Information arising from the International Committee on Hybrids

UK Chair: Dr Nick Fox

Publication date : January 1999

Revised May 2004

THE USE OF EXOTIC AND HYBRID RAPTORS IN FALCONRY

Information arising from the International Committee on Hybrids

UK Chair : Dr Nick Fox

Publication date : 18 November 1997

1 Introduction

2 The issue

3 How do we assess invasiveness?

4 Are raptors invasive?

5 Case studies of interbreeding attempts in the wild

6 Genetic implications of hybrids on wild gene pools

7 The scale of production and use of hybrids

8 Why hybrids and exotics are used

9 The identification of hybrids

10 Political issues

11 Possible options to reduce risk

12 The technicalities of sterilisation

13 Legislation in relation to hybrids and exotics

13.1 International/Pan-European legislation

13.2 National/UK legislation

14 Acknowledgements

Appendix 1 List of participants and contributors.

Appendix 2 Extracts from ‘The Regulation and Control of the Release of Non-Native Animals and Plants into the Wild in Great Britain’, Department of the Environment 1997.

Appendix 3 NAFA policy on hybrids 1978.

Appendix 4 The RRF policy on hybrids 1997.

Appendix 5 Letter from Robert A. Witzeman, m.d. of the Maricopa Audubon Society to Harold Olson, NM dept. of game and fish. 1982.

Appendix 6 Extract from: recommendations of the Colorado Wildlife Federation task force on raptor law and regulation. 1984.

Appendix 7 The use of exotic peregrines to repopulate lost range. Tom Cade. 1980.

Appendix 8 Discussion on parrot hybrids - Catherine Quinones. From the www. 1997.

Appendix 9 Some Species Definitions - notes to a paper by R.M. Zink given at the Linnean Society, March 1996

1. INTRODUCTION

1.1. This is a discussion document designed to help us clarify our thoughts on these issues. It arises from submissions made (1996 & 1997) by the UK Hawk Board on the issue of introductions of non-native species under the Bern Convention, from concern about the DFO’s attitude to the use of hybrids in Germany, and from discussions at the NAFA Board meeting, November 1996 in which NAFA decided to establish an informal international working group to tackle the issue.

This is the first time that falconry has tried tackling an issue on an international, non-allied basis using the internet as a medium for a discussion group. All participants have had their own pressures of other work, also it has been impossible, because of the large numbers of documents, to properly include those with a fax but no e-mail. The difficulties have also been compounded by having two chairmen and, because of the pressure of completing this document for a deadline of 23 November for the NAFA meet in US and the DETR Wildlife Inspector’s conference in UK, we have at the last minute, decided to keep the two documents separate. They do not overlap unduly and have their own flavours. Interestingly, both documents reach very similar conclusions, and both chairmen are still talking to each other!

1.2. The Mission Statement from NAFA is as follows:

a). To discuss the variety of issues related to the production of hybrid raptors and their use in the sport of falconry, and to achieve a level of consensus amongst falconers internationally regarding these issues.

b). To recommend a set of standards to the NAFA Board of Directors that might serve to govern the production/use of hybrids in the interest of maintaining the value added to our sport by hybrids, while maximizing opportunities to protect the integrity of native raptor populations and benefit raptor conservation.

This document, compiled by the UK Chairman from submissions from contributors (see 1.3) fulfills part (a) of the mission statement above. Although we have suggested some options, we have not made any recommendations as requested in ( b).

1.3. During 1997 a list of people (Appendix 1) have participated or contributed to this discussion, chaired by Dr Steve Sherrod (USA) and Dr Nick Fox (UK). As chairpersons, we have as far as possible compiled material and presented blocks of text verbatim from contributors without editing. However we have had to both present a structure and summarise arguments: and these derive from a submission made to the UK Government by the British Hawk Board in September 1997 which successfully defended us against proposed restrictions on the use of non-indigenous species and hybrids.

1.4. This document attempts to clearly present information and data, but, contrary to NAFA’s second Mission point, and pro the UK Hawk Board and IAF’s requests, we have not attempted to outline policy. As producers of hybrids, both Chairmen recognise that we may be perceived to be biased on this issue, and we also recognise that the situation with respect to this subject differs in each individual country. Therefore we present the material, but leave it to each club or country to formulate its own policy, bearing in mind, as Christian de Coune warns, the potential precedent, for better or worse, a national policy may have on other countries. We have seen this happen with Germany.

1.5. We assume the following three tenets:

a). We support as a first priority, the sustainable future health of wild raptor populations.

b). We support as a second priority the freedom of individual falconers to practice legal and sustainable falconry.

c). We support as a third priority the freedom of individual falconers to decide if they wish to fly a hybrid or non-indigenous raptor.

2. THE ISSUE

2.1. Some falconers object to the use of hybrids on the basis of ‘purism’. To the outsider the issue is not simply the use of hybrids in falconry, but the possibility of non-indigenous raptors becoming established in the wild, either as pure populations or by hybridising with indigenous species. This is a much wider subject which carries a potentially major impact on falconry. For example, in the UK the Harris Hawk Parabuteo unicinctus and the Redtailed Hawk Buteo jamaicensis are the mainstay species of the ‘average’ falconer. To lose these species would be a major blow for the sport. The Convention on the Conservation of European Wildlife and Natural Habitats (the ‘Bern Convention’) had several times verged on classing falconry as a ‘release’ of non-native organisms, and we have had to fend this off at different legislative levels. Now it is accepted that falconry is not a ‘release’ because falconers intend to get their birds back and use various techniques including training and telemetry, to achieve this.

2.2. Some countries or states allow only the use of native species in falconry, others the use solely of non-natives. Some are more concerned about the risk of introductions, others about the risk of thefts from the wild.

2.3. The basic premise is as follows: that the release, particularly the unplanned release, into the wild of non-native organisms is deleterious to the indigenous ecosystem and its species components.

2.4. There are examples in which organisms have been invasive, have ecologically outcompeted equivalent indigenous species, or have impacted on inadequately adapted prey populations. The major impact of invasive organisms has been on isolated endemic ecosystems which tend to be less competitively adapted than continental systems. Obvious examples include mammals such as rabbits, foxes, cats, dingos, horses and camels outcompeting indigenous marsupials in Australia. Smaller islands have been drastically affected by cats, rats, goats, pigs or rabbits. In ecologically sensitive situations such as these it is clear that the introduction of exotics poses great risk and that the basic premise holds good.

2.5. Continental ecosystems are, or have recently been, geographically linked. There is no sea barrier between Cape Town, Bergen and Hong Kong. These ecosystems are intrinsically more robust and competitive. It is less easy for exotic species to become invasive, although there are examples, such as the Rhodendron in western Scotland, and the Japanese Knotweed. Examples of competitive introductions or interbreeding in New Mexico (Witzeman 1982, see Appendix 5) include European Starling, Rainbow Trout interbreeding with Apache Trout and Rio Grande Cutthroat Trout, and dogs breeding with Red Wolves. However, over the years, many species have been deliberately or accidentally introduced by man and have become part of our flora or fauna. Most of these introductions, rather than displacing natives, have enriched the biodiversity of the ecosystem by exploiting new man-made niches. It is probable (certainly in Britain and Western Europe) that there are virtually no areas of land remaining which have not been modified by man to a greater or lesser extent.

2.6. The USA already hosts more than 2,000 species of imported plants, 2,000 species of insects and spiders, more than 140 land mammals, 70 species of fish, 90 terrestrial slugs and snails, and more than 200 microscopic plant pathogens. 29% of New England’s plant species are introduced.[1] In addition, many state wildlife agencies have for many years adopted an aggressive policy of releasing exotic game and fish species for sporting purposes.

2.7. While many of our British ‘wild’ animals are introduced, such as the Pheasant Phasianus colchius Red-legged Partridge Alectoris rufa, Rabbit Oryctolagus cuniculus and Sika Deer Cervus nippon, as are many of our trees, such as the Sycamore Acer pseudoplatanus, many more of our introduced species were introduced for agricultural production, forestry or horticulture. Our crops and farm animals, our timbers and our gardens, our entire so-called ‘natural’ British landscape, is dominated by introduced species. Attempts to deny or freeze this process, according to the premise of the Bern Convention, are doomed to failure. Rather, we must examine the process, avoid introductions of invasive and harmful organisms, but benefit from the use of other organisms and genetic material. The concept of an utopia in which one strolls through a pristine natural ecosystem is an idealised dream. If all introduced species were removed from UK agriculture and food production would collapse. For a wider discussion of this see McNeely, J. A. 2001. The Great Reshuffling: Human Dimensions of Invasive Alien Species. IUCN, Gland, Switzerland and Cambridge UK.

3. HOW DO WE ASSESS ‘INVASIVENESS’?

3.1. Let us take as the basic premise (whether or not we agree with its implications) a plant or animal establishing a new genetic population or strain breeding sustainably in an uncontrolled manner, for example, in the UK. What steps have to take place for this to happen, and what are the risks at each step?

3.2. Take first the situation of an exotic species establishing itself, for example the domestic cat Felis catus. We need to measure each of the following steps:

a) How many cats are allowed free range each year?

b) How many of these wander off each year?

c) How many of these survive to breeding age?

d) How many of these meet either another feral partner or a temporarily freed domestic one for breeding?

e) How many of these breed young?

f) How many of these young survive to breeding age?

g) How many of these young breed?

The numbers in each step can be reduced by various means, for example by marking each cat with its owner’s name to enable it to be returned, by killing stray cats, by neutering and releasing feral cats to hold territories; and the risks are also modified by other factors such as provision of food for feral cats, competition with existing cats and by differential survival rates of progeny in comparison with existing cats.

3.3. What about an exotic species inter-breeding with an existing species and introducing exotic genes into its gene pool? Take for example the domestic cat and the Scottish Wild Cat Felis sylvestris:

a) How many cats are allowed free range each year?

b) How many of these wander off each year?

c) How many of these survive to breeding age?

d) How many attempt to breed with a Wild Cat?

e) How many Wild Cats attempt to breed with Domestic Cats?

f) How many pairings produce young?

g) How many young survive to breeding age?

h) How many of these are fertile and breed young?

i) What is the differential production and survival of cross-bred cats through genetic compatibility or physical adaptiveness compared to pure Wild Cats?

j) What is the ratio of genetic challenge in terms of the size of the Domestic Cat population compared with the size of the Wild Cat population in the areas where they meet?

3.4. These factors apply to all free-living populations, which is why animals and plants are found in certain areas, but not in others. In their own area they are able to sustain their breeding populations, but outside their own range these attrition factors cause the organism to die off. If the factor is temporary, such as winter, bird species may migrate and survive to return again to breed in spring: otherwise they cannot survive. Therefore most species which are introduced outside their natural range, die off. Some of them, such as most of our agricultural species and garden species, survive through constant support and management by man. A field of wheat is not a stable ecosystem. The wheat would quickly be outcompeted by other species until in a few years none would remain. These species, in these places, are non-invasive.

3.5. On the other hand, a few species, usually those from competitive continental ecosystems entering less specialised island ecosystems, can out-compete indigenous species or utilise unfilled niches in the ecosystem. These species, in these places, are invasive. The Caucasian race of H. sapiens has been invasive in many parts of the world, establishing breeding populations either where none previously existed or by supplanting and outcompeting indigenous populations. Therefore the concept of invasiveness depends not just on the species concerned, but also on the place.

3.6. The breeding strategy of a species is also a major factor in its potential invasiveness. At the simplest level, an asexually reproducing organism has the best invasive potential because a single individual could start a new colony. Species which do not establish a strong pair bond, such as cats, have mating strategies which tend to be competitive, indiscriminate, and therefore more easily accomplished. With raptors, which often have a long courtship and nest building phase and form strong pair bonds, both potential partners have to accept each other as a mate and even if the exotic newcomer makes overtures, if the indigenous partner does not accept him or her, then pairing will fail. The implications of this are seen in the case studies below. In order for successful breeding to occur it is also necessary for two birds of the opposite sex, with a suitable nest site, and suitable habitat, to all come together at the same time. If someone in Ohio loses a female Lanner falcon Falco biarmicus and a male is lost in Oregon, the chances are that they will die before they meet. In other species, such as ducks, an individual exotic duck can join into an existing flock and contribute genes fairly promiscuously.

3.7. Therefore, it is clear that certain conditions have to be met for a species to become invasive. Blanket criticisms of raptors on the basis of their potential for invasiveness are not only ill-founded but are also hypocritical. It is certain that the critic will either have eaten or worn material from introduced organisms or have them in their garden or house.

4. ARE RAPTORS INVASIVE?

4.1. We will use the UK as an example for this general discussion.

4.2. Firstly, many species of birds of prey are flown in the UK, and hybrids are also flown. Each of these poses different risks and must be evaluated independently. For example, while humans are invasive in UK, chimpanzees Anthropopithecus troglodytes are not. Of the genus Falco, the Gyrfalcon F. rusticolus, the Saker F. cherrug, and the Lanner are the most commonly flown exotic species. For climatic reasons, and through the lack of a small mammal prey base, these species are not indigenous to the UK. They have been flown here in falconry since the Middle Ages and no breeding has been known to occur. We can confidently say that from both biological and historical evidence these species are non-invasive here. Of 771 international introductions recorded by Ebenhard, none were of genera used in falconry.[2]

4.3. Some species, such as the Cooper’s Hawk Accipiter cooperii, occur in similar ecotypes in the USA to those found in the UK. Could it breed here? In the USA, it occupies a niche mid-way between the larger Goshawk Accipiter gentilis atricapillus and the Sharp-shinned Hawk A. striatus. Interspecific mechanisms prevent breeding between these species and there, the prey base is sufficient for the Coopers.

In Britain, there is less habitat suitable for the Coopers Hawk; the Sparrowhawk A. nisus is larger than the Sharp-shinned Hawk and would compete with the Coopers’ for mid-sized prey. The Sparrowhawk is also a widespread and well-established native, whereas few Coopers Hawks are flown by falconers. Therefore the chances of introducing this species are infinitesimal, even if one mounted a major effort such as that made with peregrines by The Peregrine Fund in the USA. From experience with various support and/or reintroduction efforts of birds of prey into their own original range, it appears to require a protracted conservation effort to introduce a population of a certain critical mass for it to sustain itself. Until this point is reached, a withdrawal of effort, or even minimal persecution, will cause the potential population to die off.

4.4. There is not enough space in this submission to provide an evaluation of every single falconry species and its potential for invasiveness as new species in the UK. The point is that each species is different and policy should be handled on a species by species basis, not on a catch-all basis. Definitions of ‘species’ are given in Appendix 10.

