IUCN/SSC - Caprinae Specialist Group
Workshop on Caprinae taxonomy
Ankara, Turkey, May 8-10, 2000
|I am not sure what this is||
These look like Ovis canadensis ????
|Who could get us some DNA?||But can you tell what subspecies?|
Goals of the workshop:
Update the current status of taxonomy of mountain ungulates
Identify knowledge gaps, particularly those most relevant to conservation
Provide a forum for a frank and open exchange ideas about caprin taxonomy and conservation
Foster the establishment of collaborations among researchers
Favor coordination of future research and conservation efforts
For those taxa for which a consensus exists, produce a guide to the identification of the world's mountain ungulates
|Could you help a
identify this skull?
Is this a CITES species?
Can you tell where it comes from?
From the 1997 IUCN Caprinae Action Plan:
12.2.2 Taxonomy and Genetic Diversity
A total revision of the entire subfamily is necessary if long-term Caprinae conservation is to be effected. Not only is there biological justification for this task, but as Val Geist observed, a taxonomy has major implications for conservation legislation. The immediate benefits of a revised Caprinae taxonomy include: a) improved CITES identification sheets to aid law enforcement officials in their task of controlling the illegal trade in taxa, b) clarification of biological and legal aspects for conservation laws, and c) an advance in our understanding of the phylogenetic relationships within the Caprinae.
However, it should also be borne in mind that most conservation actions do not necessarily require a formal taxonomy, especially where management units can be defined by other criteria.
Recommended actions and implementation:
1) Taxonomy Working Group. Establish a Taxonomy Working Group within the IUCN/SSC Caprinae Specialist Group. This will require additional members to the Specialist Group (see below) with responsibilities to: 1.1) organise an international workshop on Caprinae evolution and taxonomy; and 1.2) revise the taxonomy of the Caprinae. To be useful, a revision of Caprinae taxonomy will require a multi-disciplinary, integrated approach involving DNA analysis, internal and external morphology, behaviour, zoogeography, palaeontology, and evolutionary history. It will also require large sample sizes if it is to be a valuable revision. A revision would be most effectively achieved if a Taxonomy Working Group of the Caprinae Specialist Group is established with scientists from around the world representing disciplines that directly bear on taxonomic criteria. This action requires additional new members to the IUCN/SSC Caprinae Specialist Group. The working group's task would be to work co-operatively towards the goal of publishing a revised taxonomy of the subfamily, either in a single or a series of scientific reports. The working group should also seek input from outside the scientific community, from hunting organisations and government agencies.
The Taxonomy Working Group needs to meet quickly to organise its philosophy/approach and to determine a division of labour. Its first tasks must be to establish guidelines and criteria, and decide the division of taxa among members. This may be best achieved through an international workshop on Caprinae evolution and taxonomy aimed at developing a recommended program for resolving taxonomic uncertainties pertaining to wild Caprinae populations.
2) Conservation of Maximum Genetic Diversity. While it is clear that Caprinae taxonomy is problematic, taxonomic confusion must not prevent or delay conservation. Our aim is maintenance of maximum genetic variation, not merely conservation of "taxa". It will not be sufficient to preserve just 1 or 2 "representative" populations of a given taxon, or even 1 or 2 populations within a country. We must ensure that a taxon's broad geographic distribution is well represented through our conservation actions, to effectively maintain current levels of potential genetic variability along with the supporting habitats.
Taxonomy is an area where regional co-operation can be vital. International co-operation and concerted action can ensure not only that adequate samples are available, but that the necessary wide geographic ( genetic) representation is preserved. The alternative, for countries to address their own Caprin conservation issues in isolation, will most probably lead to an unsatisfactory, piece-meal solution that fails to address the real conservation issues.
12.2.3 Maintaining Genetic Integrity
At least 3 issues are involved. First, for some Caprinae, threats to genetic integrity exist because non-indigenous taxa, often closely related, have been introduced into their range. Examples include Alpine chamois introduced into Chartreuse chamois range in France, and into Balkan chamois range in the former Yugoslavia; European mouflon into chamois range in France and Italy; and Barbary sheep into some desert bighorn ranges in the USA. The dangers arise from hybridisation, increased competition, or both. If hybridisation occurs, even between subspecies, unique gene pools adapted to local conditions are lost. Second, wild species are sometimes used for cross breeding in an effort to develop better adapted breeds of domestic livestock (e.g., domestic goat x Nubian ibex in Israel - domestic goat x Asiatic ibex and markhor in Pakistan). If successful, such crosses can only threaten the genetic integrity of the wild taxon. Last, are cases where "re-introductions" have been made without using animals of the appropriate taxon (e.g. muskox from Greenland into Alaska, Alpine chamois into several European countries).
