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 threatene ...

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 (topography), summit area, and ...

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 Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

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 Agonistic behaviour is any social behaviour related to fighting. The term has broader meaning than aggressive behaviour because it includes threats, displays, retreats, placation, and conciliation. The term "agonistic behaviour" was first implemen ...

. After being

extirpated Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...

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 Gran Paradiso National Park (Italian: ''Parco nazionale del Gran Paradiso''; ), is an Italian national park in the Graian Alps, between the Aosta Valley and Piedmont regions. The park is named after Gran Paradiso mountain, which is located in the ...

Aosta Valley , Valdostan or Valdotainian it, Valdostano (man) it, Valdostana (woman)french: Valdôtain (man)french: Valdôtaine (woman) , population_note = , population_blank1_title = Official languages , population_blank1 = Italian French ...

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 re ...

), 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 Vanoise National Park (french: Parc national de la Vanoise) is a French national park between the Tarentaise and Maurienne valleys in the French Alps, containing the Vanoise massif. It was created in 1963 as the first national park in France. ...

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, the ...

in 1758. It is classified in the

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

Capra Capra may refer to: * ''Capra'' (genus), comprising the goats * Capra (goat dance), a Romanian custom * Capra (titular see), a titular see in the Catholic Church * Capra (car), a pick-up brand from the Iranian Bahman Group People * Buzz Capra ...

'' (Latin for "goat") with at least seven other species of wild goats. Both ''Capra'' and ''

Ovis ''Ovis'' is a genus of mammals, part of the Caprinae subfamily of the ruminant family Bovidae. Its seven highly sociable species are known as sheep or ovines. Domestic sheep are members of the genus, and are thought to be descended from the wil ...

'' (sheep) descended from a

goral The gorals are four species in the genus ''Naemorhedus''. They are small ungulates with a goat-like or antelope-like appearance. Until recently, this genus also contained the serow species (now in genus '' Capricornis''). Etymology The original ...

-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 The Iberian ibex (''Capra pyrenaica''), also known as the Spanish ibex, Spanish wild goat, or Iberian wild goat, is a species of ibex endemic to the Iberian Peninsula. Four subspecies have been described; two are now extinct. The Portuguese subs ...

and it probably evolved from the extinct Pleistocene species ''

Capra camburgensis Capra may refer to: * ''Capra'' (genus), comprising the goats * Capra (goat dance), a Romanian custom * Capra (titular see), a titular see in the Catholic Church * Capra (car), a pick-up brand from the Iranian Bahman Group People * Buzz Capra ...

''. 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 ibex The Siberian ibex (''Capra sibirica''), also known as the Altai ibex, Central Asia(n) ibex, Gobi ibex, Himalayan ibex, Mongolian ibex or Tian Shan ibex, is a species of ibex that lives in central Asia. It has traditionally been treated as a subs ...

es (''C. sibirica'') were previously considered to be subspecies of the Alpine ibex, giving populations in the Alps the

trinomial In elementary algebra, a trinomial is a polynomial consisting of three terms or monomials. Examples of trinomial expressions # 3x + 5y + 8z with x, y, z variables # 3t + 9s^2 + 3y^3 with t, s, y variables # 3ts + 9t + 5s with t, s variables # a ...

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 In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...

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 ar ...

, 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 with ...

, 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 togethe ...

, 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 France, 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. ...

Valley in the

French Alps The French Alps are the portions of the Alps mountain range that stand within France, located in the Auvergne-Rhône-Alpes and Provence-Alpes-Côte d'Azur regions. While some of the ranges of the French Alps are entirely in France, others, such as ...

but it was

reintroduced Species reintroduction is the deliberate release of a species into the wild, from captivity or other areas where the organism is capable of survival. The goal of species reintroduction is to establish a healthy, genetically diverse, self-sustainin ...

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 betwe ...

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 Macedon ...

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, an ...

. 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 Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains furt ...

and mixed larch-spruce woodland if no snow has fallen. Males spend the winter in

coniferous Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extant ...

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 A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

, 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 ''Avena'' is a genus of Eurasian and African plants in the grass family. Collectively known as the oats, they include some species which have been cultivated for thousands of years as a food source for humans and livestock. They are widesprea ...

'', ''

Calamagrostis ''Calamagrostis'' (reed grass or smallweed) is a genus of flowering plants in the grass family Poaceae, with about 260 species that occur mainly in temperate regions of the globe. Towards equatorial latitudes, species of ''Calamagrostis'' general ...

'', ''

Festuca ''Festuca'' (fescue) is a genus of flowering plants belonging to the grass family Poaceae (subfamily Pooideae). They are evergreen or herbaceous perennial tufted grasses with a height range of and a cosmopolitan distribution, occurring on every ...

'', ''

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 20 ...

'', '' Poa'', ''

Sesleria ''Sesleria'' is a genus of perennial plants in the grass family. The are native to Eurasia and North Africa. They are found in Albania, Austria, Baleares, Baltic States, Belarus, Belgium, Bulgaria, Corsica, Czechoslovakia, East Aegean Islands, Fi ...

'', and ''

Trisetum ''Trisetum'' is a genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biologica ...

''. The climbing ability of the Alpine ibex is such that it has been observed standing on the sheer face of the

Cingino Dam The Cingino Dam is a gravity masonry dam located southwest of Antrona Schieranco, Province of Verbano-Cusio-Ossola in Piedmont, Italy. The dam creates Lago Cingino (or Cingino reservoir) which is fed by the Antigine and Troncone streams. The reser ...

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 estrous cycle (, originally ) is the set of recurring physiological changes that are induced by reproductive hormones in most mammalian therian females. Estrous cycles start after sexual maturity in females and are interrupted by anestrous p ...

. The higher the male's rank, the closer he can get to a female. Males perform

courtship display A courtship display is a set of display behaviors in which an animal, usually a male, attempts to attract a mate; the mate exercises choice, so sexual selection acts on the display. These behaviors often include ritualized movement ("dances"), ...

s. 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 Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...

and

poaching Poaching has been defined as the illegal hunting or capturing of wild animals, usually associated with land use rights. Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set a ...

. 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 , photo = Gran_Paradiso.jpg , photo_caption = The Gran Paradiso , elevation_m = 4061 , elevation_ref =Geoportale IGM owww.pcn.minambiente.it/ref> , prominence_m = 1879 , prominence_ref = , range = Graian Alps , parent_peak = , listing ...

and Vanoise Massifs, then both part of the

Kingdom of Sardinia The Kingdom of Sardinia,The name of the state was originally Latin: , or when the kingdom was still considered to include Corsica. In Italian it is , in French , in Sardinian , and in Piedmontese . also referred to as the Kingdom of Savoy-S ...

. 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 Victor Emmanuel II ( it, Vittorio Emanuele II; full name: ''Vittorio Emanuele Maria Alberto Eugenio Ferdinando Tommaso di Savoia''; 14 March 1820 – 9 January 1878) was King of Sardinia from 1849 until 17 March 1861, when he assumed the title o ...

, 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. T ...

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. In ...

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