The goat-antelopes, caprids or caprines (subfamily Caprinae), represent a very diverse range of species. These sure-footed, adaptable, hardy and often spectacular animals are popular zoo species which are relatively easy to cater for in captivity once their basic husbandry has been established. Although once popular and widespread in U.K. collections, they have enjoyed an even greater popularity in many other European zoos, a situation which continues today. During the late 1980s and 1990s there was a drop in their popularity in Britain and a reduction in keeping wild caprine species. In recent times, however, there seems to have been a turn-around and a renewed interest in keeping them, which has resulted in three species – Rocky Mountain goat, Japanese serow and Mishmi takin – producing their first U.K. breedings during 2006 alone. The realisation that many caprine species are becoming endangered and would benefit from captive-breeding programmes has accelerated that interest. Britain in particular seems to be moving towards keeping a wider variety of species than has been the case historically.
There are still a few collections holding Barbary sheep: it is unlikely that this species will become as commonly seen or as widespread in British collections as it once was, but we could see a move towards keeping subspecific groups instead, in accordance with recent EAZA recommendations.
Historical background – London Zoo
London Zoo was the first British collection to exhibit a wide variety of caprines. In fact the zoo achieved the first U.K. breedings of ten wild caprine species, eleven if you include Whipsnade’s breeding of the musk ox (Ovibos moschatus) in June 1969 (Table 1.) The first Mishmi takin (Budorcas taxicolor) to reach Europe alive was a male which was presented to London Zoo on 22 June 1909 by John C. White (Edwards, 1996). The animal was probably the first to be exhibited anywhere in the world. (Mr White had previously donated takin material to the Natural History Museum: the specimens were considered distinct enough for a new subspecies, B. t. whitei, to be named after him.) The takin was housed near the clock tower and lived at the zoo for nearly nine years before dying on 7 May 1918. A female was obtained on 25 January 1923; a much younger animal, she was actually used to wearing a collar and lead (ZSL, 1976). She was presented by Major F.M. Bailey and was kept at the foot of the Mappin Terraces until she died in 1935. Up to 1959 these were the only takins to have left Asia alive (Edwards, 1996).
The first Rocky Mountain goat (Oreamnos americanus) reached London Zoo in 1900; this adult male was believed to be the first living specimen brought to Europe (Ashby, 2009).
A pair of blue sheep or bharal (Pseudois nayaur) were purchased on 19 February 1880; in 1885 they were housed in a small enclosure near the Lion House at London Zoo. Between 1882 and 1908 some 47 lambs were born at the zoo. The original male died on 15 August 1893 and the female on 4 March 1896 (Edwards, 1996). In another similar enclosure, located to the east of the Elephant House, near the site of the Insect House, Himalayan tahrs (Hemitragus jemlahicus) were kept. Between 1876 and 1937 some 115 offspring were born at London Zoo. Interestingly, one female born on 17 June 1897 lived until 3 June 1914 – nearly 17 years!
Table 1. Wild Caprine First U.K. Breedings. (Data from Ashby, 2006)
Mouflon (Ovis orientalis) – pre-1785, private estate.
Barbary sheep (Ammotragus lervia) – 1854, London Zoo.
Nubian ibex (Capra nubiana) – 1854, London Zoo.
Urial (Ovis vignei) – 1858, London Zoo.
Chamois (Rupicapra rupicapra) – 1864, London Zoo.
Markhor (Capra falconeri) – 1867, London Zoo.
Wild goat (Capra aegagrus) – 1869, London Zoo.
Bharal (Pseudois nayaur) – 1882, London Zoo.
Himalayan tahr (Hemitragus jemlahicus) – 1885, London Zoo.
Argali (Ovis ammon) – c. 1902, Woburn Park.
West Caucasian tur (Capra caucasica) – 1911, London Zoo.
Bighorn sheep (Ovis canadensis) – 1932, London Zoo.
Musk ox (Ovibos moschatus) – June 1969, Whipsnade Zoo.