4.5. But what of the risk of introducing exotic genes into an existing endemic species, as with the Domestic Cat inter-breeding with the Scottish Wild Cat or the Racing Pigeon with the Rock Dove? Examples might include the Red-tailed Hawk Buteo jamaicensis from the USA interbreeding with the closely related Common Buzzard B. buteo. One would suspect that of all the species flown in falconry, this would be the one case where interbreeding might occur. And yet, despite over thirty years of large-scale use in falconry in UK, there has been only one failed attempt at interbreeding (see case studies).

4.6. The British Goshawk A. gentilis gentilis, extirpated by shooters in the 19th century, was reintroduced to the UK by falconers with major releases of wild-trapped juveniles of Scandinavian and European stock. Although of the same species, these came from phenotypically different sub-populations. This genetically mixed stock is now gradually selecting for the optimum types for the British situation. A similar situation pertains with the Red Kite Milvus milvus in the UK and the peregrine in the USA.

Similarly, if a Spanish Peregrine Falco peregrinus brookei was released in Britain, and survived and bred with a British Peregrine, not only would its genetic contribution be very rapidly diluted out (50% at each generation), but also would be weeded out through natural selection, through failure to compete with the ecologically more optimal British stock. Many sub-species of Peregrine have been flown in Britain over the centuries with no noticeable impact whatsoever. A similar situation would prevail if a European Goshawk Accipiter gentilis gentilis were lost in North America and bred with a Northern Goshawk A.g.atricapillus.

A similar case exists with the hybrid falcons, for example the gyr/peregrine. These are even less likely to have a genetic impact, because both sexes possess markedly lower fertility both in the first generation and in the second than pure peregrines. The genetic selection against them is therefore extreme. They also have major difficulties in pairing because either potential partner will fail to respond to the other’s courtship signals: and ecologically they are even less able to compete with existing Peregrines because they are so far removed from the optimum wild type. The ease with which breeding raptors can be disrupted by human interference, while of long concern to conservationists, also demonstrates how easy it is to prevent interbreeding. With species which breed more privately, such as the Scottish Wild Cat, it is less easy to disrupt breeding attempts.

4.7. Where hybridisation occurs in the wild, it is through interbreeding of two established closely related populations, such as Domestic Cat/Wild Cat, Racing Pigeon/Rock Dove, Red Deer/Sika Deer, where there is a significant large-scale gene flow. Individual hybrid birds attempting to breed with a wild population are genetically outcompeted by the first generation.

4.8. Together with the biological hurdles that introduced birds would have to overcome, falconers add others: in compliance with Section 14 (3) of the Wildlife and Countryside Act, domestic birds of prey carry a ring and anklets to identify them as domestic and in most cases, to trace them to their owner. They are also trained to come back and supervised during the period of flying free. Most birds also carry radiotelemetry to assist prompt recovery. Falconers in the UK have supported the compulsory close ringing of exotics, but this was discontinued two years ago, and we would support the compulsory use of radiotelemetry on exotics. We take all reasonable steps under 14 (3). Do all other user groups match up to the responsible standards already set by falconers?

• · Individual identification
• · Trained to return
• · Supervision during release
• · Radio-tracking for control

4.9. Some of these discussion points will be examined in more detail in the following sections. (See 11.1)

5. CASE STUDIES OF INTERBREEDING ATTEMPTS IN THE WILD

5.1.

a). Red-tailed Hawk Buteo jamaicensis Western race with Common Buzzard Buteo buteo in the UK. John Murray. A female redtail was flown at hack, without parental imprinting, in British Columbia and then trapped - virtually as a passage bird - and sent to the UK. She was lost while being flown for falconry but survived on rabbits and settled on the Arniston Estate, 15 miles south of Edinburgh in 1969. As a third year bird she paired with a pale male Common Buzzard and nested in a solitary oak near the river. She was still recognisable by her jesses, and easily located but would not come to a trap. She laid 4 eggs but these were taken by crows and Mr Murray could not tell from the shells whether or not they were fertile. She moved away to another estate where she preyed on domestic chickens and ducks. She was out in total for two and a half years but got heavily into taking pheasants and the keeper shot her and Mr Murray stuffed her.

b). Peregrine Falco peregrinus male with female Prairie Falcon F. mexicanus in Colorado in 1949. Vern Siefert cited by Lynn Oliphant 1991. These were both completely wild birds.

c). Peregrine Falco peregrinus male with female Prairie Falcon F. mexicanus in Utah in 1986. Clayton White cited by Lynn Oliphant 1991. The male had been hacked out in a release programme.

d). Peregrine Falco peregrinus male with female Prairie Falcon F. mexicanus in Saskatchewan. Lynn Oliphant 1991. The male had been hacked out in a release programme without parents:

 

 

e). Peregrine Falco peregrinus female with male Gyr/peregrine F. rusticolus/peregrinus in Germany in 1996. Christian Saar and Walter Bednarek. The female was captive bred and hacked out without parents. The male was an escaped falconer’s bird. Two young were produced in the wild. The male was trapped and the young removed. DNA tests were made by Professor Wink, proving the male gyr/peregrine to be the father.

f). Peregrine Falco peregrinus female with male Gyr/peregrine F. rusticolus/peregrinus in Germany in 1997. Christian Saar and Walter Bednarek. The female was juvenile, unringed. The male had been lost at hack. They paired near Hamburg but the male was trapped without breeding.

g). Unidentified hybrid occupying nest box but not paired up. Belgium 1995 Christian de Coune.

 

h). Female gyr/peregrine paired with wild male anatum peregrine in Colorado 1996. See Steve Sherrod’s report. Eggs laid but did not hatch, fertility status unknown.

i). A. Corso and D. Forsman (1997) also cite the following examples of successful hybridisation in wild raptors

- Black Kite Milvus migrans and Common Buzzard Buteo buteo in Rome

- Male Pallid Harrier Circus macrourous and Female Montagus Harrier Circus pygargus Finland 1993

- Golden Eagle Aquila chryseotus and Imperial Eagle Aquila pomarina in Spain

Corso and Forsman also cite the following examples of non-successful breeding attempts:

- Male Pallid harrier Circus macrourous with Female Hen Harrier Circus cyaneus

- Male Pallid harrier Circus macrourous with Female Montagus harrier Circus pygargus

5.2. Of the above examples, one, (b) occurred in 1949 before the advent of captive breeding, and is an example of natural hybridisation, although it was not confirmed. It is probable that peregrines and prairie falcons in this area were nesting in close proximity in 1949. In examples (c), (d) and (e) the pure peregrines had all been hacked out in re-introduction programmes without parents. It is likely therefore that they lacked full parental imprinting. The male gyr/peregrine in (e), the subject of major controversy in Germany, was not (as an escaped falconers’ bird) the only factor in this pairing, because the ‘wild’ female was actually captive bred and hacked out without full parental imprinting. In the second case, (f), the female was juvenile and no breeding took place. It is possible that she was responding to the male as a supplier of food, i.e. as a parent rather than as a mate. In the third case in which a hybrid was involved (g), the bird refused to pair and drove away peregrines. In case (d) Oliphant reported 2 hybrid chicks hatching from a minimum of 5 eggs. When breeding pure chicks the female produced broods of 5 and 3. In h) the eggs did not hatch at all.The little evidence we possess shows that full species are as likely to mispair as hybrids and that inadequate parental imprinting in hacking predisposes birds to mispair. This was a factor in cases (a), (c), (d), (e) - both birds - and (f). Imprinting on saker parents was likely to have been a factor preventing pairing in case (g) but we have been unable to verify this.

5.3. These records represent the total number of global, verifiable attempts that we have traced. [Note, since this was written in 1997, Gene McCarthy has been documenting hybridization of birds for a book due out shortly. It contains long lists of hybrid birds of all taxa that have occurred in the wild, and in captivity, including inter-generic hybrids. It is clear that genetic isolating barriers are by no means as water-tight as previously thought by speciesists and that occasional genetic re-combination through hybridization , although rare, is a normal event in nature.] For raptors, these records are the total known genetic threat despite hundreds of exotic falconers’ birds being lost in many countries over the centuries. It is noteworthy that of these case studies, NONE include an exotic falconry species starting to colonise. There have been no nesting attempts between two individuals of the same exotic species. We can confidently say that in the last thirty years there has probably been more intensive monitoring of nesting large falcons than of any other taxon. It is not a case that nobody has been looking. Significant numbers of birds have been lost, but almost none have bred. These are the known facts. The evidence therefore is conclusive: exotic raptors used in falconry are non-invasive in the current situations. We will compare these data with examples of other taxa which are invasive:

6. GENETIC IMPLICATIONS OF HYBRIDS ON WILD GENE POOLS

6.1. Genetic mixing to existing native populations is not necessarily a bad thing. For example the Welsh population of Red Kites Milvus milvus suffer depressed breeding as a result of genetic impoverishment. Genes contributed by a German kite in the 1960’s have provided much-needed genetic vigour. New introductions have since been made of Spanish and Swedish kites and the kites now show improved brood sizes in UK.

6.2. For non-native genes to enter a wild population a number of barriers have to be crossed:

6.3.a. Behavioral barrier.

The introduced bird has to be sufficiently close taxonomically to produce viable offspring with a native individual. The more closely related the two birds, the more likely this is to occur. For example, a non-local subspecies of peregrine, such as a Spanish F. p. brookei in the UK, is most likely to interbreed successfully with local F. p. peregrinus peregrines.

Other, full species, are very unlikely to interbreed if they already exist sympatrically somewhere in the world. For example, gyrfalcons and peregrines exist together in many areas of the world and possess mechanisms which stop them even attempting to interbreed. Similarly, sakers and peregrines do not normally attempt to interbreed. However, where gyrs and sakers link in the wild, they do seem to interbreed and form an intermediate form or natural hybrid, the Altai falcon.

This barrier, which prevents breeding attempts, is behavioral. Where two species have only recently become distinct through geographical isolation, such as the red-tailed hawk and the common buzzard, the natural mechanism keeping them genetically distinct is geographical, rather than behavioral. If a red-tail is released into a wild population of common buzzards there appears to be virtually no behavioral or genetic barriers preventing inter-breeding. Having said this, considering the numbers of red-tails lost in UK each year (10+?), there are no authenticated records of inter-breeding except the 1969 record of a failed attempt. There are several sympatric species of Buteo in America and it may be their specific behavioural barriers are quite subtle.

6.4. The mechanisms by which this behavioral barrier operates are not yet fully understood. Calls and displays given by a bird during courtship appear to be of genetic origin, ie. they are instinctive, ‘hard-wired’ and unalterable. On the other hand, the image in the bird’s mind of what constitutes an appropriate partner to display to is, to a large extent, imprinted during the bird’s early life. For example, any species of falcon reared by a peregrine is likely to treat an adult peregrine as a potential mate. Any species of falcon reared by a gyr is likely to treat an adult gyr as a potential mate. Thus, by imprinting a young falcon on a human or any other non-native species, a behavioral barrier is created which reduces the risk of hybridisation with a native species. As humans are unlike any type of raptor, imprinting on humans is a possible way of creating a behavioral barrier against inter-breeding in the wild. Imprinting may also result in reduced survivability in the wild, further decreasing the probability of successful breeding attempts in lost imprint birds (see extract from letter from Bill Burnham, below:)

 

6.5. This barrier actually goes a step further. For a bird to attempt to breed with another, the mate must not only physically resemble the imprinted image, but must also respond with its own, correct courtship signals to stimulate pairing. Take for example, a gyr reared by a peregrine and lost to the wild. What would happen if it saw a wild peregrine and attempted to court it? The peregrine would be inhibited by the courting gyr through its own behavioral barriers. It would not therefore make courtship signals and copulation would be prevented. However, a gyr/peregrine, reared by a peregrine and later meeting a wild peregrine, might look and behave sufficiently like a peregrine for the peregrine to be stimulated and return its courtship signals, leading to successful copulation.

6.6. b. Genetic barrier.

We know that if the behavioral barrier is bypassed through using artificial insemination, these species can produce viable offspring. In other words, there is not a complete genetic barrier. Hybrids can be formed between the three species.

Contrast this with another example, the peregrine and the goshawk. These two species have a behavioral barrier preventing attempted breeding, but also, if one tried artificial insemination, no embryos would develop; there is a genetic barrier.

In our current state of knowledge, it seems likely that all of the species in the genus Falco can be hybridised by artificial insemination to produce first generation young. The close sub-groups in Falco, such as the gyr/saker/lanner complex, or the peregrine complex, appear to form hybrids within their complex which usually show full fertility over indefinite generations. The less closely related species, such as the gyr and peregrine, form hybrids which normally exhibit reduced fertility; the spermatazoa show a high percentage of deformities and some females are completely sterile. A high percentage of them show physical deformities, particularly skeletal and cardiovascular deformities, some of which appear to be fatal in the embryo stage. The result? In the wild gene pool, the genes from hybrids such as these would be diluted out through an inability to compete with pure wild genes.

6.7. Less related hybrids within Falco, such as between peregrine and New Zealand falcon Falco novaseelandiae, are possible, but the embryonic mortality is very high. We have no data yet on second generation fertility. Also we have no data on the possibilities of inter-generic hybrids in the Falconidae.

6.8. Inter-generic hybrids have been produced between Parabuteo unicinctus and Buteo regalis (harris and ferruginous hawk) and between Parabuteo unicinctus and Accipiter cooperii (harris and coopers hawk). We do not know what is happening at chromosome level in these cases, or about the fertility of such offspring.

6.9. c. Genetic attrition.

If the behavioral and genetic barriers are overcome and genes enter the wild gene pool, there is then the question of assessing gene flow and proportions. Although a hybrid individual may show ‘hybrid vigour’ in terms of energy, genetically hybrids perform less well than pure breds. Hybrid semen shows reduced fertilising capacity, hybrid embryos show reduced viability, hybrid chicks show increased physical defects, adult hybrids have a higher rate of partial or complete sterility. The overall result of this is that when hybrid genes enter the gene pool, not only are they heavily diluted by pure genes, (due to the size of the gene pool and to genetic turnover between generations) but also they are out-competed. Their performance is not as good as the pure genes and therefore their frequency in the population falls. This dilution rate could be calculated based on numbers and probabilities. Phenotypically too, the hybrid is less well adapted to the local wild conditions than the pure wild type and it will tend to be less well able to support a mate, or show higher mortality, or inappropriate migratory behaviour, to maintain its gene frequency. This is the mechanism by which populations maintain their wild type. For example the Rock Dove in the UK maintains its wild type in the face of massive genetic challenge from racing and domestic pigeons, through preferential selection.