A summary of discussion on the taxonomy of mountain ungulates and its conservation implications.
Ankara, Turkey, May 8-10, 2000
Prepared by Marco Festa-Bianchet
Readers please note that this summary is mostly based on my own notes and recollections of the discussion, although I have asked for input from participants to the workshop. It is posted here for 3 reasons:
To report some of the main points of agreement and disagreement that emerged during the discussion.
To assist in identifying priorities for future work on the taxonomy and conservation of Mountain Ungulates
To provide delegates to the workshop and other caprin taxonomists the opportunity to contribute to this summary by reminding me of points that I have missed, by sending me copies of their own notes, and by participating in the electronic discussion group.
This document will be updated as I receive input from IUCN Caprinae Specialist Group members, other participants to the workshop, and taxonomists or other biologists that did not attend the workshop but have knowledge or opinions that they wish to contribute. I will post additional or dissenting opinions that will be forwarded to me by scientists interested in Caprin taxonomy. Contribution to the discussion that direct attention to new published data will be particularly welcomed.
This summary has neither been edited nor reviewed, it may contain misinterpretation of what was said, most of it is based on opinions that may or may not be backed up by solid taxonomic data, it has not gone any kind of peer-review process and it definitely SHOULD NOT be cited as a scientific publication. All taxonomists and conservationists, including delegates to the workshop, are encouraged to publish their data and ideas in the appropriate peer-reviewed journals. In addition, information or results that are of local interest, reports on initiatives, progress reports and reflections on Caprinae taxonomy are always welcomed in the IUCN Caprinae Newsletter.
I thank David Shackleton, Marinus Hoogmoed, Amjad Virk, John Wehausen, Paul Weinberg and Elsa Gagnon for comments on previous drafts of this document.
There were 3 parts of our workshop discussion, in addition to formal presentations and posters:
1. A general discussion of the implications of taxonomic problems for the conservation of mountain ungulates
2. A consideration of how to produce an identification guide for Caprinae.
3. A specific discussion of the taxonomic of Caprinae.
Taxonomy and CITES
Dr. Marinus Hoogmoed, Chairman of the CITES Nomenclature Committee, provided some information on how scientific nomenclature is used by CITES and how it can be modified. This is a very important point because the Conference of Parties to CITES must agree to adopt changes to the currently accepted taxonomy if those changes are to be part of this international agreement. Dr. Hoogmoed pointed out that CITES takes very seriously the recommendations provided by IUCN/SSC Species Specialist Groups and would very much appreciate and support any initiative by the Caprinae Specialist Group in producing an identification manual.
CITES currently uses the taxonomy used in Wilson and Reeder as their standard reference for mammals. CITES is open to changing its official taxonomy in light of new findings. To change the official taxonomy, there needs to be a request to the CITES nomenclature committee, which must be supported by published evidence. Changes accepted by the nomenclature committee are then recommended, through the CITES secretariat, to the Conference of Parties that then decides whether or not to accept the new taxonomy. The level of consensus among taxonomists over the proposed changes would be a very important factor affecting the decision. The Nomenclature Committee revises the taxonomic list used by CITES every 2-3 years and proposes changes to the Conference of the Parties, based on new taxonomic publications - their recommendations can vary from reconsideration of a genus to reclassification of a specific population.
The Wilson and Reeder text does not have subspecific designations. This is a particularly important problem for Caprinae, where in many cases different subspecies of the same species have widely different conservation status. David Shackleton, editor of the 1997 IUCN Caprinae Action Plan, mentioned that the Wilson and Reeder text was of little use in preparing that document. CITES could adopt any other standard reference for subspecific designations, but will not develop its own list of subspecies. Again, the key issue here is good, published taxonomic evidence, accepted by experts in the field. Gradual or small changes from the currently accepted CITES taxonomy are more likely to be accepted by the Conference of the Parties than large-scale revisions of taxonomic status. It would also be possible for CITES to adopt a web-based list of taxa, as long as it was supported by published evidence and enjoyed widespread support among taxonomists. A web-based list of Caprinae taxa is a potential product that the Caprinae SG Taxonomy Working group may consider working on, possibly including the production of a CD-ROM.
Development of an identification guide
There are problems with identification of Caprinae taxa at customs offices. Some protected taxa may be reported as something else and go through customs undetected. Therefore an identification guide to trophies to help customs agents would be very useful.
To be useful, such an identification guide should be simple and should be made for use by non-experts. It may refer customs agents to seek additional input in difficult cases, rather than attempt to identify all taxa. Any guide to taxonomic identification of Caprinae should explicitly point out what can be identified based on different amounts of available material (horns, skull, cape, entire specimen etc.) and what level of identification requires the consultation of an expert.