Alpine ibex (Capra ibex) – 1971, Norfolk Wildlife Park.
Rocky Mountain goat (Oreamnos americanus) – May 2006, Colchester Zoo.
Japanese serow (Capricornis crispus) – 2006, Edinburgh Zoo.
Takin (Budorcas taxicolor) – 2006, Marwell Zoo (followed shortly by Highland Wildlife Park).
The first chamois (Rupicapra rupicapra) to arrive at London Zoo were presented in 1831; others followed later.
In 1842 London Zoo imported some Barbary sheep or aoudad (). After the Mappin Terraces were opened in May 1914 a single pair of Barbary sheep were installed there, along with ibex (Capra ibex) and others. The so-called ‘Goat Hills’ consisted of four areas capable of housing four separate species; these varied over time, but in later years the enclosures housed Barbary sheep, mouflon (Ovis orientalis), markhor (Capra falconeri) and bighorn sheep (O. canadensis) fairly consistently. In the following 23 years nearly two hundred Barbary sheep were born at the zoo; as only one extra pair were brought in, the genes of these animals were rather concentrated, with a degree of inbreeding as time went by. Despite this, however, the stock remained in first-class condition, although some minor defects – such as poor horn formation, hoof problems and patches of white just above the hind hooves – have subsequently been seen in related groups in other British collections (O’Grady, 1998). Most of the Barbary sheep in the U.K. were, and still are, descended from London Zoo’s stock. In the 1920s some went to Edinburgh Zoo from London, and from Edinburgh to other U.K. collections. Bristol Zoo, for example, had some listed in 1937 and 1938 at least (Keeling, 1998).
In 1932 the Goat Hills contained Himalayan tahr, Barbary sheep, ‘Grecian ibex’ (Capra aegagrus, today known as bezoar or Cretan wild goat), and some hybrids between Himalayan ibex (C. sibirica) and West Caucasian tur (C. caucasica). In 1929 a pair of Canadian bighorn sheep arrived as a donation from Banff, Alberta. A Mishmi takin and a musk ox were also still to be seen in the collection at that time.
Whilst London Zoo remained the stronghold for wild caprines in the U.K., over the years more collections gradually started keeping them. The majority were Barbary sheep, but other species could occasionally be found in individual zoos. For example, the Alpine ibex (Capra ibex) became established at both the Highland Wildlife Park (Kingussie) and the Norfolk Wildlife Park (Great Witchingham). The latter had the first U.K. breeding in 1971, and both collections kept and bred the species for many years; but no more were imported and no stock were moved elsewhere to form new herds, so they eventually all died out. Philip Wayre had also kept some Barbary sheep in his private collection before he opened the Norfolk Wildlife Park in 1962.
In 1973, prior to the opening of Marwell Zoological Park, a pair of common goral (Naemorhedus goral) arrived. They remained off-show, and by 1975 only the female was left; by 1976 she too had gone (International Zoo Yearbook, 1974, 1975, 1976). They were not replaced, and no more goral were kept in the U.K. until very recent times.
Woburn Park kept Himalayan tahr, and in 1966 they bred six animals; two more were born in 1981. Interestingly, a pair of Dall’s sheep (Ovis dalli) were kept at Flamingo Gardens, Olney, around 1977: they were recorded in the International Zoo Yearbook (IZY) as a pair in 1980 and 1981, but they had departed by 1982. As far as I am aware, these were the only Dall’s sheep ever kept in the U.K. Between January 1977 and April 1978, two (2.0) musk oxen arrived at Chester Zoo, but only one was still alive in 1979 (IZY, 1980). In 1995 Whipsnade’s male musk ox died, leaving just one female by the end of the year.
Bristol Zoo took on Barbary sheep again in 1979 with 1.2 animals; they were kept with the crab-eating macaques in the Monkey Temple enclosure.
