|Illustration from 1898|
|Subspecies:||C. p. pyrenaica|
|Capra pyrenaica pyrenaica
The Pyrenean ibex (Capra pyrenaica pyrenaica), Spanish common name bucardo, was one of the four subspecies of the Spanish ibex or Iberian wild goat, a species that is endemic to the Iberian Peninsula. Pyrenean ibex were most common in Cantabrian Mountains, Southern France and in the Northern Pyrenees. This species was common during the Holocene and Upper Pleistocene, during which their morphology, primarily some skulls, of the Pyrenean ibex was found to be larger than other Capra subspecies in southwestern Europe from the same time.
In January 2000, the Pyrenean ibex became extinct as scientists attempted to clone DNA from one of the last females. The Pyrenean ibex left behind the Western Spanish ibex, or Gredos ibex, and the Southeastern Spanish ibex, or Beceite ibex, while the Portuguese ibex became extinct. Since the last of the Pyrenean ibex went extinct before scientists could adequately analyze them, the taxonomy of this particular subspecies is controversial regarding whether it was larger than other subspecies or not.
There are multiple theories regarding the evolution and historical migration of the C. pyrenaica into the Spanish peninsula. Scientists, including paleontologists and geneticists, should continue to find paleologic, phylogenetic, and other evidence to discover a molecular connection between the different subspecies. Knowing the history of the species can better teach scientists how the ibex and C. pyrenaica affected the region and could provide a new understanding for the relationship between subspecies.
One possibility is that the C. pyrenaica evolved from an ancestor related to C. caucasica from the Middle East at “the beginning of the last glacial period (120-80 ky).” The C. pyrenaica most likely moved from the northern Alps and traveled through southern France into the Pyrenees area at the beginning of Magdalenian period at about 18 kya. If this is the case then the C. caucasica praepyrenaica may have been more different from the other three ibex species that lived in the Spanish peninsula than scientists currently know. For example, this would mean that the C. pyrenaica (possible migration 18ky) and C. ibex (300 ky earlier migration) would have evolved from different ancestors and been morphologically more different from their separate genes. It is known that all four subspecies lived together in the Upper Pleistocene time, but scientists are unsure of how much genetic exchange there could have been. The problem with this theory is that genetics suggest that the C. pyrenaica and C. ibex could have shared a more common origin, possibly the C. camburgensis.
There are many variations of when the C. pyrenaica or C. ibex migrated and evolved in the Spanish peninsula first. It is a possibility that the C. pyrenaica was living in the Spanish Peninsula by the time that the ibexes began to migrate through the Alps. Genetic evidence also supports the theory that multiple Capra subspecies migrated to the Spanish region at around the same time period. Hybridization may have been possible, but the results are not conclusive.
Behaviour and physical characteristics 
The Pyrenean ibex had short fur which varied according to seasons. During the summer, its hair was short and come winter, the hair grows longer and thicker to accommodate the weather. The abundance and length of hair on the ibex’s neck remained the same throughout all seasons. Male and female ibexes could be easily depicted due to color and fur differences. A male ibex is commonly a faded greyish brown during the summer. Males are decorated with black in several places on the body such as the mane, forelegs and forehead. In the Winter, the ibex is less radiant with its color. A male transforms from a greyish brown to a dull grey and where the spots were once decorated with black, it became dull and faded as well. A female ibex on the other hand can easily be mistaken for a deer since its fur was brown throughout the summer. Unlike the male ibex, a female will lack black coloring. However, the offspring of the ibexes would result to looking like a female during the first year of life. 
A male ibex is known for its intricate horns which appear large and thick “curving outwards and backwards then outwards and downwards then inwards and upwards.” The surface of the horn is rough containing ridges that develop progressively with age. Its rings are said to each represent a year, so if one were to count an ibex’s rings, the total would correspond to the ibex’s age. Just like in color, the female is different than a male in its horn structure as well. Unlike the long, complex horn of males, a female possesses a short, cylindrical horn. ibexes (male and female) had the diet of a herbivore. They mostly fed on greens such as grass and herbs.