6.10. Where a local population is already depleted owing to other factors, as is the peregrine in Germany, there may be less intense competition for nest sites and mates. A lost hybrid thus has more chance to be accepted and pair up in these circumstances than in a saturated population as in most parts of Britain.

 

 

 

 

7. THE SCALE OF PRODUCTION AND USE OF HYBRIDS

7.1. This is a global situation. Birds bred in Europe or North America may be sold in the Middle East and flown in Asia or North Africa. Birds from Asia are imported into North America. The risk is not necessarily in the country in which the bird is bred, but is in the country where the bird is flown free.

7.2. It may be useful to look at the frequencies of hybrids in captive collections and the reasons for them:

7.3. Hybrids between various buteos and accipiters are infrequently made because they are difficult to produce, have few obvious advantages as hunting birds, are probably sterile, and therefore do not attract much of a market. They will probably remain marginal curiosities, and of no significance to wild populations.

7.4. Hybrids between Gyrfalcon and Saker seem to occur in all gradations both in the wild and in captivity and may not merit the title hybrid as this group appears to be a super-species. In falconry this intermediate form is larger and faster than a typical saker and more heat resistant and disease resistant than a typical gyr. All gradations appear to breed readily and indefinitely in captivity. For details of this group see: Eastham, C.P. 2000. Morphological Studies of the Taxonomy of the Saker Falco cherrug Gray 1833, and other species. PhD thesis, University of Kent at Canterbury, UK.

7.5. Hybrids between Gyrfalcon and Peregrine are popular in Arab falconry because they are large and fast. When lost to the wild in desert areas they do not survive long because there is usually not a sufficient prey base of medium sized birds for their survival and they are slow to adapt to small mammals, having been trained solely for houbara. The falcons are flown sharp and probably have at most only three days of energy reserves in which to make a kill. Those which do not perish often come to humans and are eagerly trapped by locals thinking of reward money. However they are poorly cared for and without proper resuscitating treatment, also die. When we did deliberate large-scale (80+) releases of health-screened, fat, fit ex-falconry passage pure sakers and peregrines in hunting type habitat eg Baluchistan and the Gulf, large numbers died or were re-trapped. We have been more successful by releasing the falcons in Kirghistan where there are more birds and mammals, in April when the spring migrants are returning and the wintering mammals are emerging. The local peregrines tend to be small, desert adapted varieties: Red Shaheens F. p. babylonicus and Barbary Falcons F. pelegrinoides, unlikely to pair with big gyr/peregrines.

Arab falconers have learnt not to release hybrids and we have not heard of any hybrid being ordered to be released since 1995. Instead, Arab falconers have little qualms about having unwanted birds destroyed.

In Europe female gyr/peregrines are used for pheasants, ducks and crows and males for grouse, partridges and rooks. In America they are used for desert grouse and for ducks. In most ways they hunt like large peregrines.

7.6. The peregrine/saker is used primarily in Europe where it combines the speed of the peregrine with the tenacity and willingness to take prey on the ground of the saker. It is thus more suitable for hunting crows than the pure peregrine or saker.

7.7. The National Avian Research Center’s monitoring programmes for source breeding populations of Arab falconry peregrines and sakers have not revealed any recogniseable captive-bred or hybrid falcons at nests. However, we have discovered two breeding wild calidus peregrines in Siberia wearing Arab jesses.

8. WHY HYBRIDS AND EXOTICS ARE USED

8.1. Unlike wild raptors which may catch any species they wish to attempt, falconry raptors are restricted to certain legal species and seasons. Falconers themselves are restricted to certain hunting areas and these areas are becoming more and more crossed with man-made hazards such as roads, wires and encroaching urbanisation.

The falcon must be closely matched to the prey in order to achieve a successful flight, not only to a species but also to very limited types of habitat. Classic flights thus evolved, such as the peregrine at Red Grouse Lagopus lagopus, partridges Perdix perdix/Alectoris rufa or Rooks Corvus frugilegus, and the Merlin at the Skylark Alauda arvensis, all taking place in very open landscapes.

Most falconers do not live in these open landscapes. They cannot afford a grouse moor, and have to fly in more enclosed land. Many flights are ruined by prey escaping into cover. Therefore the falconer is looking for a falcon which can hunt in the open but which can also, if need be, catch prey at or in cover at the conclusion of an aerial chase. The falconer also seeks to hunt some quarry which, although legally available pest species, cannot be caught by pure-bred falcons in falconry on a regular basis. In UK the pest list includes the Carrion Crow Corvus corone, the Rook, the Jackdaw Corvus monedula, the Magpie Pica pica, the Collared Dove Streptopelia decaocto, the Feral Pigeon Columba livia, and the Woodpigeon Columba palumbus. Among the smaller birds are the House Sparrow Passer domesticus and the Starling. Among this list, certain hybrids have proven themselves capable of providing quality falconry in circumstances in which pure-bred birds would fail. For example the peregrine/saker is far more effective at crows and rooks in poor country than a pure peregrine. The New Zealand/peregrine F. novaseelandiae x peregrinus is one of the few falcons which can catch jackdaws, magpies and pigeons consistently in poor country. The peregrine/merlin F. peregrinus x columbarius can take starlings where a pure merlin would fail.

Thus in the artificial landscapes of modern Europe, hybrids allow falconry to continue in places and at quarry which were previously unfeasible. Bad farming practices now have made such an impact on the UK population of skylarks that traditional flights with merlins are becoming a very rare sight. Access to good partridge or rook hawking ground for peregrines is now so limited that for the falconer in lowland Britain the opportunities for flying a falcon are few, unless he has access to a suitable hybrid.

8.2. As more experience is gained with the different types of hybrids, certain types will establish their potential in falconry and others will be found to confer no additional benefit. The supply of hybrids will mirror the market closely because most are made by artificial insemination and can be changed from year to year.

8.3. In Arabia, large hybrids are in demand because they are larger, faster and more glamorous than pure breds, and because they are less prone to disease, seldom carry internal parasites (such as serratospiculum) and, being less stressed in captivity, moult better. Now that Arabs have learnt to manage them, these birds consistently outperform wildcaught falcons. I do not see this trend in Arab falconry as reversible; many Arabs have set up their own breeding programmes and even if western countries stopped producing hybrids, production would be taken over in Arabia or Pakistan or in the former USSR countries.

Since 1993, as the former USSR opened its borders, trapping pressure on wild sakers in Asia has increased tremendously and the National Avian Research Center in Abu Dhabi has supported local biologists to monitor this species. At present, because of the inbalance of the economies among Asian countries, we do not see any practical means on the ground for reducing this trapping rate. However, captive bred hybrids are now gaining a major share of the Arab falconry market. More than 50% of Sheikh Zayed’s falcons (out of an annual purchase of about 300 birds) are captive bred, mainly hybrids. We also monitor the markets in Pakistan and the Gulf : prices for wild falcons were down at least 50% in 1997 and many birds remained unsold. Thus captive breeding is the key element at present to reduce the market pressure on Asian wild falcon populations. [Note : in 2004, 40 Sakers in one batch in Pakistan, remaining unsold for falconry, were allegedly killed and stuffed for taxidermy]. At the Consultation on International Trade in Falcons for Falconry, hosted by CITES in UAE in May 2004, it was unanimously agreed that captive breeding should be actively supported in order to offset the trade in wild falcons.

8.4. The Arab market has stimulated large scale commercial production of hybrids which has had both positive and negative effects. While the availability of female hybrids for western falconers has decreased, the males, which are a by-product, are increasingly available and at prices well below production costs. Commerce has stimulated research and development into many aspects of producing birds for falconry, and in ancillary equipment such as radiotelemetry. This has produced spin-off for western falconers and for conservation. In terms of resources, it is more expensive to breed hybrids than to breed pure-breds and it should be borne in mind that the commercial breeder, from the business standpoint, produces hybrids not because he wants to, but as a response to market demand. If falconers did not demand hybrids, breeders would not continue to produce them. In 2003 and 2004, Hunting Falcons International euthanased 10% of its young hybrids, all at 5 days old. These were all small males for which there was no market.

9. THE IDENTIFICATION OF HYBRIDS

9.1. In the case of hybrids between peregrine, gyr or saker; these three species are themselves very variable and there are many specimens from the wild in Asia which cannot be positively identified as being either gyrs or sakers. To identify hybrids on a legal basis is even more difficult. Work on DNA and karyotyping continues but will probably never be definitive. This begs the question as to whether some hybrids between recognised species are actually hybrids or whether we should revise the taxonomic divisions within Falco. Watch this space!

9.2. For specialist interest we have morphometric data on the following birds at the Falcon Facility in Wales, together with colour transparencies and also quite a good collection of study skins of falcon hybrids:

9.3 Table 1. Data on hybrid falcons collected and stored at the Falcon Facility, Wales.

Last updated 14th November 1997. Father is first mentioned species.

Type of hybrid	Sex	Morphometric data	Blood for DNA analysis	Breeding data	Photographs
Gyr / Saker	M F	18 23	17 20	6 3	11 12
Gyr / Peregrine	M F	7 10	7 10	1	4 10
Peregrine / Gyr	F	1	1		1
Peregrine / Saker	M F	12 22	4		8
Barbary / Saker	F	2	2		2
Gyr (1/4) / Saker (3/4)	M F	11 19	7 19	2	2 10
Gyr (5/8) / Saker (3/8)	M F	1 1	1 1		2 1
Gyr (3/4) / Saker (1/4)	M F	2	2		1 1
Gyr (1/4) / Peregrine (3/4)	F	2			2
Gyr / Saker x Peregrine	M F	1 2	2		1
Gyr / Peregrine x Saker	F	1
Peregrine x Gyr / Saker	M	2	2
Peregrine x Gyr / Saker x Saker	F	1	1		1
Peregrine / New Zealand falcon	M	1	1		1
Gyr / Peregrine x New Zealand falcon	M	1	1		1
Gyr / Prairie	F				2

10. POLITICAL ISSUES

10.1. There are many pressures on falconry at present, with attempts to ban all fieldsports or to nibble away at them using legal restrictions. As falconers, our first responsibility is to the raptors both in the wild and in captivity. While we naturally desire to do whatever is of biological benefit for the birds, we have to be careful not to over-restrict ourselves and our children for the sake of political correctness, expediency or pressure. German falconry is already subject to heavy restrictions on who may breed and fly birds, what species they may fly and what quarry they may fly. This has resulted in no benefit to falconry in Germany and therefore we should not let similar restrictions be imposed on us. Rather we should fight for the future of falconry and resist political pressures.

10.2. While a restriction on the use of non-native sub-species of peregrine would not be a great loss to falconry, the removal of the redtail from British falconry would be a major blow, and the loss of falcon hybrids would create a much greater loss than most people currently realise. The loss of non-native hybrids, such as the gyr/saker in the UK, would be a major triumph to our opponents and achieve absolutely nothing in conservation terms. If action is deemed necessary it should be targeted at the biological problem rather than a catch-all approach which would do far more harm to falconry than the situation warrants.

 

11. POSSIBLE OPTIONS TO REDUCE RISK

11.1 As stated in 4.8, Falconers already:

· Mark their birds;

· Train them to return;

· Supervise them during the release period.

This is more than can be said for many user groups including cat owners, fish stockists, etc. For clubs supporting the use of exotics and hybrids, there are a number of options to reduce risk still further:

11.2 It would not be a major hardship if falconry clubs made it part of their codes of conduct that:

· Any raptor flown free should be marked with its owners’ name and telephone number;

· Any exotic or hybrid raptor flown free should carry telemetry.

11.3 In the case of allopatric hybrids it would be advisable for them to be imprinted on the absent parent species: e.g. gyr/peregrines flown in Britain should be imprinted on gyrs or sakers rather than on peregrines; failing that on humans or any other species than peregrines.

Incomplete parental imprinting appears to have been a key factor in most of the case studies. It appears that the future identification of sexual partner depends on exposure to a parent figure, not just during fledging, but also during the flying dependency period. We have found occasionally, when sending hacked falcons to Arabia, that certain individuals started to scream. Analysis discovered that these birds had been reared by peregrines and that they screamed when placed on a perch near an adult peregrine. Those which had been reared by sakers did not scream in the presence of an adult peregrine, or in the presence of an adult saker. Adult sakers do not have a significantly different plumage to juveniles, as peregrines do. There may be something going on here, especially in the imprinting process of peregrines, which would repay more careful study and documentation. We are conducting trials here in UK on imprinting captive bred falcons and the subsequent sexual signals which act as behavioral triggers. This is being done using models.

Bearing in mind that the risks involve not just hybrids, but also exotics, a blanket legal requirement to imprint these groups onto humans is not justifiable. For example, an imprint redtail being flown in a crowded island such as Britain, is a definite safety risk, especially to children. Catch-all solutions are not appropriate; we must look at individual situations.

12. THE TECHNICALITIES OF STERILISATION

Over the past three years we at the National Avian Research Center in Abu Dhabi, and the Middle East Falcon Research Group, have been assessing and discussing whether sterilisation for falcons destined for Arab falconry is both necessary and feasible. The balance at present is that it is not necessary (see discussion 7.5 and 8.3).

Is it feasible? The vets (see items below and Steve Sherrod’s paper) consider that, in specialised hands, the operation itself is feasible. But from the practical point of view, how could it be achieved? If one insisted that birds had to be sterilised before they left the breeder, could this be done consistently and safely before the birds were 45 days old and scheduled to go into the hack box? If so, what effect would this have on their feather development at such a critical stage, and on their mental development? Would they all have to be travelled to a clinic for this to be done? Is this in the bird’s welfare interest at this age and would it be legal? What happens about the birds which die or have complications and what follow up would there be to establish that each bird, subsequently, was indeed sterile? From the buyer’s point of view, who would want to buy, let alone pay more for such a bird, when non-sterilised birds are available from other countries which anyway have lower labour and production costs? Careful consideration of these questions, particularly with reference to the large Middle East market, concludes that sterilisation is impractical; Arab buyers will set up their own programmes and not sterilise the birds produced. Western falconers will have penalised themselves for little conservation benefit. However, details of techniques are included here for completeness.

13 THE LAW IN RELATION TO HYBRIDS AND EXOTICS

13.1 The law in relation to hybrids and exotics, as it applies to the individual falconer, is many-tiered. We have international agreements, such as CITES (Convention on International Trade in Endangered Species), which covers participating countries which are signatories to the Convention. CITES is not enforced evenly across the countries and in many third world countries it will never have the level of control and enforcement applied by Management Authorities in Western countries. CITES is primarily aimed at trade and movement, but also tries to define terms such as ‘captive bred’ and ‘commercial purposes’ which tend to set precedents of definition for national legislation.