An identification system for big-game trophies, specifically aimed at customs agents, was the goal of a project partially financed by Environment Canada. It appears, however, that funding for this project was cut before its completion. There was a strong consensus among participants that this was a very worthy project, much of the required work has already been done, and there should be strong support for its completion. The delegates were particularly impressed with the partially finished guide already produced by Elsa Gagnon and her collaborators.
The Office of Enforcement of Environment Canada is currently looking for partners to help fund the development, printing and distribution of the guide to trophy identification. The funds that Environement Canada allocated to this project are limited, and the growing demand for such guides increases the development, printing and distribution costs. For this reason, organizations who share an interest in this project are invited to take the challenge of making this guide a reality by funding, distributing it, or contributing in other ways.
Tissue sample from capes or other body parts should be collected in problematic cases, along with a series of pictures of the specimen. Much recent advance in Caprinae taxonomy is based on molecular techniques, and DNA analyses could help with the identification of difficult cases.
There is an urgent need for photographic documentation of several wild Caprinae taxa. In particular, very few pictures exist for females of some species, and most of the available pictures are of trophy-hunted large males. Consequently, little is known about color variation in other sex-age classes, or on all sex-age classes outside the hunting season. Genetic profile data are also required for many subspecies.
The guide book to hunting trophies that was being developed with the support of Environment Canada was meant to be a broad guidebook suitable for use by customs officials. It first tried to show how to identify several groups, including when only the horns are available, as is often the case with hunting trophies. These are some of the problems Elsa Gagnon encountered while working on this guide:
Capra - the only CITES-listed species is the markhor, which is relatively easy to identify. Nevertheless, there is much variation in markhor horn types, and they all need to be shown in the guide. If the horns are removed from the skull, the direction of the spiral can be difficult to determine. The Chiltan goat is a taxonomic problem: is it a markhor (falconeri), a wild goat (aegagrus), or an hybrid? (see taxonomic discussion of Capra below). Because all markhor and the Chiltan goat are currently listed in Appendix I, however, this is not a major concern for CITES at this time.
Rupicapra - the genus is easy to identify, but the species (rupicapra or pyrenaica) are difficult or impossible to identify unless the cape in winter coat is available.
Goral and serow species are very difficult to identify from just the horns.
Ovis - While the argalis (O. ammon) as a species are easy to identify from the horns, subspecific identification is very difficult and relies mostly on horn measurements. The subspecies nigrimontana and severtzovi are particularly difficult to identify, and small or immature specimens present additional problems. Some subspecies can be easily identified from the cape in its winter coat, but not in summer coat.
While a guide for taxonomic identification would find its primary conservation use for customs agents and wildlife law-enforcement agencies, it could also be useful as a field guide for ecotourism operators, and would illustrate the diversity of Caprinae to the general public. Several delegates pointed out that an identification guide should not only be seen as a law-enforcement tool. It would be very useful for conservation and hunting organizations as well, and it would be of limited use at least in North American courts where an expert witness would not be considered as such if he/she had to rely on an identification guide..
The Conseil International de la Chasse has been working for several years on the production of a World Atlas of Caprinae that should include pictures and distribution maps of most taxa. The IUCN Caprinae Specialist Group strongly supports this initiative. Despite a great amount of work by a great number of dedicated people, however, that project has not yet been completed.
In addition to a means to identify specimens based on morphological features, it would be very useful to have information on the amount of variability in characters both within and between taxa. Such a study would also allow identification of the most useful diagnostic characteristics and provide a guide to the reliability of taxonomic identifications. Local adaptations, genetic variability and environmental influences lead to a large amount of within-subspecies variability in morphological characteristics of many Caprinae; therefore there is much phenotypic variation that is not useful for taxonomic identification, and there is a strong need to associated taxonomically diagnostic features with genetic differences among taxa.
Other points that were raised regarding identification:
- A program to extract and coordinate analysis of tissue samples from harvested animals is urgently needed.
- The Taxonomy Working Group could write forensic descriptions for the identification of those taxa that can be reliably identified, and list those whose identification is currently difficult or impossible.
- There is a need to explore the forensic side of several taxonomic issue, to learn what types of data and statistics will hold up in court.
- There should be a prioritized list of problem taxa: some are difficult to identify but do not face great conservation threats at the moment. The IUCN Caprinae Specialist Group should elaborate a prioritized list of taxa for which taxonomic research is urgently needed. An IUCN-sanctioned list could assist national and local governments and NGOs in setting research and funding priorities. Several of these priorities are presented in bold in the taxonomic discussion below.
- Results published in peer-reviewed journals are the only kind of evidence that is useful for both taxonomy and conservation, and must be the ultimate goal of all taxonomic research. Published evidence is also much more useful for legal cases.