Around the time of the opening of Blackpool Zoo in 1972 a fine adult male markhor, who had been the mascot of a Royal Air Force squadron, was presented to the zoo, as the animal had become somewhat unmanageable. This male is thought to have been the original breeding animal at Blackpool (C. Bloor, pers. comm., 2010). A small herd of 2.6 Barbary sheep arrived in Blackpool from Belle Vue Zoo, Manchester, on 17 May 1972 (Keeling, 1992), and more were added in 1974/75. On 4 September 1973 1.6 chamois also arrived at Blackpool from Belle Vue’s quarantine station, two kids having been born while they were there (Keeling, 1992). They were installed in an island enclosure with rocky sandstone outcrops, and in 1975 three more young were born. In 1976 the group numbered 2.6 animals (IZY, 1977). Numbers of chamois were kept constant through the late 1970s and early 1980s, with young being regularly born, though adult numbers were kept to a small group. Around 1984 Blackpool received London Zoo’s herd of markhor as London started to clear the Goat Hills of stock: they were installed in Blackpool’s former chamois island enclosure, the chamois having been moved to a grassy paddock. In April 1985 the last two Barbary sheep – descendants of Belle Vue’s original stock – left Blackpool for the animal dealers, Ravensden. In 1985 Blackpool still had 1.3 chamois (IZY), and the species was still listed in the 1987 guidebook, though I did not see any when I visited in 1987. Apparently they were still there off-show: some left the zoo in April 1989 to go to Palacerigg Country Park, Cumbernauld (Keeling, 1992), and another 1.2 left Blackpool during the first six months of 1990 (Ratel). London Zoo also had 1.3 chamois in 1985, for the first time in many years, but it is not clear whether these came from Blackpool Zoo stock. There was no further mention of the London Zoo chamois (IZY), so perhaps they were held temporarily. Palacerigg had 1.3 chamois in 1995/96 and were still breeding them in 1997.
In 1990, Blackpool Zoo had 6.2 markhor; three more were born in 1990 and two in 1991 (IZY) – they had been breeding since their arrival and were doing very well. At one point, in fact, the markhors took up two enclosures, the island and a paddock. They continued to prosper throughout the 1990s. The late Martin Bourne’s collection near Manchester also had a pair by 1995; it is likely that these came from Blackpool Zoo, as 1.0 markhor had left in 1993 followed by 0.1 in 1994 to a ‘private collection’ (Ratel). Another pair went to the Bourne collection in 1996, but the last survivors, a pair, left in 1998. Three (0.3) more were born at Blackpool Zoo in 1996. At around this time a sign on the zoo’s markhor enclosure stated that some had moved to Knowsley Safari Park; a spokesperson confirmed to me that they were briefly kept at the park, but firm details are lacking. In 1998 Blackpool’s markhors were moved off their island enclosure; at this time they were still the only breeding group in the U.K.
A return to Britain – after a very long absence – was that of the Rocky Mountain goat (Oreamnos americanus). None of this spectacular species had been kept in Britain since the first specimen reached London Zoo back in 1900, so great excitement surrounded the first pair which arrived at Colchester Zoo in 1997. They bred in May 2006, achieving a U.K. first (Ashby, pers. comm., 2010), and did relatively well, despite husbandry issues involving the regular, but difficult, procedure of hoof trimming. These are large and potentially dangerous animals to restrain manually, especially the adult males. I believe it was ultimately for this reason that Colchester made the reluctant decision to go out of keeping them. The adult male was the first to leave the U.K., followed by two females to Rostock and Chemnitz (Germany) respectively. This left the female, Polar, and her offspring Lupin in July 2009 (Vaughan, pers. comm., 2010), who were also to leave in due course, though according to ISIS (March 2010) one female was still apparently awaiting departure off-show.