Pyrenean ibexes migrated according to seasons. During Spring, the ibex would migrate to more elevated parts of mountains where females and males would mate. After the mating sessions, females would normally leave the males behind so they can give birth in a more isolated area. Birthing season is typically during the month of May. Like most mammals, the ibex only gives birth to one offspring. Twins aren’t impossible to give birth to but it is highly unlikely. During the winter, the ibexes would migrate to valleys that aren’t covered in snow. These valleys allowed them to continue their diet regardless of the change in season.
The species has been often spotted throughout the areas of France, Portugal, Spain, and Andorra but not as much in northern areas of the Iberian Peninsula. In areas like Andorra and France in the mainland, the Pyrenean ibex was extinct originally in the northern tip of the Iberian Peninsula. The Pyrenean ibex was estimated to have 50,000 individuals with more than 50 other sub groups that ranged from the Sierra Nevadas to Sierra Morena and Muela de Cortes. Many of these sub groups lived and thrived in sierras and ranges of the sort, which expanded into Spain and into Portugal. The last remains of the Pyrenean ibex was spotted in areas of Middle and Eastern Pyrenees all of which were below twelve hundred meters of altitude. However, in areas of southern France and surrounding areas, the ibex was founded to Upper Pleistocene and Holocene which would range from 350–925 meters and between 1190–2240 meters, making it the highest elevations for Pyrenean ibexes to be spotted at.
Pyrenean ibex was quite abundant up until the 14th century and numbers did not dwindle down in the region until the mid 19th century. Pyrenean ibexes tend to live in rocky habitats that are filled with cliffs and trees interspersed with scrub or pine trees. However, small patches of rocks in farmland or various areas along the Iberian coast would be livable conditions for the Pyrenean ibex. The ibex was able to thrive well in its environment as long as the appropriate habitat is available because they were able to disperse rapidly and colonize quickly. Pyrenean ibexes were also good sources of revenue to the local areas that they thrive in, which may have been a cause to their eventual extinctions. Researchers say that the eventual downfall of the Pyrenean ibex may have been caused by continuous hunting and even perhaps that the animal could not compete with the other live stock in the area. However, definite reasons for the extinction of this animal is still unknown.
The subspecies once ranged across the Pyrenees in France and Spain and the surrounding area, including the Basque Country, Navarre, north Aragon and north Catalonia. A few hundred years ago they were numerous, but by 1900 their numbers had fallen to fewer than 100. From 1910 onwards, their numbers never rose above 40, and the subspecies was found only in a small part of Ordesa National Park, in Huesca.
The Pyrenean ibex was one of four subspecies of the Spanish ibex. The first to become extinct was the Portuguese ibex (Capra pyrenaica lusitanica) in 1892. The Pyrenean ibex was the second, with the last individual, a female called Celia, found dead in 2000.
In the Middle Ages, Pyrenean ibex were very abundant in the Pyrenees region but decreased rapidly in the 19th and 20th centuries due to hunting pressure. In the second half of the 20th century only a small population survived in the Ordesa National Park situated in the Spanish Central Pyrenees.
Competition with domestic and wild ungulates also contributed to the extinction of the Pyrenean ibex. Much of the range of the Spanish ibex was shared with sheep, domestic goats, cattle and horses, especially in summer months when it was in high mountain pastures. This led to interspecific competition (Fandos, 1991; Martínez, 1992) and overgrazing, which affected the ibex particularly in dry years. In addition, the introduction of alochtonous wild ungulate species in areas occupied by the ibex (e.g. Fallow Deer and Mouflon in the Sierras de Cazorla, Segura y Las Villas Natural Park) increased the grazing pressure, as well as the risk of transmission of both native and exotic diseases (Fandos & Reig,1992).