- There are differences between scientific and hunting taxonomies. Although in most cases there is broad overlap, hunting record categories have been established for a number of forms that are not recognized by taxonomists. There was a general consensus among the delegates that scientific taxonomy must be based on genetic differences: subspecies cannot be simply recognized on the basis of environmentally-induced morphological differences, or differences in horn size. Those differences, however, can be justifiably used for classification and ranking of hunting trophies.
- Better knowledge of Caprinae taxonomy is required to help guide future reintroduction efforts to use the animals best suited for any particular site. If Asian countries enter into a reintroduction phase such as has occurred in North America, how will they choose stock for translocations? It was noted that in Pakistan the current emphasis is on saving what is left rather than on restoration. Rich Harris pointed out that translocation of sheep has already occurred in Mongolia. Translocations may be driven and funded by hunting interests and taxonomic guidelines would help chose translocation stock. Dr. Hoogmoed noted that the IUCN reintroduction specialist group has already put out general guidelines.
- There was a brief discussion of what taxonomic level should receive priority. There seemed to be consensus that subspecies would receive priority. It was suggested that the concept of Evolutionarily Significant Unit (ESU) be considered as a criterion for subspecies designation, but the ESU concept requires substantial genetic data to establish reciprocal monophyly. Because these data are lacking for many caprinae taxa, it is necessary to make the best judgements based on current information. Gordon Luikart suggested that taxa should be ranked by evolutionary significance in the sense of a risk assessment so that a cost/benefit analysis would identify priorities for conservation/research efforts.
Taxon by taxon discussion
The participants in the workshop examined the taxonomic list of Caprinae provided on pages 10-14 of the 1997 IUCN Caprinae Action Plan, to identify points of agreement, of disagreement, and of lack of adequate knowledge.
One important general point that was raised by several delegates is that we should not underestimate the potential anthropogenic influences over the current distribution of Caprinae. Most conservationists are aware that in recent years there have been various attempts (successful and not) of introducing and reintroducing different Caprinae into different areas, including some inappropriate introductions of exotics that have led to problems in conservation and may have affected the genetic make-up of different species or populations. People, however, have co-existed with Caprinae for hundred or thousands of years, during which there may (or may) not have been numerous possibilities for genetic pollution, including hybridisation with domestic forms, or artificial movements of wild or semi-wild individuals from one mountain range to another. Even without any active translocation program by wildlife agencies, freelance initiatives have led to Caprinae transplants. Therefore, current patterns of genetic and morphological differences are not necessarily devoid of anthropogenic influences.
For the goral (Naemorhedus) and serow (Capricornis ), the current taxonomy is in need of revision, and several delegates lamented the fact that so little work is being done on these species, possibly because they are not actively sought by foreign hunters. For example, the Japanese and Formosan serow (Capricornis crispus and C. swinhoei) may or may not be the same species, and some Naemorhedus subspecies appear to be mostly based on geography. Much of the subspecific designation of these 2 species is very dated and based on very few specimens they need to be re-examined with a larger sample size and modern techniques (morphological, genetic and statistical).
Delegates agreed that there is one species of Mountain goat, Oreamnos americanus, with no recognized subspecies. The phylogenetic relationships of Oreamnos with other rupicaprini (and other Caprinae), however, are unclear.
There was a general acceptance of the 2 species of chamois (Rupicapra rupicapra and R. pyrenaica), but some delegates expressed doubts about the validity of some subspecies, particularly within the R. rupicapra species.
The subspecific designations of the Barbary sheep, Ammotragus lervia, were doubtful, this is a particularly problematic taxon for conservation because several populations are threatened while others are possibly extinct, and therefore a knowledge of subspecific taxonomy is required to aid potential future reintroduction programs. The subspecies angusi, blainei and fassini may not justify subspecific status and should possibly all be reclassified as fassini. The subspecies omatus from Egypt may be extinct (although some may survive in captivity), while the saharensis subspecies is likely valid. It is unclear which subspecies was (or were) released into North America, where Barbary sheep are now a widespread exotic species, with large population in the southwestern USA and in Mexico. Exotic populations also exist in Spain.
For the blue sheep, Pseudois, there was disagreement as to whether the dwarf blue sheep (P. schaeferi) is really a separate species or just a population of P. nayaur that is small because of environmental constraints, not genetic differences.
The 3 species of tahr (Hemitragus jemlahicus, H. hylocrius and H. jayakari) are good species with no subspecies subdivisions.
The subspecific designations of the wild goat, Capra aegagrus, require clarification. The two subspecies aegagrus and blythi may not be sufficiently different to deserve subspecific status, and the Chiltan wild goat (C. a. chialtanensis) may be a subspecies of aegagrus, a subspecies of markhor, or a hybrid form between markhor and wild goat. Therefore the Chiltan goat urgently requires more taxonomic work. The Cretan agrimi (C. a. cretica) is a domestic form and should not be considered a subspecies of wild goat. A poster presented at the workshop further substantiated this idea. (During the field trip, the delegates worked hard and looked hard to see wild goats, but mostly failed to do so!)