Of all the goat-antelope species held in the U.K. the Barbary sheep has been the most commonly kept, represented in a large number of British collections. Most of them, however, have been of ‘mixed race’, as are the 900+ animals in EAZA zoos at the moment. It would be almost impossible to genetically identify most of the present captive stocks and separate them into any subspecific groupings. There are now at least five recognised subspecies or regional races, one of which, the Egyptian Barbary sheep (Ammotragus lervia ornatus) is now extinct in the wild. The Saharan race (A. l. sahariensis) is listed as Vulnerable by the IUCN, so ideally efforts should be made to try to establish separate populations of this subspecies as a priority. There are currently moves by some countries to start phasing out the mixed-race stock in favour of keeping those subspecies which are in particular need of conserving, but this process is currently very slow, and as yet has not started in the U.K.
London Zoo maintained a herd of between 40 and 50 Barbary sheep on the Mappin Terraces throughout the 1960s and 1970s. Dudley Zoo had a small herd in 1937 which all came from London; they were installed in a small hillock enclosure to the left of the entrance (Keeling, 1989), but were later moved to a more suitable enclosure, with no access to grass, in which they remain to this day. Although Barbary sheep will readily consume grass, it can cause digestive problems. Himalayan tahrs, which are even less tolerant of it, were also initially kept on grass at Dudley in 1948; these animals also probably came from those bred at London Zoo. Following the arrival of Barbary sheep at Edinburgh Zoo in the 1920s (O’Grady, 1998) and subsequent breeding successes over the years, Edinburgh sent five animals to Glasgow Zoo in 1968; over sixty were born there, and many went on to other U.K. collections and overseas. Chessington Zoo obtained Barbary sheep around 1960 (Keeling, 1996), probably from London Zoo, and their inventory for 1963 records three (1.2). They were initially kept in a paddock to the right of the entrance, but after a year or two they were given their own enclosure. This was essentially a narrow hard-standing with a concreted rocky area and a little grass; there was a shed at one end which they rarely used, and as a result they were almost impossible to shut in while the enclosure was being cleaned. They remained in this enclosure until about 1985, when it was given to the guanacos.
Three collections which now no longer exist – Weyhill Wildlife Park (Andover), Thorney Wildlife Park (Wisbech) and Sherwood Zoo (Hucknall) – all kept Barbary sheep around 1970. Dudley Zoo’s second enclosure for the species, which still stands today, was built just prior to 1970. By 1972/73 Glasgow Zoo’s herd numbered around 20 animals, and in 1974/75 some 6–8 animals were sent to Blackpool. Newquay Zoo, Hotham Park Zoo (Bognor) and Basildon Zoo all kept Barbary sheep around 1976. West Midlands Safari Park also kept some: the initial group of 2.3 animals arrived in 1977 from Belle Vue when it closed down (Keeling, 1992), and two lambs were born in 1979 (Ratel).The largest breeding herds were kept by London, Dudley, Edinburgh, Glasgow and Blackpool Zoos (O’Grady, 1998); but Windsor Safari Park, which had added the species just prior to 1985, built up its herd in a relatively short time and had some 30 animals by the time the park closed in 1993. Smaller groups were kept at Paignton Zoo, Paultons Park (Romsey), Suffolk Wildlife Park, and Belle Vue (until it closed in 1977).
For several years the species had proliferated, with London Zoo being the largest producers by far, and a surplus of animals was regularly available. Several were sent overseas from other U.K. zoos as well, usually via Ravensden, to help alleviate the problem. Some collections started to control breeding where possible, and numbers gradually became more manageable.
Barbary sheep are robust and adaptable, though they tend to do better in the more urban collections. These are certainly better placed to provide more suitable facilities such as artificial or natural rockwork and hard-standing areas, rather than completely grassy paddocks, although a combination of the two works fairly well, with more of the former and less of the latter. Where groups have been maintained in open grass paddocks they have not flourished as well as elsewhere, as they do not like damp and soft conditions for too long. They obviously require a good amount of hoof-wear to control overgrowth, and this can still be a major problem in this species even where hard substrates are provided.