The last natural Pyrenean ibex, a female named Celia, was found dead on January 6, 2000, next to a fallen tree. Although her cause of death is known, the reason for the extinction of the subspecies as a whole is a mystery. Some hypotheses include the inability to compete with other species for food, infections and diseases, and poaching. The Pyrenean ibex became the first taxon ever to become "un-extinct", for a period of seven minutes in January 2009, when a cloned female ibex was born alive and survived a short time, before dying from lung defects.
Cloning project 
The biotechnology company Advanced Cell Technology, Inc. announced on October 8, 2000 that the Spanish government has agreed to their offer to use nuclear transfer cloning technology in collaboration with other scientific partners to clone the Pyrenean ibex from the tissue that was taken in 1999. The cloned individual was captured in National Ordesa Park in Huesca, Spain; skin biopsies were taken and cryopreserved in nitrogen. Celia, the last ibex, died a year after donating tissue from her ear.
It was expected to be easier than the cloning experiment of endangered gaur (Bos gaurus), as the reproductive biology of goats is better known and the normal gestation period is only five months. In addition, only certain extinct animals are candidates for cloning because of the need for a suitable proxy surrogate to carry the clone to term. ATC has agreed with the government of Aragon that the future cloned Pyrenean ibexes will be returned to their original habitat.
Celia provided suitable tissue samples for cloning. However, attempts to clone her have highlighted a major problem: even if it were possible to produce another healthy Pyrenean ibex, there are no males for the female clone to breed with. To produce a viable population of a previously extinct animal, there will need to be genetic samples from many individuals to create genetic diversity in the cloned population. This is a major obstacle to reestablishing an extinct species population through cloning. One solution could be to cross Celia's clones with males of another subspecies, although the offspring would not be pure Pyrenean ibex. A more ambitious plan would be to remove one X chromosome and add a Y chromosome from another still-existing subspecies, creating a male Pyrenean ibex, but such technology does not yet exist and it is not known whether this will be feasible at all without irreparably damaging the cell.
Three teams of scientists, two Spanish and one French, are involved in the cloning project. One of the Spanish teams was led by Dr. Jose Folch of Zaragoza, from the Centre of Food Technology and Research of Aragon. The other teams had researchers from the National Research Institute of Agriculture and Food in Madrid.
The project is coordinated by the Food and Agricultural Investigation Service of the Government of Aragon (Spanish: Servicio de Investigación Agroalimentaria del Gobierno de Aragón) and by the National Institute of Investigation and Food and Agrarian Technology (Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria). The National Institute of Agrarian Investigation of France is also involved in the project.
Researchers took adult somatic cells from the tissue and fused them with oocytes from goats that had their nuclei removed. The purpose of removing the nuclei from the goats’ oocytes was to extract all the DNA of the goat, so there would be no genetic contribution to the clone from the egg donor. The resultant embryos were transferred into a domestic goat (Capra hircus), to act as a surrogate mother. The first cloning attempts in 2003 failed. Of the 285 embryos reconstructed, 54 were transferred to 12 mountain goat and mountain goat-domesticated goat hybrids, but only two survived the initial two months of gestation before they too died. In 2009, one clone was born alive, but died seven minutes later, due to physical defects in the lungs. There was an atelectasis and an extra lobe in the left lung. This is not surprising, as lung defects have occurred in sheep clones before; the famous clone Dolly had a different sort of lung defect. Atelectasis doesn’t allow normal oxygen absorption to healthy tissues and is often related to a pneumothorax. DNA decomposes even when frozen, and the DNA is worse off because it was taken from an aged individual (Celia was 13). Aged DNA is known to cause cloned animals problems because of shortened telomeres, resulting in decreased lifespans. This means that had the clone survived the initial problems, it still would have had a shortened lifespan.
This was the first attempt to revive an extinct subspecies, although the process technically began before the extinction of the species.
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