The three subspecies of markhor (Capra falconeri falconeri, C. f. hepteneri and C. f. megaceros) were generally accepted, although they are not easily distinguished because there is much variation in horn shape within subspecies. The current occurrence of markhor is highly fragmented relative to its former range.
The Capra ibex group requires a major revision. All the workshop delegates with knowledge of Capra taxonomy were unanimous in their opinion that there are likely 5 or 6 different species (with significant geographic separation) that are now recognized as subspecies of C. ibex. Some of these subspecies may be full species and contain their own subspecies. These are:
Alpine ibex (Capra ibex), which survived extinction only in the Gran Paradiso National Park in Italy and has now been reintroduced into much of its former range in the Alps.
Nubian ibex (Capra nubiana), a warm-climate species. The walia ibex, found in a single, highly endangered population in Ethiopia, is either a subspecies of the Nubian ibex and should be referred to as C. n. walie, or is a separate species, Capra walie.
Spanish ibex (Capra pyrenaica, which may have 2 extant subspecies (C. p. victoria in the Sierra de Gredos and C. p. hispanica in the rest of Spain) and 2 extinct subspecies, including the Pyrenean ibex C. p. pyrenaica which disappeared very recently.
Asiatic ibex, Capra sibirica, which has a very vast geographic distribution and may include 3 different subspecies.
The Kuban or West Caucasian tur (Capra caucasica).
The Daghestan or East Caucasian tur, Capra cylindricornis, was already recognized as a distinct species in the Caprinae Action Plan.
While the official taxonomy now lists 4 wild Capra species (aegagrus, ibex, falconeri and cylindricornis), in reality there may be 8 or 9. Most Capra taxa are easily differentiated by body size, size and shape of the male's horns, and color patterns, although seasonal coat differences can be somewhat confusing. Most of the species differences coincide with major differences in habitat and are presumably the results of genetic differences, although they could also result from phenotypic plasticity. Vast geographic barriers have existed for a long time among the proposed species. Given the increasing threats to conservation of many Capra populations, and the increasing importance of these animals as a trophy-hunting species, a taxonomic revision is urgently required. Potential human influences in the current distribution and genetic make-up (including hybridization with domestic goats) must be taken into account, as most of these taxa have co-existed with pastoralists for hundreds or thousands of years.
There was much discussion of all Ovis species, and delegates unanimously agreed that much more taxonomic work is required for this genus, particularly for the Asian taxa. Many delegates underlined the urgency of this work, as several taxa are threatened with extinction over the short term. Differences of opinion over the taxonomy the Ovis group are currently causing a number of conservation problems, including a dispute over some CITES designations.
The 1997 IUCN Caprinae action plan lists one species, Ovis orientalis, for all mouflon and urial. Most delegates disagreed with that taxonomy, partly because of differences in the number of chromosomes: mouflons have 54, urials have 58.
The mouflon should be referred to as Ovis gmelini, with a possible 4 to 6 subspecies:
O. g. gmelini, the Armenian mouflon (possibly including the surviving population in Turkey, which however may be a distinct subspecies, O. g. anatolica). (The delegates very much appreciated the opportunity of observing these beautiful animals and their habitat during the field trip);
two subspecies restricted to Iran: O. g. isphahanica, the Esfahan mouflon and O. g. laristanica, the Laristan mouflon;
the European mouflon, O. g. musimon, which is the descendent of feral sheep and may not warrant its own subspecific designation. This sheep evolved as free-ranging populations in Sardinia and Corsica but has been widely introduced as an exotic in Europe, North America, Hawaii and elsewhere. Many of these exotic populations have apparently been subject to hybridization with domestic sheep.
The Cyprus mouflon, O. g. ophion, is also the result of an artificial introduction into that island and may or may not be a different subspecies. Some delegates pointed out that given that both the European and the Cyprus mouflons are descendants of domestic sheep, they should be referred to as O. aries.
The red sheep (listed as Ovis o. orientalis on page 13 of the IUCN Caprinae Action Plan) is a hybrid form found in Iran, the result of interbreeding of different mouflon and urial subspecies. In general, the taxonomy of mufloniform sheep in Iran is complicated by geographical and genetic gradients among subspecies, and different levels of hybridization among subspecies and species.
Some delegates felt that the scientific name for urial should be Ovis orientalis, others that it should be O. vignei. The Caprinae Action Plan uses orientalis, but that name may have first been used to designate the red sheep, which is a hybrid. Both CITES and the Conseil International de la Chasse use vignei, and orientalis may be a nomen nudum. The important outcome of the discussion was, however, that mouflons and urials are different species. The usual taxonomic rules should decide which specific name should take precedence.