As a species they are popular with the public – the lambs especially so, although the sight of a mature adult male is a very impressive one. Adult males have a distinct aroma – like other adult male caprines, they have the habit of urinating on themselves. In the 1980s, early on in my career, I briefly worked with the small herd of Barbary sheep at Chessington Zoo. We had the interesting task of moving the dominant adult male, Godfrey, at various intervals to control breeding activity. He was strong, but we walked him to and from his enclosure – a common, hands-on way of moving stock in those days that today would not be allowed due to health and safety risks. When isolated, he was held in a pen at the nearby Children’s Zoo area. Obviously he disliked being taken away from his herd: the return journey proved rather quicker for us and the end result more enjoyable for him! He was a great character.
The decline in the keeping of Barbary sheep started around the mid- to late 1980s, though Edinburgh Zoo had already stopped by the late 1970s or early 1980s: this was a significant step, combined with London Zoo’s decision to stop keeping Barbary sheep on the Mappin Terraces. Only their bighorn sheep (Ovis canadensis), a large herd of 20 (7.13) animals, remained on the Mappins long after the area had been closed in 1984: these were the only ones left in the U.K. at that time and were, incredibly, still there in 1990. (Some were later moved to Whipsnade when London’s famous ‘goat hills’ were finally cleared of caprines. Chessington’s Barbary sheep had moved on around 1986 after a rather unsuccessful attempt to mix them with Barbary macaques in the redeveloped former bear enclosures in 1985. By 1987 that enclosure was also swept away to become a themed train ride.
Dudley and Paignton still had groups of Barbary sheep in the late 1980s, Glasgow was still breeding them (though restricted in numbers), and Bristol had two males briefly in 1989, which may have been their last. By 1995 Dudley Zoo, Glasgow Zoo, Paignton Zoo, Suffolk Wildlife Park and Paultons Park kept the main groups of Barbary sheep in the U.K. (Zoo Federation inventories). Dudley and Glasgow always kept the larger groups; Paignton’s enclosure was limited in size and Paultons’ animals were kept in a small concrete yard with a rocky outcrop (similar to Paignton’s), but with access to a large grassy field. Incidentally, on one occasion an unfortunate animal at Paultons was struck by lightning and died (pers. comm., ex-Paultons keeper, 2010). In 1996/97 the same five collections were continuing to keep Barbary sheep: numbers remained consistent, only Dudley showed a slight decrease. Personally, I have always felt that Dudley Zoo’s Barbary sheep enclosure is an excellent example of a good artificial urban zoo exhibit: despite its perhaps ‘unnatural’ appearance, it has done particularly well with the species over the years, and also shows the animals off to their best, providing the public with a very good, close view of their behaviour.
To bring things right up to date as of March 2010 (ISIS), Barbary sheep in the U.K. can be found at Dudley Zoo (still a good group and still breeding), Africa Alive! (formerly Suffolk Wildlife Park) and West Midlands Safari Park (still maintaining large groups), and Paignton Zoo. Paignton have four females left now, in a new enclosure which has been formed out of a natural escarpment, a very unusual and interesting exhibit which utilises the natural environment to best effect. An adjoining hardstand and indoor stabling is a useful addition, and a larger group could potentially be housed here (or indeed it could equally be used for other caprines). Paultons gave up keeping Barbary sheep when it cut down on all its larger hoofstock in the late 1990s: they were not really thriving on that grass paddock, in any case.
Affectionately known as ‘Falconer’s goat’, the largest and most impressive member of the Caprine family is the markhor, an endangered species, with currently fewer than 2,500 adult animals left in the wild. There are three subspecies recognised by the IUCN. The captive European population numbers around 200 animals, and in the late 1990s Helsinki Zoo initiated the European Studbook for the Turkmenian or Bukharan markhor (Capra falconeri heptneri).