Countries, or groups of countries such as the European Union, also adhere to other agreements such as the ‘Bern Convention.’ These are listed below.

13.1.2 INTERNATIONAL LEGISLATION AND GUIDANCE CONCERNING RELEASES OF NON-NATIVE SPECIES

13.1.3 The Bern Convention

A4.1. The Convention on the Conservation of European Wildlife and Natural Habitats (the "Bern Convention"), adopted by the Council of Europe on 19 September 1979, aims to conserve wild flora and fauna and their natural habitats, especially those species and habitats whose conservation requires the co-operation of several States, and to promote such cooperation. Particular emphasis is given to endangered and vulnerable species, including endangered and vulnerable migratory species. Article 11(2) of the Convention requires that:

"Each Contracting Party undertakes:

a) to encourage the reintroduction of native species of wild flora and fauna when this would contribute to the conservation of an endangered species, provided that a study is first made in the light of experiences of other Contracting Parties to establish that such reintroduction would be effective and acceptable;

b) to strictly control the introduction of nonnative species." (note: this applies to ALL exotic forms, including hybrids)

The Bern Convention forms the basis for the Council Directive 92/43/EEC, the "Habitats Directive" (see A4.3. below).

A4.2. Further to this Convention, the Committee of Ministers to Member States have made Recommendations concerning the introduction of non-native species[5] and on the reintroduction of wildlife species[6] . These recommendations state that the introduction of non-native species into the natural environment be prohibited where adverse effects on the ecosystem may occur. However, certain exceptions to prohibitions may be authorised on the condition that the possible consequences are assessed beforehand.

Where reintroductions are considered, they should only be undertaken after carrying out research and implemented under scientific supervision. It is recommended that interested parties be informed of such reintroductions. Also, collecting stock for reintroductions should be prohibited from populations which would be threatened as a result.

13.1.4. Council Directive 92/43/EEC on the Conservation of Natural Habitats and of Wild Fauna and Flora

A4.3. The "Habitats Directive", adopted by Member States on 21 May 1992, also aims to conserve wild flora and fauna and their natural habitats and is based on the Bern Convention. Article 22(b) states that, in implementing the provisions of this Directive, Member States shall:

"ensure that the deliberate introduction into the wild of any species which is not native to their territory is regulated so as not to prejudice natural habitats within their natural range or the wild native fauna and flora and, if they consider it necessary, prohibit such introduction. The results of the assessment undertaken shall be forwarded to the committee for information."

13.1.5. Council Directive 79/409/EEC on the Conservation of Wild Birds

A4.4. The "Birds Directive"aims to protect wild bird species and their habitats. Article 11 states that:

"Member States shall see that any introduction of species of bird which does not occur naturally in the wild state in the European territory of the Member States does not prejudice the local flora and fauna. In this connection they shall consult the (European) Commission".

13.1.6. Convention on Biological Diversity

A4.5. The Convention on Biological Diversity was opened for signature on 5 June 1992 at the United Nations Conference on Environment and Development at Rio de Janeiro, after adoption on 22 May 1992 at the Nairobi Conference for the Adoption of the Agreed Text of the Convention on Biological Diversity. The Convention entered into force on 29 December 1993. The objectives of this Convention are the conservation of biological diversity, the sustainable use of its components and the fair and equitable sharing of the benefits arising out of the utilization of genetic resources.

The Convention specifically addresses the introduction of non-native species. Article 8(h) states that:

"Each Contracting Party shall, as far as possible and as appropriate:

(h) prevent the introduction of, control or eradicate those alien species which threaten ecosystems, habitats or species.

13.1.7. IUCN Position Statement on Translocation of Living Organisms: Introduction, Re-introductions and Re-stocking.

A4.7. The World Conservation Union (IUCN) position, published in 1987, is that introductions should only occur if there are clear and well defined benefits and if no suitable native species are available. It sets out general principles for determining the desirability of intentional introductions in natural, seminatural and human-made habitats, discouraging accidental introductions, undertaking eradication measures and administering introductions in national and trans-boundary contexts.

See also Shine, C., N. Williams and L. Gundling. 2000. A Guide to Designing Legal and Institutional Frameworks on Alien Invasive Species. IUCN Environmental Policy and Law Paper No 40.

13.1.8. FAO Code of Conduct for the Import and Release of Exotic Biological Control Agents

A4.8. The United Nations Food and Agriculture Organisation's Code of Conduct, published in February 1996, sets out the responsibilities of Government authorities, exporters and importers of biological control agents capable of self-replication (parasitoids, predators, parasites, phytophagous arthropods and pathogens) used in research and for environmental release, and describes three responsibility phases; pre-export, pre-import and post-import.

13.1.9. ICES Code of Practice on the Introductions and Transfers of Marine Organisms 1994

A4.9. This Code of Practice, adopted by the International Council for Exploration of the Sea in September 1994, recommends practices and procedures to diminish risks of detrimental effects from the introduction and transfer of marine organisms. It provides recommendations for new intentional introductions and suggests that member countries submit proposals to ICES for an opinion on a proposed introduction.

13.2 NATIONAL LEGISLATION AND GUIDANCE CONCERNING RELEASES OF NON-NATIVE SPECIES.

We then have group legislation such as the European Community Birds Directive. This is applied in each European country as national legislation, therefore differing in each place, and in Britain at least (in the form of the Wildlife and Countryside Act 1981 and the Import/Export Act) tends to be stricter. In the US there is a Federal, and a State level of legislation. Below the national legislation, we have voluntary codes of conduct such as those applied by the IAF and by falcony organisations such as NAFA, the Hawk Board, DFO, BFC, etc.

Note: In the USA the legislaion is USFWS: 50CFR21:30. The UK legislation is as follows:

The UK is governed by Section 14 of the Wildlife and Countryside Act 1981 as a legal framework for the application of the ‘Bern Convention’:

With respect to the release of animal species, Section 14(1) of the Act states that:

'Subject to the provisions of this Part, if any person releases or allows to escape into the wild any animal which -

(a) is of a kind which is not ordinarily resident in and is not a regular visitor to Great Britain in a wild state;

or

(b) is included in Part I of Schedule 9, he shall be guilty of an offence.

With respect to the release of non-native plant species Section 14(2) of the Act states:

"Subject to the provisions of this Part, if any person plants or otherwise causes to grow in the wild any plant which is included in Part II of Schedule 9, he shall be guilty of an offence."

Thus, sections 14(1) and (2) of the Act respectively prohibit the release of all non-native animal species (see paragraphs 2.2 - 2.5 below) and certain plant species (see paragraphs 2.6 - 2.7 below). There are certain groups of organisms to which the Act does not apply which are listed in paragraph 2.8 below.

Non~native animals

2.2. Section 14(1) of the Act prohibits the release of any non-native animal into the environment, no matter what its purpose or circumstances. In addition to direct releases to the open environment, including natural and semi-natural habitats, it is considered that "releases or allows to escape into the wild" includes semi-confined situations such as gardens and glasshouses. This is because animals will escape into the wider environment via available routes such as vents in commercial glasshouses, or via removal of soil or plant matter in which they may shelter or feed. Similarly, the keeping of non-native aquatic animals in cages or pens floating in lakes, sea lochs and coastal regions is also considered as a release, because non-native freshwater or marine fish and invertebrates are likely to escape in these situations.

2.3. Section 14 applies to "any animal". It is considered that this is any species that is currently accepted by scientists to fulfil the criteria of the Kingdom Animalia. This includes all nematodes (including microscopic species), mites, insects and all other invertebrates, in addition to all vertebrates (fish, amphibians, reptiles, birds and mammals). The "kind" of animal may be specified down to the sub-species level, and includes hybrids between native and non-native species. It is also important to note that all life-cycle stages of the non-native animal are controlled. Thus the Act applies, where appropriate, to eggs, semen, embryos, larvae, pupae and adult stages.

Query: 14 (1) (a).... is of a kind which is not ordinarily resident in and is not a regular visitor to Great Britain in a wild state. ‘Kind’ refers not to species, but to an individual which could be distinguishable phenotypically, but be taxonomically identical. For example, the Racing Pigeon, Town or Feral Pigeon and the various fancy pigeons, are all the same taxonomically as the native Rock Dove, i.e. they are all Columba livia. But obviously they are phenotypically different because they are of different ‘kinds’.

However, most Schedules to the Act maintain that common names shall not apply, only scientific names: in other words, phenotypic differences will be ignored in favour of taxonomic differences. So, what is a ‘kind’? As far as human introductions are concerned, the criterion for entry is at species level - Homo sapiens - and does not discriminate against non-indigenous races or phenotypes: indeed it is illegal to do so!

2.4. In section 14, a non-native animal is one that is "not ordinarily resident in and is not a regular visitor to Great Britain in a wild state". This includes all species of animals which, according to scientific records, do not naturally occur in Great Britain. In order to be resident, it is considered that a species must be breeding in the wild and producing young which reach maturity without the deliberate assistance of man, ie: under natural environmental conditions. Section 14 does not extend to non-native animals that regularly visit Great Britain in a wild state, for example migratory birds and some species of insects which, as an established behavioural trait, seasonally migrate either for the summer or winter months to utilize environmental resources for feeding and reproduction.

Query: And rather than or, indicates that the animal must be of a kind which is neither a resident nor a visitor in a wild state. For example, the domestic cat is not ordinarily resident in a wild state, although feral cats do breed in the UK, nor are cats regular visitors to Britain because they cannot swim the Channel. It is therefore illegal to release domestic cats on both counts. Gyrfalcons on the other hand, although they visit Britain, are not ordinarily resident and therefore only fulfil one of the two mandatory requirements. It is therefore legal to release them.

Regular refers to the even spacing of the visits and has been misused here. Regular could mean once every ten minutes or once a century. What was intended was some measure of frequency.

In a wild state refers here not to a place, but to a condition of the ‘kind’ of animal which already resides or visits Britain (ie. not the individual being released). It may mean that the ‘kind’ is independent of man in terms of food supply, nest sites or breeding; it may mean that the species is ‘mentally wild’, in the sense of unapproachable, or genetically wild type rather than a feral domestic type. It cannot mean a place - because the animal could not visit and take the wild place with it.

2.5. Section 14(1)(b) prohibits releases of animal species into the wild which are specifically listed in Part I of Schedule 9 of the Act (see Annex 1), which is referred to in section 14(1)(b). In practice, most of the animals listed are those known to be established in the wild in Great Britain and causing damage to the environment.

Non-native plant species

2.6. Section 14(2) of the Act only prohibits the release of the non-native plants specifically listed in Part 11 of Schedule 9 of the Act. These are plant and algal species which have become established in habitats in Great Britain and have been specifically listed because of the damage they are causing to the natural environment (see Annex 1).

2.7. The Act states that any person who "plants or otherwise causes to grow in the wild" the listed plant species shall be guilty of an offence. It is considered that this includes the deliberate planting of these species in the wild. It also includes the cultivation of these species in confined situations but allowing the removal or transfer of plant material to where it may become established in the wild. All life cycle stages and parts of these listed species which are capable of dispersal, regeneration growth and reproduction are controlled: including, where appropriate, reproductive and dispersal structures, seeds and live vegetative or root material capable of regenerating, for example rhizomes of Japanese knotweed (Fallopia japonica).

Query: Whereas subsection (1) essentially covers the introduction of new exotic animal species, subsection (2) does not make the same provision for exotic plant species. Provided one does not introduce any of the four plants in Schedule 9, two of which are seaweeds, there are no controls on exotic plants under this Act.

Species not covered by section 14 of the Act

2.8. Species in the following groups of organisms are not controlled by section 14 of the Act:

• Monera (including viruses and prokaryotic microorganisms, including all bacteria);
• Protista (including all protozoa and eukaryotic algae, including seaweeds), except those listed in Part II of Schedule 9 (listed in Annex 1);
• Fungi (including lichens);
• Plants (including mosses, liverworts, horsetails, ferns, gymnosperms and angiosperms) except those listed in Part II of Schedule 9 (listed in Annex 1).

Re-introductions of animal species endangered or extinct in Great Britain

2.9. Where native animal species are declining or have become extinct from areas of Great Britain, there may be attempts to reintroduce populations using non-native stock from outside Great Britain. These releases are also prohibited by section 14 of the Act, if the non-native stock consists of a sub-species which is not native to Great Britain. It is considered that the non-native sub-species of the animal from another country may be genotypically and phenotypically significantly different from the original native populations, and thus may behave differently in the environment after release. Re-introductions should be planned within the context of UK Biodiversity Action Plans[7], and applications for licences will be considered with existing plans and policies in mind. Further information regarding Species Action Programmes may be obtained from English Nature, Scottish Natural Heritage, the Countryside Council for Wales or the joint Nature Conservation Committee (addresses at Annex 6).

Population studies of animal species listed in Part I of Schedule 9 of the Act

2.10. Where field studies are being carried out with populations of species listed in Part I of Schedule 9 of the Act which are established in Great Britain, it may be necessary to apply for a licence. This would apply where, in the course of a study, individuals of the list species are captured, tagged or otherwise labelled for identification purposes, and re-released. In the event of re-release, an offence shall have been committed unless a licence had been granted to the person or persons responsible for the study.

Penalties for an offence under section 14 of the Act

2.11. Section 21 (4) of the Act states:

"A person guilty of an offence under section 14 shall be liable-

(a) on summary conviction, to a fine not exceeding the statutory maximum;

(b) on conviction on indictment, to a fine.

With respect to section 21(4)(a), the maximum statutory fine, arising from a summary conviction in either a sheriff or a magistrates court is presently £5000. With respect to section 21(4)(b), the fine arising from a conviction on indictment in the High or Crown court is potentially unlimited.

2.12. If an offence is committed under section 14(1) (for releasing or for allowing the escape of non-native animals without a licence) or section 14(2) (for planting or otherwise causing to grow in the wild the plants listed in Part 11 of Schedule 9 without a licence), under section 14(3) of the Act it shall be a defence to a charge of committing an offence to prove that all reasonable steps were taken and all due diligence exercised to avoid committing an offence.

Summary of Section 2

The Act prohibits the release of all non-native animal species and sub-species into the wild in Great Britain. This includes semi-confined situations where the animals will escape into the wild.

• The Act also prohibits the release of specifically named animal species which are established in Great Britain.
• The Act also prohibits the release of specifically named plant species which are established in Great Britain.
• The penalty on a conviction for releasing non-native animals or the listed plants without a licence may be an unlimited fine.