Delegates generally agreed that there are between 5 and 7 subspecies of urial: arkal, bocharensis, cycloceros, punjabiensis, vignei and possibly blandfordi. Several of these subspecies are very difficult to distinguish, for example there are few differences between cycloceros and blandfordi, and the latter may be an ecotype, not a subspecies. In addition to problems with identifying different subspecies, some delegates pointed out that several local populations of urials may be currently classified to the "wrong" subspecies. For example the IUCN Action plan lists 2 disjunct groups of cycloceros urials in the Kopet-Dagh mountains.
Urials are clearly a high-priority group for further taxonomic research.
There was a general consensus that Severtzov's sheep in Uzbekistan is an argali and not an urial, and should therefore be referred to as Ovis ammon severtzovi. This would be a major change from the taxonomy that is currently accepted by most enforcement and conservation organizations.
There was considerable disagreement over the subspecific designations of argali, Ovis ammon. The majority of delegates felt that there are currently too many recognized subspecies, but there were some strong dissenting opinions. This is clearly one group where more taxonomic work is required.
There are 3 subspecies recognized by the 1997 Caprinae Action Plan list in the northern part of the argali range: ammon, collium and darwini. Many delegates felt that this subdivision is somewhat artificial, with morphological differences among populations due to environmental differences and not based on genetic differences and local adaptations. These subspecies are very difficult to identify without relying on geographic differences, which is not a valid or a useful taxonomic character. There is a need to further investigate this problem, and the darwini subspecies may have to be synonymized with O. a. ammon. A poster presented at the workshop supported the idea that there are very few if any genetic differences among these subspecies.
The subspecific status of Tibetan argali is particularly controversial. Most delegates were of the opinion that it should be referred to as O. a. hodgsoni, but some believed that a different form, O. a. dalai-lamae, should also be recognized. This is clearly an urgent problem for conservation, because some people have suggested that the dalai-lamae designation is used to circumvent CITES restrictions on exports of hodgsoni specimens.
The Shansi or Northern Chinese argali, O. a. jubata, is apparently extinct.
The remaining 3 subspecies were generally accepted as valid by delegates: Tien Shan argali, O. a. karelini, Marco Polo argali, O. a. polii and Kara-Tau argali, O. a. nigrimontana. The Kara-Tau argali, however, is very difficult to identify based on morphological characteristics, and may be on the verge of extinction, possibly with fewer than 200 animals remaining. It is another taxon that requires urgent taxonomic and conservation work. Val Geist (in litt.) recently pointed out that "what I illustrated in my C.J.Z. article as nigrimontana, is actually collium. As best as I can make out, Russian animal dealers sold to the zoological gardens in Frankfurt, Munich and Oklahoma the common collium for the uncommon nigrimontana. There are quite a few colour photos floating around in which the Oklahoma ram or the Frankfurt ram are labled as nigrimontana. I discoverd the error when I was examining specimen of argali rams from Kaskhstan and noted to my unhappiness that they matched my nigrimontana descriptions and sketch. As to nigrimontana: there are to date no illustrations anywhere of a mature nigrimontana ram in nuptial dress."
The subspecific divisions of the snow sheep, Ovis nivicola, were not generally accepted by the delegates. Unfortunately, however, there were no taxonomic experts on this species taking part in the workshop, and several delegates pointed out that a considerable amount of taxonomic research on this species was available only in Russian. Existence of published information in languages other than English (particularly Russian and German) is a problem that could lead to work being duplicated or existing knowledge not taken into account in future research. The Caprinae Taxonomy Working Group will therefore strive to include members familiar with non-English scientific literature. The snow sheep is another species with a very large geographic distribution, with different populations (and possibly subspecies) that vary widely in conservation status. There are 4 subspecies listed in the Caprinae action plan (alleni, borealis, lydekkeri and nivicola), but the delegates had little to say about whether these 4 are all valid, or whether additional subspecies exist but have not been identified; alleni and lydekkeri may be the same subspecies.
There are two recognized species of wild sheep in North America. The bighorn sheep (Ovis canadensis) has been the object of much recent taxonomic research, including both genetic and morphological investigations. The currently accepted taxonomy has a number of problems, because subspecific classifications were based on small sample size and poorly-developed quantitative methods. Recent research combining morphological and molecular approaches has led to a substantially revised classification, supported by evidence published in peer-reviewed journals. The extinct Audubon's bighorn, once thought to inhabit river valleys to the East of the current distribution of bighorn sheep, is not distinguishable from, and has been synonymized with, the Rocky mountain bighorn, O. c. canadensis. Similarly, populations of "California" bighorn (O. c. californiana) in Washington and British Columbia have been reclassified as canadensis because they are not distinguishable by mithochondrial DNA or cranial shape. And, the region of Oregon, Idaho, Nevada, and northeastern California once considered California bighorn range, from which all native population have vanished, has been reassigned to O. c. nelsoni on the basis of a cranial shape character. The californiana subspecies is now restricted to the central and southern Sierra Nevada in California.