London Zoo was the first U.K. collection to keep markhor, but the last ones bred there were five kids born in 1983 on the Mappin Terraces before the animals went on to Blackpool Zoo. For many years Blackpool’s animals were the only breeding group in Britain. Captive numbers in the U.K. have never been large and breeding success has been limited, though consistently steady over the years: again, though, there has been a lack of new imports of unrelated animals, combined with a reluctance of new potential holders to come forward, which has resulted in a relatively static population. Recently, however, this situation changed when, in 2007, the Highland Wildlife Park obtained 1.3 animals from Liberec Zoo, Czech Republic, and Blackpool Zoo. In addition a further 4.2 animals joined the group from Blackpool, Helsinki and Nordens Ark in 2008. In early 2010 Blackpool’s last pair of markhor travelled up to join the others at Highland. Blackpool Zoo now no longer keeps any wild caprines, for the first time since its opening.
The first Mishmi takins to arrive in the U.K. since the first two early ones at London Zoo were two young males who arrived in 2000 (Irven, 2009); one went to Colchester Zoo, the other to Marwell Zoo. They both came from Tierpark Berlin, Germany, and were believed to be related to some degree, which is unfortunately the case with many of the takins in Berlin. The Marwell animal was named Willi and became very popular with staff, but sadly he died at the age of only 14 months from pancreatic cancer. This was thought to be a genetically-linked condition, as Colchester’s animal also died around the same time. Marwell subsequently made a commitment to keeping this species in the future.
With this in mind a young pair of Mishmi takin arrived on 18 March 2004 from Nuremberg and Frankfurt Zoos respectively. At first the young pair, the male Mooshu (born March 2003) and female Mulan (born March 2002), were very nervous, but they gradually settled in. They were kept in a holding enclosure while a new enclosure was being prepared for them: an area of ‘mock rockwork’ was incorporated into the landscape, as well as several large upright logs. On 21 June 2006 the first Mishmi takin was born in the U.K. to this pair. The offspring, a male, was named Chien-Po. A second calf was born on 30 September 2007, this time a female named Mei-Ling. In early 2009 a third calf was unfortunately stillborn.
Meanwhile the Highland Wildlife Park had established their own large herd of Mishmi takin, and they achieved their first birth soon after Marwell’s first-born. Currently, Marwell and the Highland Wildlife Park are still the only U.K. collections keeping Mishmi takin. The herd at Highland is much the larger and as a result has been far more successful in terms of breeding. They are able to keep a larger social herd in an area much greater than that of Marwell’s current facilities, although this may change in the future. I look forward to being able to watch our group develop. Takin keepers in the U.S.A. have called these animals ‘goats with attitude’ – a phrase I would completely agree with. I work closely with the pair at Marwell and have conducted a successful programme of environmental enrichment with them to help reduce and redirect the adult male’s aggressive behaviour (Irven, 2009).
As mentioned above, this collection has had a history of keeping wild caprines. It is also blessed with a wonderful natural landscape and environment which is nearly ideal for their captive husbandry. The park currently has the largest and most diverse collection of wild caprines of any U.K. collection (Table 2), and there is great potential for maintaining larger groups in the future.
The Mishmi Takin EEP is now being managed and coordinated by Douglas Richardson at Highland Wildlife Park. In 2008/9 Highland had a larger group of this species, up to a dozen animals (Sellars, pers. comm., 2008), but since then they have suffered some unfortunate losses to the group. They are maintained in a wonderful enclosure incorporating an area of coniferous woodland which is ideal for takin.
Table 2. Caprines at Highland Wildlife Park. (Data from ISIS as of March 2010.)
Chinese goral (Naemorhedus griseus) – 2.1 (plus another 2.2.1 at Edinburgh Zoo).
Bharal or blue sheep (Pseudois nayaur) – 4.0.
Japanese serow (Capricornis crispus) – 1.1.
Turkmenian markhor (Capra falconeri heptneri) – 6.1.
Himalayan tahr (Hemitragus jemlahicus) – 6.6.2.
Mishmi takin (Budorcas t. taxicolor) – 2.3.
In 2008 there was a group of Afghan urials (Ovis aries bochariensis) as well, but these are no longer in the collection. Mouflon are also no longer held at the park, as the species is now deemed to be a domestic form and is not recommended by the EAZA Sheep and Goat TAG to be held in serious collections.
Caprines not held in the U.K.