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CONCLUSION:

STEVE SHERROD (US Co-chair)

1. This document evidences the miniscule threat that exists from introduced falcons to native falcon inhabitants. Nevertheless, hybrids comprised of at least one native species do impose a certain risk factor. Regardless of that fact, I believe that the requirements in the current USFWS regulations (50CFR21.30) regarding breeding and raising hybrid falcons are totally adequate to prevent any significant problems with these birds in wild populations as long as those regulations are followed as written.

2. In addition, working radio telemetry should be required to be used on any hybrid falcons flown or hacked, and;

3. a name tag with phone number should also be required on these birds always when flying freely. NAFA Members should be willing to help state authorities trap (and if not possible, remove by any means necessary, ultimately including shooting) any hybrid or exotic falcon which takes up residence at eyries. The methods of sterilization which are now being used and further developed will have to be learned by more avian veterinarians, and they will have to be available to the falconry/breeder community at reasonable costs. The owner of any such hybrid falcon will have to be held responsible for the sexual orientation (imprinted/wild raised) and reproductive state (sterilized/non-sterilized) of his/her own bird. Ultimately I believe most falconers will deal with this matter responsibly if the operation is available to them. Some, however, might not.

There has been discussion that the owners of all hybrids might be required to provide documentation regarding how their individual hybrid was raised, and to have available for inspection a certificate of sterilization signed by a licensed veterinarian for any non-imprints. This would not be acceptable because of the varying shades of gray or loopholes which exist, and therefore would be very difficult to enforce. When matters are not black and white, and when they become unenforceable, law enforcement officials react by eliminiating the problem in its entirety (i.e. hybrids).

Although the problem is in reality quite small, concern is real and not totally unfounded; the perception of a much greater problem than probably actually exists as well as some individuals' failure to abide by the reasonable regulations we already have, could well result in restrictions which eliminate this bird from American falconry.

SUMMARY: WE MUST BE WILLING TO FOLLOW THE RULES WE ALREADY HAVE.

---

CONCLUSION: NICK FOX (UK Co-chair)

I agree with Steve Sherrod’s three points, with the addition of the following:

1. The ‘wild’ hacking of exotic or hybrid species which have not been sterilised or which are not closely monitored by radio-telemetry, and which result in individuals becoming ‘hacked back’ to the wild should be discouraged. This has been the root of the problem in Germany.

2. Falconers should be strongly encouraged not to shoot themselves in the foot over this issue. Look around you at how totally dependent we are on non-native organisms, in our food, our gardens, in our landscape. Are we prepared to get rid of our dogs and keep wolves instead?

14 ACKNOWLEDGEMENTS

The chairman would like to personally thank all the members of the Hybrid Discussion Group for their contributions; Helen Macdonald, who created and moderated the list and provided valuable editorial assistance with this document; and finally, we are indebted to all those who have previously tackled this issue for NAFA and other organisations. We have not been able to include all of their work due to the fact that much of the scientific data and technical information (eg on sterilisation) have been superseded. Their work has been crucial to the tenets and scope of this document.

Special thanks are due to Mr Mohamed Al Bowardi, Managing Director of the National Avian Research Center, part of the Environmental Research and Wildlife Development Agency, Abu Dhabi, for his continuing initiative in tackling conservation issues related to Arab falconry, including support in the preparation of this document.

APPENDIX 1

PARTICIPANTS

Dr Steve K. Sherrod
Committee Co-Chair
Director, George M. Sutton Avian Research Center
PO Box 773, Bartlesville, OK 74005 USA
918 333 8346 (H), 918 336 7778 (W)
Email: BLACKJESS1@aol.com (H)

Dr Nick C. Fox
Committee Co-Chair
Director of Falcon Management and Research
The National Avian Research Center, Abu Dhabi
Falcon Facility, PO Box 19,
Carmarthen, SA33 5YL, Wales UK
Tel/Fax: 44 1267 233864
Email: office@falcons.co.uk

Dr Walter Bednarek
Deutscher Falkenorden
Haselhof 4. 4428, Rosendahl, GERMANY
4925 477129 (H)

Mr Robert B. Berry
President, North American Raptor Breeders’ Association
100 Rapid Creek Road, Sheridan, WY 82801 USA
307 672 7716 (H), 610 688 2535 (W)

Mr Frank Bond
North American Falconers’ Association
RT9 Box 68FB, Santa Fe, NM 87505 USA
505 984 2061 (H), 505 988 4476 (W)

Mr Christian de Coune
President, International Association for Falconry (IAF)
Le Cochetay, 4140 Gomze-Andoumont, BELGIUM
041 6873 7369

Mr Anthony Crosswell
Editor of the British Falconers’ Club Journal
Sneath Farm, High Green, Great Moulton, Norwich, Norfolk NR15 2HU, UK
01379 677296 (H)
Email: GYRCROSS@aol.com

Dr Robert Kenward
Institute of Terrestrial Ecology
Furzebrook Road, Wareham, Dorset, ENGLAND
Email: reke@wpo.nerc.ac.uk

Dr J. Timothy Kimmel
North American Falconers’ Association President
Route 1, Box 82A, Ellinwood, Kansas 67526 USA
316 562 3509 (H) 316 792 9335 (W)
Email: kimmelt@cougar.barton.cc.ks.us

Dr Gordon Mellor
School of Humanities, Sport and Education
De Montfort University
Landsdowne Road, Bedford MK40 2B2 UK
01234 351966 (W) 01234 713383 (H)
Email: gtmellor@zetnet.co.uk

Mr Patrick Morel
Honorary Secretary IAF
Rue de Longueville 13, B-1315 Incourt, BELGIUM
3210 889 030 (H) 3210 412 664 (W)
fax: 3210 881 177

Mr Jose-Manuel Rodriguez-Villa y Matons
Darro, 18, 28002 Madrid, SPAIN
156 42574 (H) 1766 5309 (office fax)
Email: mazdaesp@lix.intercom.es

Professor Clayton White
Department of Zoology
Brigham Young University
574 Widtsoe Building, PO Box 25255, Provo, Utah 84602-5255 USA
801 378 2006 (H)
Email: WHITEM@acd1.byu.edu

APPENDIX 2

EXTRACTS FROM ‘THE REGULATION AND CONTROL OF THE RELEASE OF NON-NATIVE ANIMALS AND PLANTS INTO THE WILD IN GREAT BRITAIN’, DEPARTMENT OF THE ENVIRONMENT 1997

1. Introduction

1.1. Numerous non-native animal and plant species from other countries have been introduced into Great Britain for a number of reasons in historical times. Some introductions may have been accidental, others to provide different sources of food, sport or for ornamental purposes. Many non-native species have become well established as part of Great Britain's fauna and flora, interacting with native species. Their success depends on their ability to adapt to the environmental conditions experienced in Great Britain, and on their biological interaction with native species.

1.2. In some cases non-native species can adapt and invade natural communities. Often, this is because they have not evolved over a long period of time with the native species with which they interact, so environmental pressures to limit their spread, such as pathogens, predators or defence adaptations are absent. Some naturalised introductions may be relatively benign, while others may be damaging to the environment, including native flora and fauna, or to human interests such as agriculture and forestry. Changes in land use may exacerbate spread, because environmental conditions and the species balance are altered, giving the non-native species a competitive advantage. There are many examples worldwide of damage to the physical environment or native fauna and flora occurring as a result of non-native species introductions. In Great Britain examples include the grey squirrel (Sciurus carolinensis), the New Zealand flatworm (Artioposthia triangulates) and Japanese knotweed (Fallopia japonica). It is therefore important to control and regulate releases of non-native species to ensure that further damage to the environment does not occur.

International controls on releases of non-native species

1.3. The introduction of non-native species is controlled by international and European legislation, particularly Article 11 (2) (b) of the Convention on the Conservation of European Wildlife and Natural Habitats (the "Bern Convention"), Article 22(b) of Council Directive 92/43/EEC on the Conservation of Natural Habitats and of Wild Fauna and Flora, Article II of Council Directive 79/409/EEC on the Conservation of Wild Birds, and Article 8(h) of the Convention on Biological Diversity. Codes of Practice have also been drawn up by international organisations, for example the Food and Agriculture Organisation (FAO) International Standards for Phytosanitary Measures "Code of Conduct for the Import and Release of Biological Control Agents", the International Council for Exploration of the Sea (ICES) "1994 Code of Practice on the Introductions and Transfers of Marine Organisms", and the International Union for Conservation of Nature (IUCN) "Translocation of Living Organisms" (1987). Further details on this legislation and guidance are provided in Annex 4 to this guidance note.

Controls of releases of non-native species in Great Britain

1.4. The release into the wild of animal species which are not native to Great Britain, whatever their proposed use, is prohibited by section 14 of the Wildlife and Countryside Act 1981 ("the Act"). The release of certain animals and plants which are already established in the wild in Great Britain is also prohibited by section 14 of the Act. Section 16 of the Act gives the Secretary of State and the Minister of Agriculture, Fisheries and Food powers to grant licences for releases so that section 14 does not apply. Releases of non-native animals may therefore be licensed under the Act for specific purposes, for example to authorize their use for the control of pests on commercial crops.

APPENDIX 3

NORTH AMERICAN FALCONERS ASSOCIATION POLICY ON RAPTOR HYBRIDIZATION

A. N.A.F.A. encourages the captive-breeding of the raptors used in falconry under the terms of those laws/regulations currently (Nov. '77) in effect. N.A.F.A. is opposed to any captive breeding attempts undertaken without appropriate permits.

B. Although encouraging captive-breeding of "pure strains" of raptors (i.e. subspecies with like subspecies), N.A.F.A. approves of intraspecific hybridization when accomplished in accordance with Item A, above.

C. N.A..F.A. similarly approves of interspecific hybridization when conducted in accordance with appropriate permits. Recognizing that much interspecific hybridization to date has occurred only because of a lack of "matched" pairs of birds or a lack of semen to match the species/subspecies of laying females, N.A.F.A. encourages those of its members participating in captive-breeding to exchange birds or semen (within appropriate legal restrictions) to facilitate "pure" breeding and to refrain from such cooperation with anyone attempting to breed without appropriate permits.

D. N.A.F.A. concurs in principle with the present federal restrictions placed on interspecific hybridization (i.e. that birds so produced be surgically sterilized or totally imprinted on humans), but supports such restrictions ONLY for birds to be used in falconry. In view of the fact that surgical sterilization is a procedure of considerable technical difficulty', N.A.F.A. highly recommends that interspecific hybrids to be used in falconry be imprinted.

E. Although the possibility of hybrid "contamination" of wild raptor populations is virtually infinitesimal, N.A.F.A. strongly urges that falconers flying hybrids utilize telemetry to reduce thus possibility of their loss.

F. The N.A.F.A. urges that individuals engaged in any captive raptor breeding keep scrupulously accurate records and participate fully and without reservation in the North American Peregrine Foundation Raptor Registry and Stud Book so as to develop maximum knowledge from all captive raptor breeding.

‘NAFA calls to the attention of all captive breeders the problem - and potential subsequent over-reaction and likely over-restriction - that could come about as a result of ‘flaunting’ hybrids of any type or the subject of captive hybridization before unknowledgeable non-falconers and other raptor breeders. The NAFA strongly recommends the utmost discretion be used in any public statements etc., concerning the subject.’

Since 1977, substantial data have become available on the status of most raptor species suitable for falconry, and depressed raptor populations have generally recovered in North America and Europe. Most of the recommendations of the WOS Conservation Committee have been achieved in the United States and Canada, and the contributions of falconers to raptor management and conservation education have been widely recognized. Yet regulation of falconry still causes controversy in some countries.

The purpose of the Falconry Position Statement by the Raptor Research Foundation, Inc., is to provide current and additional expert opinion based on available biological data on issues relating to the regulation and practice of falconry. This statement neither affirms nor disaffirms the philosophical question of the legitimacy of the sport of falconry.

DEFINITION OF ISSUES

Harvest of wild populations -- The removal of young birds from wild populations reduces productivity (directly, and perhaps indirectly through disturbance during the nesting season). However, raptors are a renewable resource, and thus the game management principle of ‘sustainable yield’ may be appropriately applied to harvest of individuals from healthy populations.

Captive Propagation -- Captive propagation of raptors has increased dramatically in recent years, and the release of captive-bred progeny has been valuable for reestablishing some endangered species. Captive breeding provides birds for falconry, but may also be used to conceal illegally acquired birds unless parentage can be proven.

Hybridization and Introductions -- The production of hybrids, especially among large falcons, has raised questions concerning the release of such birds to the wild. Genetic theory predicts that, at normal population levels, hybrids between sympatric or parapatric species would be eliminated by natural selection. Similarly, non-native species from within the same super-continent (Americas, Eurasia) are unlikely to establish themselves in the wild as introduced aliens. However, trains from hybrids between allopatric species might establish in native stocks, and species from other super-continents might become accidentally introduced if used in large numbers for falconry.

Identification of Individual Birds and Parentage -- To effectively enforce falconry regulations, individual birds must be reliably identified. Leg bands which cannot be refastened after removal would be a convenient method, but bands currently used are not entirely reliable. Alternatives include biochemical parentage tests, which should soon become available for raptors, and foot scute patterns which are expected to provide unique ‘fingrprints’ for individual identification.

Regulation and Enforcement of Falconry. --Controls are desireable; however, the intensity of regulations and their enforcement should be consonant with the risk to raptor populations.

POSITION

1. The position of the Raptor Research Foundation, Inc., with regard to the above stated issues relating to Falconry is:
2. North American raptors used in falconry have stable or increasing populations throughout most or all of their range. This is also generally true of European countries where falconry is practiced.
3. Evidence indicates that large and stable or increasing raptor populations can sustain an annual harvest of at least 10% of nestlings.
4. Any harvest of raptors from small and unstable or declining populations should be evaluated, in each instance, on a biological (e.g. population and productivity data) basis.
5. The annual harvest of wild raptors by falconers in the United States is well below 5% for any species and below 1% for most species. Percentages are not adjusted for return to wild stocks of released and escaped birds.
6. Final development of biochemical parentage tests and the use of foot scute patterns for individual identification should be encouraged as tools for regulation and enforcement.
7. Escape of sympatric and parapatric species or their hybrids is unlikely to pose any significant threat to wild populations. However we recommend that hybrids between allopatric species should not be bred for falconry, and that other hybrids or species at risk of accidental introduction between super-continents should be imprinted on humans before being used in falconry.
8. Licensing individual falconers on merit is effective for regulating falconry, espacially when combined with individual markers for raptors of special management concern (e.g., endangered species.) There is little conservation justification for the administrative costs of marking common raptors individually, and future consideration should be given to modification of this practice.
9. Many resources now being directed towards the control of falconry in the United States and elsewhere could be redirected to raptor population monitoring, habitat conservation, education and preventing the killing of wild raptors.
10. Government agencies should be more responsive to the changing status of species, both by imposing protection when necessary and by removing restrictions on use when biological data indicate such is warranted.
11. International standards for the practice and regulation of falconry are encouraged.