The number of subspecies of desert bighorn sheep has also been reduced in the light of recent work: cremnobates has been synonymized with nelsoni, while mexicana and weemsi (in Baja California) are still recognized, but are expected to be synonymized because of lack of evidence of distinction. The mexicana subspecies has disappeared from much of its former range, some of which is now occupied by exotic populations of Barbary sheep.
Clearly, the recent revision of the taxonomy of bighorn sheep has major implications for management and conservation, particularly in view of the frequent use of transplants as a management tool to reintroduce or even supplement populations in North America. The delegates recognized, however, that differences in subspecific designation are not the only criterion for deciding about the source of transplant stock. There are potentially important genetic differences that occur among populations of the same subspecies, and transplant programs must take those differences into account.
There was little discussion of Dall sheep, Ovis dalli taxonomy, as the 2 currently recognized subspecies (dalli and stonei) are easily distinguished.
The taxonomic status of ovibovini (muskox, Ovibos moschatus, and takin, Budorcas taxicolor) was only briefly discussed, partly because there were no experts of these taxa among the delegates. The four subspecies of takin should be re-examined with modern genetic and morphometric techniques.
Finally, the taxonomic status of the chiru (Pantholops) was briefly discussed: it is not clear whether or not this animal is a caprin.
Where do we go from here?
Two specific actions should result from this workshop:
1. Formation of the Caprinae Taxonomy Working Group (see below), which will coordinate sample collection and analysis, facilitate exchange of information among taxonomists and help in coordinating funding requests. This group will also provide the expertise to assist with taxonomic problems that may arise in various conservation activities for Caprinae, including CITES regulations, enforcement, transplants, identification of taxa in particular areas etc. A lack of coordination among researchers, and between researchers and hunting organisations, was frequently singled out as a problem during the workshop. Many delegates called for a system whereby tissue samples and morphological measurements are routinely collected from hunting or culling operations and made available for scientific study.
2. Continued work towards the production of a guidebook for the identification of Caprinae. Much work towards this goal has already been done by Elsa Gagnon with the support of Environment Canada, and a special effort will be made to ensure that project is completed. In addition, the IUCN Caprinae Specialist Group will produce a web-based depository of pictures and drawings of the various taxa.
Future planned activities include a workshop on the economic, social and biological consequences of trophy hunting, in cooperation with the IUCN Sustainable Use group, Safari Club International, and other interested parties, during the III World Congress on the Biology of Mountain Ungulates in Zaragoza, Spain, in June 2002.
Would you like to comment on Caprinae Taxonomy? Do you want to know what other people think?
Here is the place to do it.
TAXONOMY WORKING GROUP of the IUCN-SSC Caprinae Specialist Group
Coordinator: Gordon Luikart, Ph.D. Laboratoire de Biologie des Populations d'altitude, CNRS, University of Grenoble, BP 53, 38041 Grenoble, France Cedex 9
Working group objectives:
The IUCN Caprinae taxonomy working group will strive to promote (i) conservation and long-term viability of wild populations and their habitats, and (ii) the understanding of the systematics and evolutionary history of the Caprinae. To this end we will try to do the following:
- Facilitate communication and exchange of information among taxonomists to avoid repetitive work and to promote rapid scientific advancement (e.g., we will posted on our web site a list of DNA sequences and microsatellites being used in each species).
- Promote the publication of results in peer-reviewed journals so that data and conclusions can be used with confidence in conservation management and in forensic or law enforcement cases.
- Help find and publicize potential funding sources
- Coordinate sample collection and analysis (e.g., see Sampling Instructions... below)
- Establish tissue and DNA banks accessible to researchers conducting genetic studies
- Establish a global DNA genotype database for forensics and population structure studies, including a standard set of DNA markers for forensics and population studies
- Provide expertise and assist with taxonomic problems that may arise in various conservation activities for Caprinae, including CITES regulations, enforcement, transplants, identification of taxa in particular areas
- Promote better coordination between researchers and hunting organizations. For example to insure that tissue samples and morphological measurements are routinely collected from hunting or culling operations and made available for scientific study.
- Prioritize taxa and populations for study using criteria such level of endangerment and the cost/benefit of the study
- Follow the regulations of the Rio International biodiversity treaty. For example genetic material will remain the property of the country of origin and be used only for scientific research.