Unfortunately there are still many wild caprines that are not kept in U.K. zoos. It is a great shame that there are no ibex or chamois and no western or eastern Caucasian tur, despite EAZA recommendations. The last Rocky Mountain goat is to leave Colchester Zoo, if it has not already done so by the time you read this. There are no Dall’s or Stone’s sheep, argali, bighorn sheep, and possibly no urial. Numbers of mouflon are also greatly reduced from the days when it was a commonly kept species in the U.K.
The long-term future for takins in the U.K. looks hopeful. There will be further Mishmi takin brought in: certainly there is a strong chance that Marwell will be obtaining at least another female when available, now that we have extended the present facilities and may well expand even further in due course. There has also been some discussion about the possibility of bringing a pair into Paignton Zoo in the future; this would be excellent news, increasing the number of collections participating in the breeding programme within the U.K. itself and in the overall EEP for the species. The future also looks better for markhor, although ideally it would be nice to see other U.K. collections taking on animals as well.
It seems likely that the ‘mixed race’ Barbary sheep will eventually die out – only three collections are still breeding with the ones they have. Again, it would be good to move towards establishing pure groups of endangered subspecies and phasing out the rest.
As shown in Table 2, the Highland Wildlife Park currently has four male bharal: there are no plans to import females, as the park acts as the bachelor male holder for the European studbook. At the time of writing the park is planning to import 0.2 musk oxen that stemmed from the Czech Republic: the animals are currently in quarantine in the Netherlands, where a female calf was born, but they are expected to arrive in Scotland before the end of the year, and a new male is being sourced.
References and sources
Ashby, A (2009): We Went to the Zoo Today . . . The Golden Age of Zoo Postcards Independent Zoo Enthusiasts Society, Todmorden.
Ashby, A. (2006): First and Early Breeding Records for Wild Mammals in the U.K. and Eire. Bartlett Society (published online at http://zoohistory.co.uk).
EAZA (no date): Sheep and Goat TAG overview.
Edwards, J. (1996): London Zoo from Old Photographs 1852-1914. John Edwards, London.
Irven, P. (2009): Mishmi takin (Budorcas taxicolor) enrichment at Marwell Zoological Park. Ratel 36 (1): 5–9.
ISIS: Data taken from the International Species Inventory System (www.isis.org) listings as of March 2010.
Keeling, C. (1998): The Bristol Book – the Story of Bristol Zoo. Clam Publications, Guildford.
Keeling, C. (1996): The Chessington Story – Chessington Zoo. Clam Publications, Guildford.
Keeling, C. (1992): The Fragments that Remain – the Story of Belle Vue Zoo. Clam Publications, Guildford.
Keeling, C. (1989): They Lived at the Castle – the Story of Dudley Zoo. Clam Publications, Guildford.
O’Grady, R. (1998): Barbary sheep at Glasgow and other places. Zoo! (magazine of the Independent Zoo Enthusiasts Society) 10: 5–8.
Ratel: (Data from various years.) Association of British Wild Animal Keepers.
Zoological Society of London (1976): Golden Days – Historic Photos of London Zoo. Duckworth, London.
International Zoo Yearbook: (Data from various years.) Zoological Society of London.
Zoo Federation (Federation of Zoological Gardens of Great Britain and Ireland, now BIAZA): Data taken from the mammal inventories from various years.
Various U.K. zoo guidebooks.
I would like to thank the following people for their assistance with this paper:
Alan Ashby, Rob Vaughan, Cyril Bloor and Raymond Robinson, zoo enthusiasts and historians; Marc Fox and David White, both Marwell section heads, for their information; Douglas M. Richardson (Animal Collection Manager) and Morag Sellars (Head Animal Keeper) at Highland Wildlife Park, and Matthew Dodd of Knowsley Safari Park, for their responses to my enquiries; and the late Clin Keeling, who was a constant inspiration, and great friend, to me during my career – we are indebted to him for recording and publishing so much information on zoological history with incredible attention to detail.