1997

APPENDIX 5

LETTER FROM ROBERT A. WITZEMAN, M.D. of the MARICOPA AUDUBON SOCIETY TO HAROLD OLSON, NEW MEXICO DEPT. OF GAME AND FISH.

September 6 1982

Dear Mr Olson,

The New Mexico Department of Game and Fish is to be congratulated for recognizing the importance in limiting potential introductions of exptic wildlife which may constitute a threat to native populations and their ecosystems as well.

Historically unthinking and unplanned actions by man have led to the destruction of native wildlife populations or their habitats or both. A few examples should be mentioned:

1. European starlings here in the Southwest appear to have competed with, and crowded out many native cavity-nesting species such as Elf Owls, Gila Woodpeckers, Gilded Flickers and Purple Martins. This may have major untoward implications for the entire Saguaro-Palo Verde vegetative and wildlife community.

2. Rainbow trout in Arizona have interbred and genetically swamped Arizona’s native Apache trout, requiring extensive rescue and restoration efforts. Brown trout also introduced in the area have competed for the food supply and preyed upon the Apache trout.

3. Rio Grande Cutthroat Trout likewise have been genetically diluted by Rainbow trout and exotic subspecies of Cutthroat trout in New Mexico.

4. The Red Wolf has reached near extinction for many reasons, and the few remaining have been genetically destroyed through interbreeding with the Coyote and dog in Texas and Louisiana.

5. The re-stocking of depleted Northeastern US Bobwhite populations with genetically distinct Texas and Mexican Bobwhite populations has produced an individual which may biologists believe is less capable of wintering-over successfully in the northern ecosystem (Our Wildlife Legacy, Durward Allen, 1954.)

6. Gambusia introduced into the Southwest have competed for food and also preyed directly upon the native, endangered Gila Topminnow populations. Reintroduction of Topminnows has succeeded primarily where Gambusia have been entirely removed-- at great comittments in time and cost.

7. The Masked Bobwhite subspecies of Bobwhite Quail was reintroduced into Arizona with great care to preserve the genetic purity of the introduced population. Guardian (Texas subspecies) surrogate (male) Bobwhite parents released with the Masked population were carefully sterilized to prevent genetic mixing of the highly-esteemed indigenous Masked Bobwhite population (formerly found in Arizona until past overgrazing excesses eliminated it).

Many people do not realize the small size of our native southwestern raptor populations. There are but a few dozen pairs of Peregrine Falcons - the native breeding population - in New Mexico and Arizona. The flying of non-native Peregrines here in the Southwest, which have origins, whether from far-flung parts of the world or other adjacent North American populations, tempts disaster. The various world-wide subspecies or populations of Peregrines are not just subtle taxonomic differences. They may vary greatly from each other in size, appearance, behavior, blood chemistry or migration patterns. Each population represents hundreds of years of evolution - to enable that race to thrive best in its own ecosystem.

The ability of exotic falcon populations to provide the genetic makeup to cope with our southwestern ecosystem is unknown and when a key species such as the Peregrine Falcon is at stake, how can such chances be taken? The Department is to be congratulated for the September 25 regulations which will insure the integrity and survival of the Peregrine and other native raptor populations from potential jeopardy by exotics.

It should also be noted that the use of Peregrines from the 'eastern" Cornell mixed stock should not be flown and represent an unwarranted risk. They are composed of a highly crossbred mix containing the genes of several different Peregrine subspecies-- Including many arctic tundra birds, birds from heavily forested wet Pacific coastal areas and even European stock. This "polyglot" population has never had the benefit of centuries of adaptation to the rigors of survival In the New Mexican ecosystem. The native New Mexican subspecies, which has had the benefit of years of the natural selection process has already shown it is capable of coping with the Southwest environment.

Peale's Peregrines from the extremely humid Pacific Northwest coastal areas of Canada have eggshells of increased porosity to accommodate the transpiration of water concurrent with embryonic metabolic processes. Such genes would suggest disaster for our southwestern Peregrine population.

Because Arizona has a Peregrine Population which interacts closely with and shares the genes of those birds in New Mexico, we are especially gratified that the new regulations address this concern.

New Mexico and Arizona both have the good fortune of possessing other unusual and exciting raptor populations which are relatively small in numbers and could be adversely impacted by exotic raptor introductions:

1. The Harris' Hawk subspecies of South America is different from the Harris' Hawk population found in New Mexico or the one found in Arizona. The latter two populations are separated from each other by the Sierra Madre and Rocky Mountain continental divide. The inadvertent or unthinking release of a non-native race into any of these three populations would be a disservice to the centuries of natural selection which made each population fit to cope with its respective ecosystem. Harris’ Hawks obtained in Arizona or Western Mexico should not be flown by falconers in New Mexico nor should New Mexico-Texas birds be flown in Arizona.

2. Arizona and New Mexico may both point with pride to their Apache Goshawk population. It is to be commended that the proposed regulations prevent this southern population from being placed in jeopardy by (smaller) northern Goshawk populations of the US and Canada or by any of the half dozen other exotic populations of Goshawks which are found throughout the northern hemisphere in the old and new world.

The use of hybrid species, as permitted in the proposed regulations, is unwise even with transmitters. Such equipment is far from foolproof and costly. Species which some biologists have believed to be sterile-as-mules have not always turned out that way. The supposedly sterile Grass-Carp (White Amur-Bigmouth Carp cross) was an unpleasant surprise to a great many biologists.

A host of ‘hybrids’ of widely varying lineage have been used for falconry in the US involving Gyrfalcons, Peregrines, Prairie falcons, Lanner, Saker and many other allied and peregrine-like taxa (forms). The taxonomy and genetic isolation of many of these forms is unknown to biologists today. One glance at wolf, dog and coyote interbreeding or duck fancier breeding efforts leads one to a healthy respect for the destructive potential which such so-called hybrids could hold for our native southwestern raptor populations.

In conclusion, the Department is to be congratulated on its drafting of falconry regulations which reduce the potential for introduction of exotic wildlife. We have also commented upon the telemetrized use of hybrid species and why it is unwarranted. Thank you for this opportunity to present this commentary.

Sincerely,

Robert A. Witzeman, M.D.

Editor

APPENDIX 6

D) extract from: RECOMMENDATIONS OF THE COLORADO WILDLIFE FEDERATION TASK FORCE ON RAPTOR LAW AND REGULATION

Submitted to the Colorado Wildlife Commission

4 January 1984

The Colorado Wildlife Federation established the Task Force on Raptor Law and Regulation in September 1983 based on our concern that there had not been full and complete discussion on several raptor regulation issues and the fact that new information had become available on key issues. The Task Force members are listed below. The Task Force identified key issues, met formally four times, and has made the following recommendations. These recommendations were reviewed and approved by the Board of Directors of the Colorado Wildlife Federation on December 10, 1983. Colorado Wildlife Federation appreciates the responsiveness of the Commission to the work of our Task Force. Colorado has historically been a national leader in the development of raptor management. We believe these recommendations provide Colorado and the Colorado Wildlife Commission with the opportunity to continue this leadership role in a very positive and constructive fashion. References made in the following recommendation discussion are to the Chapter 6 Commission regulations approved in August, 1983.

1. Hybridization. The Task Force is convinced that truly negative biological effects to wild populations resulting from the occasional loss of individual intra-specific (sub-species) crosses are remote. To discourage sub-species crossing by requiring such crosses to be surgically sterilized or imprinted is over-regulation which could result in net deleterious rather than beneficial effects. Sterilization or imprinting should be required of inter-specific (species) hybrids only. To further guard against impacts on wild populations, all crosses of, and non-native, Falconidae should be flown free only with identification and telemetry.

Recommendations:

A. Delete "sub-species" and "cross-breeding" from definition of "hybridization" (p. 19, #600 e.);

B. Allow hybridization (inter-specific) in accordance with federal regulations, i.e. require surgical sterilization or imprinting of produced hybrids, and prohibit hybridizing threatened or endangered species (p. 14, #625 a.2.); and

C. Require any sub-species cross or hybrid of the Family Falconidae, and any species or sub-species of the Family Falconidae not native to Colorado, to have an owner identification tag and adequate telemetry equipment when flown free (p. 10, #611 C., under Death, Escape, or Release of a Raptor, and p. 14, #625 a.z., Hybridization).

2. Sale. The sale of captive bred raptors under federal regulations became legal on August 8, 1983. The U.S. Fish and Wildlife Service, in making sales legal, argues that breeding and selling (especially rare) raptors under the federal guidelines will benefit wild populations and serve to increase their numbers by providing a cost recovery incentive to individuals for successful propagation programs. The Service argues further that sale will alleviate human pressures on wild populations by meeting demand with captive produced birds and maintains that sale will increase the genetic diversity in captive populations by encouraging production and providing an incentive to exchange valuable breeding stock. The Task Force was disappointed to find the federal regulations to be less than clear in many cases and sometimes internally inconsistent.

The Task Force is generally very wary of commercializing Colorado’s raptor resources, particularly the rarer species. The creation of new incentives, namely maximizing profit, for removing young birds or eggs from the wild, incentives which are not related to the bird’s biological welfare, is contrary to the sound husbandry of a very special part of Colorado’s Wildlife heritage. Serious concern was also expressed about the creation of fly-by-night or non-serious propogator intending only to make money. Yet some aspects of the Service's arguments make sense. The Task Force was relieved to discover that the Service's long-heralded seamless bands do exist and seem to have some promise of effectiveness. Reports of extremely high prices for certain raptors, especially in foreign markets, appear, however, to be accurate. Moreover, the Task Force is concerned about the possible development of bird dealers whose only goal is maximizing profit in brokering desirable birds of prey.

APPENDIX 7

THE USE OF EXOTIC PEREGRINES TO REPOPULATE LOST RANGE

By Tom J. Cade, Cornell University, 10 August 1980.

"...I believe that conservation should mean the keeping or putting in the landscape of the greatest possible ecological variety--in the world, in every continent or island, and so far as practicable in every district. And provided the native species have their place, I see no reason why the reconstitution of commities to make them rich and interesting and stable should not include a careful selection of exotic forms ...” (ELTON, 1958)

After a lapse of 25 years, peregrine falcons are again nesting in the eastern United States. They are not descendants of the "duck hawks" that Archie Hagar, Joe Hickey, Dick Herbert, Walter Spofford, and other falcon watchers once knew, for that population of peregrines was totally extirpated by DDT poisoning. Nor are they nesting on the rocky crags overlooking the Connecticut, Hudson, and Susquehanna rivers as the duck hawks once did. Instead, they are nesting on special towers in coastal salt marshes and on buildings in cities. They are, nonetheless, bona-fide peregrines of the species Falco peregrinus, produced in captivity by parents taken from various geographic populations extraneous to the eastern United States.

From its beginning in 1970, everyone involved in the eastern peregrine recovery effort has clearly understood that restoration of a nesting population of peregrines in the East would depend upon the introduction of falcons drawn from non-indigenous sources. The issues involved were thoroughly discussed at a conference sponsored by the National Audubon Society in 1972 (Clement, 1974). They were further reviewed in great detail by the Eastern Peregrine Falcon Recovery Team in preparing the official recovery plan for the U.S. Fish and Wildlife Service (Bollengier et al., 1979), and the Office of Endangered Species conducted its own internal review of the matter in 1977 of exotic organisms (E. 0. 1987, May 24, 1977).

The Service then issued official policy statement supporting the use of non-indigenous peregrines release in the eastern U. S.., a policy that is still in force today.

Since the propriety of releasing non-indigenous or exotic falcons into the "natural ecosystem" of the eastern states continues to be questioned by sme people, and since the A.O.U. committee on resolutions specifically addressed the subject in a resolution passed last year and widely publicized, I would like to recapitulate, once more, the salient and, I believe, justifiable reasons for the course of action we have undertaken. My reasons are based on the following views about the conservation of biological diversity.

First, the Endangered Species Act rightly focuses on the species as the taxon of fundamental importance for preservation. It is not the "endangered subspecies act" or the "endangered deme act" for sound biological reasons. Species are populations the members of which freely interbreed and exchange genes in nature, or that have the potential for doing so, but that are reproductively isolated from all other populations of similar organism. Reproductive isolation is the key to the definition of species and also to the scope of the need for preservation, because all members of a species, regardless of how widespread or how nmy populations have been described by taxonomists as "subspecies," share a closed, cohesive,.cdadapted gene pool and a conmn epigenetic system of phenotypic development (Mayr,1963; Selander, 1971). To be sure, a wide-ranging species such as the peregrine falconlis likely to consist of a number of "ecotypes" that are to some degree specially adapted to local enviroments, but if the work of Corbin (1977) and others who have been studying "genetic distance" among avian taxa applies to falcons, then probably as mch as 90 to 95 per cent of the total genetic diversity of the peregrine falcon is shared in conmn by all local breeding populations. This estimate of similarity led Corbin (op. cit.) to conlude that "It is likely that coadapted gene pools of birds differ somewhat among geographic areas. However, the genetic identity data suggest that for management purposes the origin of individuals being used to repopulate areas following local extinctions need not be a major concern of the program."

Second, the primary purpose of work under the ESA is to restore species, as far as possible, to their original density and distribution prior to their endangement; or if that goal is not possible, at least to secure a freeliving population in some suitable range. The purpose is not necessarily or only to preserve local genotypes but to mintain the species as a population of ecologically adapted individuals and to increase their numbers in geographic regions where they have declined or disappeared.

Approximately 2.5 million square kilometers of former nesting range for peregrines now lies vacant east of the Rocky Momtains. As far as we know, the original breeding population has been totally extirpated for about 20 years. Any peregrines that become re-established as breeders within this vast range will be in some sense exotic or non-indigenous, whether they come in naturally or by human intervention.

The chances for natural re-establishment on this range are remote. The Great Plains constitute a partial barrier against dispersal frm a sparse, poorly reproducing population in the western states, and northern breeders show no proclivity to settle and nest in mid-latitude habitats over which they migrate. Thus, the situation is totally unlike that in Great Britain, where a resident population reduced to about 44 per cent of its original numbers has shown a dramatic recovery since 1963 (D. Ratcliffe, in press).