- Write forensic descriptions for the identification of those taxa that can be reliably identified, and list
those whose identification is currently difficult or impossible (It is immediately possible to conduct DNA fingerprinting and population assignment/origin tests in many Caprinae (e.g. for Ovis canadensis and Capra ibex ibex, the necessary data, statistical tests and DNA markers are available).
Members of the Caprinae Taxonomy and Conservation Genetics Working Group:
|Name & address||Species of interest||Expertise|
|Hsueh-Wen CHANG, Ph.D.
Department of Biological Sciences
National Sun-Yat Sen University
|Ovis ammon, serow..|
|Yu-Yu QUN, Ph.D.
Shannxi Institute of Zoology, Xi'an ??
|Vladimir F. SHAKULA, Ph.D.
Jabagly village Yuzhno-Kazakhstanskaya oblast '487964 Kazakhstan
|Ovis ammon...||Field biologist|
|Amjad Tahir VIRK, Ph.D.
Senior Biodiversity Specialist Biodiversity Unit
H. 26, St. 87, G-6/3, Islamabad Pakistan
|Markhor, Ibex, Urials||Field biologist, social-economic aspects of Caprinae conservation, sustainable use programs, Pakistan Caprinae information|
|Kate BYRNE, Ph.D.
Institute of Zoology
Zoological Society of London
London NW1 4RY UK
|All Ovis||Molecular systematics and populations genetics, sequencing mtDNA, microsat analyses|
|Raul VALDEZ, Ph.D.
New Mexico State University; Dept. Fishery &Wildlife Sciences; Box 30003; Dept. 4901; Las Cruces; NM; 88003; USA. 1/505/6463719; 1/505/6465975.
|Field biologist, systematics, evolution, behavior...|
|Dr. Paul I. WEINBERG
North-Ossetian State Reserve; Basijeva str. 3-8; Alagir; Severo-Osetinskaja (North Ossetia); 363200; RUSSIA,
|Field biologist, morphology, behavior, ecology|
|John D. WEHAUSEN, Ph.D.
University of California
White Mountain Research Station
3000 E. Line St.
Bishop, CA 93514
|O. canadensis||Morphometrics, multivariate statistics, population demography, genetics, microsats from feces|
|Rob R. RAMEY II, Ph.D. Environmental, Population and Organisimic Biology; University of Colorado; Boulder; Colorado; 80309-0334; U S A. 1/303/4928239; 1/303/492-8699.||All Ovis, especially O. canadensis||Morphometrics, multivariate statistics, molecular systematics, mtDNA sequencing, population genetics, microsats from feces|
|Victor ORLOV, Ph.D.
Russian Academy of Sciences
Institute of Ecology &Evolution
Laboratory of microevolution
Leninsky prospect, 33
Moscow, 117071 Russia
|Chromosomal genetics, Biometrics, Phylogenetics|
|Ettore RANDI, Ph.D.
Istituto Nazionale per la Fauna Selvatica
Via Cà Fornacetta 9
40064 Ozzano dell'Emilia (BO)
Tel: (++39) 051 6512 111
|All Caprinae, especially Chamois||Molecular systematics, mtDNA sequencing|
|Gordon LUIKART, Ph.D.
Lab. des Pop. d'Altitude, CNRS UMR 5553, University of Grenoble, BP 53, F-38041 Grenoble, France Cedex 9
Tel: 33 4 76 63 56 07
Fax: 33 4 76 51 42 79 Gordon.firstname.lastname@example.org
|All Caprinae, especially Capra sp. and O. orientalis||Molecular systematics, mtDNA, Y chromosome, growth hormone & other gene sequencing, population genetics, microsats from feces, forensics statistics|
|Bonnie C. Yates, Ph.D.
Sr. Forensic Specialist
US Fish & Wildlife Service
National Fish & Wildlife Forensics Lab
1490 East Main Street
Ashland, OR 97520
Tele: 541-482-4191 FAX: 541-482-4989
|All Caprinae and other mammals||Forensic mammalogist|
|Sandro Lovari, Ph.D.
Gruppo di Etologia e Ecologia Comportamentale
Dip. di Biologia Evolutiva, Univ. di Siena
Via P.A. Mattioli 4, I-53100 Siena, Italy
tel. +390 577 232955-4
fax +390 577 232825
|Rupicaprini, Hemitragus spp.||Behaviour, Biometrics, Phylogenetics|
*Please contact G. Luikart or Marco Festa-Bianchet if you would like to be part of this working group.
Instructions for collecting Caprinae tissue for DNA extraction. 20/8/00
using a clean knife blade to avoid cross-contamination between samples.
Note: Sample should be collected and stored in duplicate if possible to avoid loss (e.g., if one set is lost in the mail).
IUCN Caprinae SG page
Last update - 01/05/02
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