habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...

is the rocky region along the

above alpine forests, where it occupies steep, rough terrain at elevations of . Alpine ibex are typically absent from woodland areas, although adult males in densely populated areas may stay in larch and mixed larch-spruce woodland if no snow has fallen. Males spend the winter in coniferous forests. For most of the year, males and females occupy different habitats. Females rely on steep terrain more so than males.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. Males use lowland meadows during the spring, which is when snow melts and green grass appears. They then climb to alpine meadows during the summer. When winter arrives, both sexes move to steep, rocky slopes that amass little snow. They prefer slopes of 30–45° and use small caves and overhangs for shelter. Home ranges are highly variable, depending on the availability of resources, and vary in size throughout the year. Figures from have been recorded. Home ranges tend to be largest during summer and autumn, smallest in winter, and intermediate in spring. 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.

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. 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 ''Agrostis'' (bent or bentgrass) is a large and very nearly cosmopolitan genus of plants in the grass family, found in nearly all the countries in the world. It has been bred as a GMO creeping bent grass. Species * '' Agrostis aequivalvi'' ...

'', '' Avena'', '' Calamagrostis'', '' Festuca'', ''

Phleum ''Phleum'' (timothy) is a genus of annual and perennial plants in the grass family. The genus is native to Europe, Asia and north Africa, with one species (''P. alpinum'') also in North and South America. They are tufted grasses growing to 2 ...

'', '' Poa'', '' Sesleria'', and ''

Trisetum ''Trisetum'' is a genus of plants in the grass family Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses ...

''. 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 it, Piemontese , population_note = , population_blank1_title = , population_blank1 = , demographics_type1 = , demographics1_footnotes = , demographics1_title1 = , demographics1_info1 = , demographics1_title2 ...

, where it licks the stonework to obtain mineral salts.

Life history

Although the Alpine ibex is a social species, they segregate sexually and spatially depending on the season. Four types of groups exist: Adult male groups, female-offspring groups, groups of young individuals 2–3 years old, and mixed-sex groups. 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. Mixed-sex groups of adult males and females occur during breeding, which lasts from December to January. By April and May, the adults separate. 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. The males then leave their separate wintering areas and gather again. Bouquetins combat

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, while in mobile and large groups, where encounters with strangers are common, rank is based on horn size.Schaller, G. B. (1977) ''Mountain monarchs: wild sheep and goats of the Himalaya''. University of Chicago Press. Chicago, Illinois. 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. Indirect aggression is mostly intimidation displays. Alpine ibex Cima di Terrarossa 7

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. 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 Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...

, the female moves her tail and courtship becomes more intense. They copulate and then he rejoins his group and reverts to the first phase. Gestation lasts around 167 days, and results in the birth of one or two kids, with twins making up about 20% of births. 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 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.

Conservation status

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 extirpated in Switzerland and Germany by the 18th century, and was extirpated 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 en, Victor Emmanuel Maria Albert Eugene Ferdinand Thomas , house = Savoy , father = Charles Albert of Sardinia , mother = Maria Theresa of Austria , religion = Roman Catholicism , image_size = 252px , succession ...

, the first king of a united Italy. 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 A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. ...

by the

. However, Alpine ibex have low genetic diversity putting them at risk of

inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...

References

External links

Alpine Ibex European Specialists Group

{{Authority control Capra (genus) Mammals of Europe Mammals described in 1758 Taxa named by Carl Linnaeus Articles containing video clips