There are three ways to go about establishing a new population.

A. We can try to discover the existing peregrine population with the greatest number of genetic similarities to the extinct duck hawk and use that stock for our introductions, in the expectation that such falcons will have the maximum possible fitness for establishment in the East. Based on geographic proximity (since we know nothing about the actual gene pools), peregrines from Labrador mighthe closest ones; based on morphological similarities, those from northern Alberta might be. Unfortunately, all such potential candidates are themselves severely endangered, and the removal of individuals from these populations for use in an eastern recovery program would meet strong resistance.

B. We can look for peregrines that are the closest ecological counterparts of the former eastern birds and try them. None of the existing North American populations really qualifies as well as, say, French or German peregrines do. But they are exotics, and mreover they are also severely endangered.

C. A third possibility put forth by William Drury (1974) and Ian Nisbet (see Clement, 1974) represents a refreshing counter to the objections often raised about "exotics" and "mongrelization" of races. They suggest that breeders of captive peregrines should deliberately mix their stocks to achieve the greatest possible degree of genetic variability in the genotypes and then to release these "hybridized" individuals into the vacant breeding range and let natural selection pick those individuals that are fit for the present environment of the East. Such a procedure recognizes our inability to determine a priori which kinds of peregrines are adapted for survival and reproduction in the current eastern environment and also comprehends the seyere problems of inbreeding that often occur in small populations of animals, captive and wild (Ralls et al. 1979). (It is worth calling attention here to the elaborate ways in which birds have evolved social behaviors that prevent inbreeding; see Koenig and Pitelka, 1979).

Following Drury's lead, our basic working assumption has been that if we can release enough individual peregrines with some degree of fitness for the eastern environment, then natural selection will have a chance to work toward optimum fitness for the eastern environment, so that after several generations a well-adapted population of peregrines will emerge. If the same selective forces that produced the original duck hawks are still operative, then it is reasonable to predict that the new population will converge genetically and phenotypically on the old; if, as seems more likely, new and different selective forces are now associated with the much altered eastern environment, then a somewhat different peregrine will result. The difference will not be noticeable to 99.9 per cent of the people who watch peregrines, and the new population will still belong to the closed gene pool that is represented by individuals of the species Falco peregrinus.

Since 1975 we have released 272 peregrines in the eastern United States. They represent genotypes derived from breeding populations in Spain, Scotland, Chile, the Canadian tundra and taiga, the Alaskan tundra and taiga, the Aleutian Islands, the Queen Charlotte Islands, and California. Many of these released peregrines have not only managed to survive in the eastern states but have also returned as adults to the environs where they were first established. Five pairs have formed, two have successfully produced young of their own this year, and a third has reared fostered young. In addition, one female has paired with a wild mle at an eyrie in southern Quebec, and she produced two young in 1980.

The genetic backgrounds of these successfully established birds are as follows. The female in Baltimore is a "half -breed" between a California male and a Chilean female; she is mated with a male of Nearctic tundra-taiga origins. The breeding pair in Brigantine NWR consists of a female derived from the Queen Charlotte Islands (Peale's falcon) and another half-breed male of California-Chilean extraction. The Manahawkin breeding pair consists of an Alaskan tundra-taiga male and a female of mixed Alaskan tundra-Queen Charlotte Island parentage. The Sedge Island male is Spanish, the female, Nearctic tundra. The new birds in Atlantic City have not yet been individually identified. The female in Quebec is either of tundra origin or a tundra-taiga mixture; she has not been individually identified either.

Despite their diverse genetic backgrounds, these successfully established peregrines have converged remarkably close in their main biological habits to the former duck hawks. First of all, they are not highly migratory-- some are quite sedentary-- despite the fact that many of them derive from populations that are highly migratory in their natural ranges. Our most southerly reports so far are one bird seen in the Florida Keys and another that was trapped illegally by a gang of Germans on the Mexican Gulf coast. By contrast, the Canadians, who have been releasing peregrines in northern Alberta and other northern locales, have had several of their birds reported from as far away as Mexico City and Belize south into northern South America (R. Pyfe, in litt.), exactly where one would expect them to go, following the natural migratory habits of the populations into which they have been introduced. Our mst distant sumer returns have been in southwestern Saskatchewan and, possibly, the west side of James Bay just south of Churchill.

So far, based on five nestings by three females, the timing of reproduction corresponds closely to the late March-April period of egg-laying that characterized the breeding season of the former duck hawks. Response to photoperiod and other seasonal timers of reproduction does not seem to be precisely influenced by genetic background, and earlier concerns about the complexity of the natural photoperiods exerienced by the Arctic migratory peregrines, for example, have proved to be groundless.

The released peregrines have adopted trophic relations virtually identical to those of the old duck hawks, too. Around their nests they feed heavily on blue jays and other small woodland birds and on feral pigeons and mourning doves. In coastal environments, especially in late sumer, fall, and winter, they feed on a wide variety of shorebirds and on some ducks, as well as on pigeons.

The peregrine is a well known generalist and opportunist, and it appears that there is sufficient behavioural and physiological plasticity built into the phenotype, so that adaptive adjustments to specific environmental conditions can be made regardless of the precise allelic composition of the genotype. This plasticity should make it much easier to establish a founding population from exogenous sources than would be the case were the genotypes highly selected to fit specific environmental configurations.

When possible, it probably is wisest to release only birds from indigenous stock, assuming that a large enough sample exists to avoid problems of inbreeding, as we are doing in our Rocky Mountain peregrine program. This rule can be carried to gross and unnatural extremes, however, when, for example, people insist that only peregrines from New Mexico should be released in New Mexico, or only Alberta birds in Alberta, Swedish birds in Sweden, and, of course, only Falco peregrinus germanicus in Germany--as though these political boundaries have some deep biological significance to peregrines. It could be that only falcons from western montane habitat should be released in the mountains of Colorado, or only coastal dwelling peregrines on the sea-cliffs of California; but even this ecotypic principle appears to have exceptions, judging from our experience in releasing a variety of ecotypes in the eastern states.

Ornithologists and conservationists mzt learn to deal with the world the way it really is. It is man-dominated, and it will only become more so. There is no such thing as a "natural ecosystem,” if by that term one means an ecosystem unaffected by man's activities and by the introduction of exotic plants and animals. Many forms of life face extinction in the next two decades. In our efforts to maintain the maximum possibile diversity of living things, we should not exclude the intelligently considered and careful introduction of exotica into new areas, as Charles Elton (1958) recognized same time ago in his little book on the invasions of animals and plants. The natural world is not the way it was in Britain before the arrival of civilized man; but there is still an out-of-doors full of a variety of plants and animals. The English experiencewith the naturalization of about 80 species of exotic vertebrates (Lever, 1977) shows that there are many benign benefits from their presence in the fauna, as well as some problems. If the California condor can only survive in the out-of-doors through introduction onto Santa Cruz Island, for example, or the Mauritius kestrel, on Reunion, those compromises are more acceptable, it seems to me, than extinction.

The notion that "endangered gene pools" of local populations are somehow pure and sacrosanct, and should be protected against gene exchange with other populations of the species, needs to be weighed carefully against the well known problems of inbreeding depression encountered by small, relict populations and the known benefits of increased heterozygosity and heterosis from outbreeding. What are we to do about the "gene pool" of the dusky seaside sparrow, which apparently is now represented only by male genotypes? It might be saved and reconstituted by natural selection if females from some other seaside sparrow population were introduced into the Dusky’s habitat, assuming that the habitat can still support a breeding population.

REFERENCES.
Bollengier, R. M., Jr., et. al. 1979. Eastern peregrine falcon recovery plan. U. S. Fish and Wildlife- Service. Mimeographed. 147 pp.
Clement, R. C. (ed.) 1974. Peregrine falcon recovery--proceedings. Audubon Conservation Report no. 4:1-36.
Corbin, K. W. 1977. Genetic diversity in avian populations. Pp. 291-302 in Endangered birds edited by S. A. Temple. University of Wisconsin Press.
Drury, W. H. 1974. Rare species. Biological Conservation 6(3):162-169.
Elton, C. 1958. The ecology of invasions by animals and plants. London: Methuen & Co., Ltd. 181 pp.
Lever, C. 1977. The naturalized animals of the British rsles. London: Hutchinson & Co. 600 pp.
Koenig, W. D., and Pitelka, F. A. 1979. Relatedness and inbreeding avoidance: Counterploys in the communally nesting acorn woodpecker. Science 206:1103-1105.
Mayr, E. 1963. Animal species and evolution. Harvard University Press. 797 pp.
Ralls, K., Brugger, K., and Ballou, J. 1979. Inbreeding and juvenile mortality in small populations of ungulates. Science 206:1101-1103.
Selander, R. K. 1971. Systematics and speciation in birds. Pp. 57-147 in Avian biology, vol. 1 edited by D. A. Farner and J. R. King. Academic Press.

APPENDIX 8

Giving some idea of the issues as they are perceived by the parrot-keeping community.

HYBRID BIRDS: SHOULD THEY BE CREATED ON PURPOSE? SOLD?
Catherine Quinones

From the WWW 1997

‘This is one topic that can generate incredibly heated arguments. The issue is whether hybrid birds (the result of the breeding of parents that belong to two different species) should be ‘made’ and if so, should they be sold to the public. I don’t claim to have covered all the points of this discussion, by the way.

For starters, lets agree on the definition of a species. A species is defined as a group of individuals that are more related to each other than they are to other such groups, and this hsa come about because of genetic (reproductive) isolation. This is to say that a set of individuals (a popualtion) have been breeding amongst themselves, and in doing so, natural selection, mutation and chance have caused each population to become genetically different from other populations (if there is migration among populations, that will have a certain effect in mixing things up.) By ‘different’ I mean that genetically, each popualtion will eventually develop distinct, characteristic frequencies of alleles, with allelles being the different ‘versions’ of given genes. One key reason those populations remain genetically distinct is that the populations do NOT intermingle, and if they do, they trade genetic material at a very low rate (this level is arbitrarily set).

At least part of the confusion comes from Mayr’s biological species concept, which spells out that the offspring of a mating among individuals belonging to different species is sterile; this DOES happen is some cases. Unfortunately, in using that definition, people often stick ‘to the letter of the law’ and fail to see ‘the spirit of the law.’ In such cases, the fertile hybrid is then seen as the ‘missing link’ that proves the unity of two otherwise discrete populations and the sterile individual is seen as the culmination of a series of gentic steps, the ultimate proof of true separation between the two populations. Yet, equally thorough genetic isolaiton can happen well before there is the production of infertile young. As long as the populations don’t mate, whether that is due to behavioural, physical or geographical reasons is irrelevant: genetic exchange doesn’t happen: period! Genetic isolation among populations is THE effective parameter in this discussion, as long as people are willing to define the species involved according to their behaviour and geographical range in their natural, UNDISTURBED habitats.

When we take up the discussion of whether two individuals capable of mating and producing viable, non-sterile offspring are the same species or not, we must decide WHY WE CARE about the outcome of the argument. To some people, the relevance hails to their belief that allowing a mating in captivity that would never happen in the wild IS doing a disservice to the evolutionary trajectory of each species/population in question. Alternatively, others argue that the fact that the breeding animals are no longer in their natural habitat already introduced a very chaotic factor into the population’s genetic futures, and that anything else that may happen in captivity and which doesn’t directly involve the wild populations is irrelevant to the future of the wild populations. Obviously, the people that support the first interpretation oppose hybridisation in captivity, while the others don’t. I am not trying to convince anyone that hybridisation in captivity is good or evil; I’m here simply trying to summarise the arguments offered by both sides.

There are some incontrovertible facts that also affect this discussion: many populations of wild birds are in imment danger of extinction due to the loss of their habitat. Even if sufficient habitat was secured and a population was established, population genetics models predict that, unless the population is sufficiently large, it will eventually become extinct due to imbreeding (which accelerates the expression of inheritable characteristics, whether they are good or bad.) Another fact is that there are representatives of at least some of these species being kept in captivity, and that some or most of these individuals are capable of breeding.

The following are arguments commonly offered by people who oppose the creation and sale of hybrids in captivity:

• Each bird that is bred to an individual of another species COULD have been breeding with a member of its own species and creating young that can help maintain genetic variability. (Remember that birds must form a bond with their mate: this takes time and is different from the more ‘casual’ mating rituals of other species).
• Each non-hybridised individual we have is a vessel that stores valuable genetic information. This genetic information may be used in the future to help natural populations recover from the damage that has been done by human intrusion (in the form of hunting, capturing birds for the pet trade, and destruction of habitat).
• ‘Pure’ (non-hybrid) birds are more beautiful that any hybrids anybody can create. Hybrids are aberrations that were not ‘meant to be.’
• If hybridisation doesn’t occur in the wild, it shouldn’t be allowed to occur in captivity.
• We still don’t know a lot about breeding birds in captivity, inparticular about the nutritional requirements of baby birds, and we may be using nutritionally incomplete formulae. Hybrid birds may have different requirements from what non-hybrids of either parent species are believed to need, and because of our ignorance we may not properly nurture hybrid offspring.
• There is no way to ENSURE that a hybrid bird will not be bred. Even if the owner swears they won’t breed the bird, unforeseen circumstances may come into play. Ask any breeder and they will tell you that many of their breeding birds were formerly pets: that means a lot of birds DO get passed on to others. The same thing may happen with a hybrid, and the new owner may not be as careful about upholding the promise not to breed the bird.
• The reports of hybrid parrots occurring in nature are rare. Many such matings may have occurred due to human intervention, or disturbance of natural habitats, or to people facilitating the mating of animals that would otherwise prefer to mate with their own species. May such reports are anecdotal and unverified. Also, hybrids are rare and only occur in the zone where the geographic boundaries of two species meet. Hybridisation s NOT a common occurrence nor is it part of the ‘natural scheme of things’ in particular when the species involved DO NOT have overlapping geographical ranges!

People who don’t think creating hybrids is a problem offer the following counterarguments (numbered in correspondence to the item they dispute):

• Not all birds kept in captivity are capable of breeding. Hand-fed birds in particular seem to have a high incidence of showing no interest in bonding with members of their own species. Even if they were capable of breeding, their owners may not be interested in allowing them to be bred. Because of these variables, it is not truthful to say that each pet bird counts in this conservation effort.
• There is no unified program controlling the matings that take place in captivity. For the most part, no records were kept of where the initial breeding stock originated, so for all practical purposes we may have lots of hybrids in the pet trade that are believed to be ‘pure’ representatives of one species or another. If the anti-hybridisation people don’t feel those distinctions are important, then they should be just as happy with repopulating the habitats with whatever parrots can be had, regardless of their species or of whether they are hybrids.
• Beauty is in the eye of the beholder, and some people DO like the colouration of some hybrids. For instance, there are many colour mutations in some species of captive birds (e.g. budgerigars) that, although they didn’t originate from hybridisation, are very beautiful in their own right. Hybridisation (mixing of genetic material from two distinct groups or species) has had a place in the domesticaiton of every grain we eat and also played its part in the domestication of most or all of our farm and companion animals. Hybridisation has been a key part in the domestication process and perhaps it will help us create pet birds that are more comfortable and happy in a human home.
• There are lots of things that don’t occur in the wild, like having a veterinarian care for the animals, and offerring a balanced diet, and keeping the birds in human homes. Or keeping birds in cages and clipping their wings. Captivity IS an unnatural circumstance, and whatever birds are bred in captivity are NOT ‘playing by the rules’ that regulate the lives of wild birds. Because of this, we may in fact be breeding birds that won’t have what it takes to survive in the wild.
• Hybrid chicks seem to be doing just fine. If they weren’t, they wouldn’t be for sale, and we wouldn’t be having this discussion, right?
• Hybrids occur in nature. Hybridisation is natural. Don’t accuse me of failing to care for my bird for the length of its lifespan, or of providing for its care if I can’t do so, you don’t know me, blah blah blah etc.
• There aren’t very many hybrids being created, therefore their numbers don’t pose a threat to any ‘domestic breeding for future repopulation’ efforts. Besides, there will be no return to the natural world as it was, so we must start talking about modern evolution, which happens when humans intervene.

The counter-counter argument to that is something along the lines of ‘fine, even if all that is true, AND the world is going to hell in a hand-basket, AND there is a hole in the ozone, AND the rainforests are gone, not creating hybrids AT LEAST DOESN’T COMPOUND ON THE PROBLEM and COULD help preserve species as we know them.’

As it stands, making, selling, nor owning hybrids is illegal. I think most rational people can see that there ARE arguments for both ‘pro’ and ‘con.’ Ultimately, as long as there is a market for hybrids, there will be hybrids. As is, I can’t imagine domestic breeders as a whole ‘saving’ any wild populations via their breeding stock because of point #2 above. As long as there is no agreed, encompassing course of action, all we have is a bunch of people running their own breeding and domestication programs (programs that may include inbreeding, hybridisation, breeding of unsuitable or unhealthy birds, dubious record keeping...) Which is to say, the great majority of breeders (and the birds they own and produce) are NOT (and probably CAN’T be) part of pro-conservation army, because more likely than not they are breeding PETS for the pet TRADE, or at worst, are just pumping out a commodity the market demands. Most certainly they are not breeding wild birds meant to be the basis of a re-release program.

With that said, I hope people think about these issues before they buy a hybrid bird strictly on its appearance. As is the case with other purchases, people need to decide if they are OK with being part of the pro-hybridisation crowd because whenever a hybrid is purchased, the buyer IS supporting hybridisation in general. Likewise, let us all remember that the birds are NOT to blame for what is going on. Hybrids deserve as much loving care and respect as any other animal. Their owners likewise don’t deserve to be ostracized becaus (1) they may have NOT been informed that they were buying a hybrid (both ignorance and false advertisement can be to blame on this one), or (2) they may have rescued the animal - meaning they were not directly, deliberately contributing to the demand for hybrids - and (3) because, after all, the impact of domestic hybridisation on parrot conservation hasn’t been assessed.

APPENDIX 9

SOME SPECIES DEFINITIONS

From notes by

R. M. Zink March 1996,

Bell Museum, 100 Ecology Bldg., Univ. of Minnesota, St. Paul, MN 55108

1, 'No one definition has as yet satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species' (Darwin 1859).

2. 'A species is a set of populations capable of combining with each other but not with other similar sets of populations on the basis of affinity and co-direction in ecological speciation.' (Shaposhnikov 1966: 13)

3. 'A species is a group of organisms not itself divisible by phenetic gaps resulting from concordant differences in character states (except for morphs such as those resulting from sex, caste, or age difrerences), but separated by such phenetic gaps from other such groups.' (Michener 1970: 28)

4. ‘We may regard as a species (a) the smallest (most homogeneous) cluster that can be recogmzed upon some given critenon as being distinct from other such clusters, or (b) a phenetic group of a diversity somewhat below the subgenus category, whether or not it contains distinct subclusters.' (Sneath and Sokal 1973: 365)

5. ‘Somit ist die Art als das Kollektiv von Lebewesen zu bestimmen, das gemeinsam eine okologische Nische behauptet.' (von Wahlert 1973: 249)

6. 'Species may then be defined as groups of phenetically similar populations that have the capability to interbreed, and share similar ecological characteristics.' (Doyen and Slobodchikolf 1974: 240)

7. ‘Species, then, are the most extensive units in the natural economy such that reproductive competition occurs among their parts." (Ghiselin 1975: 538)

8. ‘A species is a lineage (or a closely related set of lineages) which occupies an adaptive zone minimally different from any other lineage in its range and which evolves separately from all lineages outside its range.’ (Van Valen 1976: 233)

9. 'Species are the smallest groups that are consistently and persistently distinct, and distinguishable by ordinary means.' (Cronquist 1978: 15)

10. ‘A species is a single lineage of ancestral descendant populations of organisms which maintains its identity from other such lineages and which has its own evolutionary tendencies and historical fate." (Wiley 1978: 18)

11. ‘A ‘species’ is merely a population or group of populations defined by one or more apomorphous features, it is also the smallest natural aggregation of individuals with a specifiable geographic integrity that can be defined by any current set of analytical techniques.' (Rosen 1979: 277)

12. ‘A species is a group of animals or plants all of which are similar enough in form to be considered as minor variations of the same organism. Members of the group normally interbreed and reproduce their own kind over considerable periods of time.' (Trueman 1979: 764)

13. 'A species is a diagnosable cluster of individuals within which there is a parental pattern of ancestry and descent, beyond which there is not, and which exhibits a pattern of phylogenetic ancestry and descent among units of like kind.' (Eldredge and Cracraft 1980: 92)

14. ‘Species are simply the smallest detected samples of self-perpetuating organisms that have unique sets of characters.' (Nelson and Platnick 1981: 12)

15. ‘Each species is an internally similar part of a phylogenetic tree." (Willis 1981: 84)

16. 'We can, therefore, regard as a species that most inclusive population of individual biparental organisms which share a conunon fertilization system." (Paterson 1985: 25)

17. ‘An ‘evolutionaryspecies’ is a single lineage of ancestor-descendant populations which maintains its identity from other such lineages and which has its own evolutionary tendencies and historical fate.’ (Wiley 1978: modified from Simpson 1961)

18. ‘A species is what a good taxonomist says it is.' (anon.)

19. ‘At the outset I confess a disbelief in species, as that word is commonly understood to refer to the basic taxonomic unit or to the taxonomic unit of evolution... There seem to be no basic taxonomic units and no particular taxonomic unit of evolution... and as Agassiz said in 1859 'species do not exist in nature in a different way from the higher groups’ (Nelson 1989).

20. '... species as the most inclusive group of organisms having the potential for genetic and/or demographic exchangeability' (Templeton (1989).

2I. ' a species is 'the smallest aggregation of populations (sexual) or lineages (asexual) diagnosable by a unique combination of character states in comparable individuals (semaphoronts)’ (Nixon and Wheeler 1990).

22. Species 'refer to groups of actually or potentially interbreeding populations isolated by intrinsic reproductive barriers from other such groups. Evidence for reproductive barriers . . will involve concordant genetic differences among the populations involved [...] Subspecies are groups of actually or potentially interbreeding populations phylogenetically distinguishable from, but reproductively compatible with, other such groups, Importantly, the evidence for phylogpnetic distinction must normally come from the concordant distributions of multiple, independent, genetically based traits' (Avise and Ball 1990).

23. ‘Species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups' (Mayr 1942).

24. 'A species is a reproductive community of populations (reproductively isolated from others) that occupies a specific niche in nature' (Mayr 1982).

25. 'A species is the smallest diagnosable cluster of individual organisms within which there is a parental pattern of ancestry and descent' (Cracraft 1983).

26. ‘Species are "lineages whose components (if distinguishable) are not incontrovertibly on different phylogenetic trajectories (i.e., sublineages, if distinguishable, are reproductively compatible), as long as these sublineages do not form a paraphyletic group in recovered history. [...] The species category ... would represent the largest entities that have evolved whose parts, if distinguishable, are not likely to be on dilterent phylogenetic trajectories' (Frost and Hillis 1990).

27. 'if a given historical group of hybridogens is persistent and is not affecting the evolutionary trajectory of its Mendelian ancestor (as indicated by biogeography, habitat preferences, or genetic divergence), it should be considered a separate species' (Echelle 1990).

28. ‘A species is the 'smallest recognizable monophyletic or unresolved unit’’ (Donoghue 1985). [see also de Quciroz, K. & M. J. Donoghue. 1988]

29. 'Phylogenetic species can be delimited by a procedure (population aggregation analysis) that involves a search for fixed differences among local populations, followed by successive rounds of aggregation of populations and previously aggregated population groups that are not distinct from each other [...] descent relationships among [phylogenetic species] must be hierarchic'. (Davis and Nixon 1992).

30. See # 1.

References

Avise, J. C., & R. M. Ball. 1990. Principles of genealogical concordance in species concepts and biological taxonomy. Oxford Surveys Evol. Biol. 7:45-67.

Cronquist, A. 1978. Once Again, What is a Species? In: Biosystematics in Agriculture. Ed. by Romberger, J. A. pp. 3-20.

Cracraft, J. 1983. Species concepts and speciation analysis.Current Ornithology 1: 159-187.

Darwin, C. 1859. On the origin of species. London: John Murray.

Davis, J. I. & K. C. Nixon. 1992. Populations, genetic variation, and the delimitation of phylogenetic species. Syst. Biol. 41:412-435.

de Queiroz, K. & M. J. Donoghue. 1988, Phylogenetic systematics and the species problem. Cladistics 4:317-338.

Donoghue, M. J. 1985. A critique of the biological species concept and recommendations for a phylogenctic alternative. The Bryologist 88:172181.

Doyen, J. T., & C. N. Slobodchikoff. 1974. An operational approach to species classification. Syst. Zool. 23:239-247.

Echelle, A. A. 1990. In defense of the phylogenetic species concept and the ontological status of hybtidogenetic taxa. Herpetologica 46:109-113.

Eldredge, N. & J. Cracraft. 1980. Phylogenetic Patterns and the Evolutionary Process. New York. Columbia Univ. Press.

Frost, D. R, & D. M. Hillis. 1990. Species in concept and practice: herpetological applications. Herpetologica 46:87-104.

Ghiselin, M. T. 1975. A radical solution to the species problem. Syst. Zool. 23:536-544.

Mayr, E. 1942. Systematics and the Origin of Species. Columbia Univ. Press.

Mayr, E. 1982. The Growth of Biological Thought. BelknapPress, Harvard University.

Michener, C. D. 1970. Diverse Approaches to Systematics. Evolut. Biol. 4: 1 38.

Nelson, G. 1989. Species and taxa: systematics and evolution. Pp. 60-84 in D. Otte and J. A. Endler (eds.), Speciation and its consequences. Sinauer Assoc. Inc., Sunderland, Mass.

Nelson, G. & N. Platnick. 1981. Systematics and Biogeography. New York. Columbia Univ. Press.

Nixon, K. C., & Q. D. Wheeler. 1990. An amplification of the phylogenctic species concept. Cladistics 6: 211-223.

Paterson, H. E. H. 1985. The Recognition Concept of Species. In: Species and Speciation. Ed. by Vrba, E. S. olt. cit pp. 21-29.

Rosen, D. E. 1979. Fishes from the uplands and intennontane basins of Guatemala: revisionary studies aid comparative geography. Bull. Am. Mus. Nat. Hist. 162:267-376.

Shaposhnikov, G. Ch. 1966. The origin and the breakdown of reproductive isolation and the species criterion. Entomol. Rev. 45:1-18.

Sneath, P. H. A., & R. R. Sokal. 1973. Numerical Taxonomy. San Francisco. W. H. Freeman.

Templeton, A. R. 1989. The meaning of species and speciation: a genetic perspective. Pp. 3-27 in D. Otte and J. A. Endler (eds.), Speciation and its consequences. Sinauer Assoc. Inc., Sunderland, Mass.

Trueman, E. R. 1979. Species concept. In: The Encyclopedia of Paleontology. Ed. by Fairbridge, R. W.; Jablonski, D., Stroudsburg, Pa.: Dowden, Hutchinson & Ross. pp. 764-767.

Vaten, L. van. 1976. Ecological species, multispecies, and oaks. Taxon 25:233-239.

Wahlert, G. von. 1973. Phylogenie als okologischer Proecls. Naturwiss. Rundschau 26:247-254.

Wiley, E. 0. 1978. The evolutionary species concept reconsidered. Syst. Zool. 27:17-26.

1981. Phylogenetics-The Theory and Practice of Phylogenetic Systematics. New York. J. Wiley.

Willis, E. 0. 1981. Is a species an interbreeding unit, or an internally similar part of a phylogenetic tree? Syst. Zool. 30:84-85.

[1] Heywood, ‘Invasions by terrestrial plants’, 40; US Congress, Harmful Non-indigenous Species, 95. See also Drake, J. A. et al. 1989. Biological Invasions: A Global Perspective. SCOPE Report 37, John Wiley, New York.

[2] Ebenhard, T. 1988. Introduced Birds and Mammals and their Ecological Effects. Viltrevy 13 (4): 3-76; see also Long, J.L. 1981. Introduced Birds of the World, David and Charles, London.

* Note: Saar’s 1994 report refers to hybridisation in captivity, not in the wild. (Bednarek, pers. comm.))

* Note: see case history of the female (Ed).

[3] See section 13

[4]Lever, C. 1977. The naturalised animals of the British Isles. Hutchinson, London

[5] Recommendation No. R(84)14 adopted on 21 June 1984

[6] Recommendation No. R(85)15 adopted on 23 September 1985

[7] Biodiversity: The UK Steering Group Report

Volume 1: Meeting the Rio Challenge HMSO ISBN 0 11 7532185

Volume 2: Action Plans HMSO 1995 ISBN 0 11 7532282

[WD1]