Nature versus nurture: the relative importance of genetic and environmentally determined grazing characteristics erosion

David Anthony Murray (presently unpublished paper), August 2011

Introduction:
The physical attributes, hardiness and grazing characteristics of British native breed ponies illustrate their potential value to plant and animal wildlife and habitats. Anecdotal evidence from the gradually increasing day-to-day use of native ponies in official grazing schemes and projects throughout the United Kingdom, coupled with significant, albeit still insufficient, empirical studies, are gradually confirming the extent of their usefulness as conservation grazers. This usefulness might be extended in view of changing British climate. Native ponies, particularly those sub-populations living in semi-feral, extensive grazing, acclimatised, closed herd system - no breeding mares are brought into the herd from another herd - are likely to be suitable conservation grazers in a wide variety of grazing habitats and landscapes. Their grazing conservation value is at least partially dependant upon the extent to which breed genetic characteristics, manifested through the breeds variety of breed-lines, have been protected, preserved and conserved.

Native breed ponies: a definition

The Fell pony, together with several other pony breeds, is a British native breed, which is, by definition, included in the UK National Breed Inventory (6). Firstly, it must be recognised as a livestock breed, an interbreeding population of husbanded or formerly husbanded domesticated animals of consistent genotype and phenotype with a recognised history and administrative framework (6). Secondly, it fulfils, or potentially fulfils, a role in the rural economy. Thirdly, less than 10% of the aggregate genetic contributions to the population over 4 generations are derived from other resources distinct from foreign herd books recognised as representing the same breed (6). Fourthly, breed history documents the breed origin within the UK and the UK has formed the primary environment for the development of the breed. Additionally, breed history documents its presence in the UK in its current adapted form for (at least) 40 years and 6 generations (6). Finally, maximally 20% of the genetic contributions come from animals born outside the UK (other than those imported for an approved conservation project) in any one generation for the last 40 years plus 6 generations (6).

Extensively grazing ponies: a definition

The Rare Breeds Survival Trust defines a feral breed as a breed which is not subject to routine handling of any kind; and more than 90% of the population have been born to feral parents Semi-feral populations (within a native breed) have some level of human input to their husbandry (RBST, 2011). I have previously discussed the meaning of feral in the context of British native ponies with reference to the Section A Welsh Mountain pony, which I considered should be more properly regarded as semi-feral (3): There are no true feral, unhandled and not owned, ponies living in Wales. A semi-feral pony is not fully tamed or brought under control by breeders. It is owned but handled only occasionally for weaning and welfare purposes, is allowed to run in natural herds, cannot be haltered, is not fed by the public and grazes extensively on unimproved land Differences between the registered semi-feral breed population and the remainder of the Section A breed are a source of debate. A key distinction between these and other ponies of the same breed is that semi-feral ponies are extensively grazing, live within a closed system and are acclimatized within that system. They are hefted animals. Some UK scientists do not recognize the concept of semi-feral from a genetic perspective. I propose that, in future, sub-populations should be identified more precisely by a set of criteria that define native pedigree ponies as either extensively grazing (if they are also hefted within a closed system) or non-extensively grazing. Particularly, whether extensively grazing Welsh Mountain ponies, and similar subpopulations of several other native breeds including the Fell pony, represent genetically distinct groups from ponies that may not be described as extensively grazing is a matter of considerable confusion. A clearer understanding of the genetic spectrum within pony breeds and between breeds is likely in future, now that genomes of each breed and individual breed lines can be assessed. At present, the genome of the Fell pony is being compared with that of over 30 other breeds at the College of Veterinary Medicine, University of Minnesota (1 - Personal Communication). To determine genetic differences between these two distinct sub-populations within the Fell pony breed, if they exist, controlled studies separating genetic material of extensively grazing and non-extensively grazing populations are needed. Scientific and conceptual difficulties associated with properly defining the term extensively grazing will be critically assessed in a future paper. However, the use of this term becomes immediately useful when considering genetic erosion and environmental erosion, particularly, but not exclusively, within the Fell pony breed.

Genetic Grazing Characteristics Erosion

The breeding of any native pony breed by that breeds official society members, and sometimes unofficially by organisations or individuals, has historically and generally been determined by the ponys contemporary uses. If there is a perceived human need for a breed it will survive, often aided by human genetic manipulation. Uniformity of appearance of native breeds, including extensively grazing subpopulations, has been established largely by prescriptions laid down by breed societies over a century ago. Historically, a breed populations profile and its perceived and actual use have favoured some characteristics and some animals, which ultimately has, to greater or lesser degree, limited, that is, reduced, biological diversity (biodiversity). This gradual loss of biodiversity within a breed may be regarded as genetic erosion by the voluntary stripping of genetic assets (3, p.118). Genetic erosion, its extent, causes and consequences, has been little researched and is often poorly understood by native pony breeders. A presently increasing, but at any time more or less limited, use of hardy, free-living, extensively grazing native breed ponies by official conservation organisations and groups for specific conservation grazing regimes and schemes is well documented. In any specific grazing situation specific habitats, vegetation types and geographical indicators will usually be recognised. Some breed-lines and some breeds will be more optimally suited to a given, clearly defined grazing situation than others. Personal pony preferences of land managers are sometimes based upon inconclusive empirical or anecdotal evidence are determined by breed availability. Too often what is deemed fashionable is chosen or selection is determined by excessive pro-activity of a particular breed society, which ensures that a particular breed or breed line is preferentially and repeatedly selected to help manage an identified habitat type. However, the chosen breed may not be the most optimally suited for the predominant, local grazing conditions. If any breed or breed-line is favoured, i.e. used more frequently than another for habitat conservation purposes, whether for scientifically justifiable or other reasons, the ultimate, long-term survival probability of that breed or breed-line will be increased. Conversely, any extensively grazing breed population, or breed line within that population, that is not used in grazing schemes is more likely to face extinction over a greater or lesser time period, unless government support programs can ensure the survival of genetically viable herds. Loss of a semi-feral breed-line leads to enhanced and rapidly accelerated genetic diversity reduction, referred to in previous discussions as genetic erosion. Within the context of appreciating native breeds as important conservation grazers, genetic erosion might be more appropriately described as genetic grazing characteristics erosion. This must be halted immediately because important hardiness and feeding abilities might relatively quickly not be retrievable, in view of economic and logistical constraints of husbanding extensively grazing herds. Of course, frozen sperm and egg storage now allows the ex-situ conservation of important genes for future generations. However, in-situ ponies are constantly

adapting to changing geographical conditions, sometimes aided by man, sometimes by equine instinct, allowing specific genes to be emphasised that aid survivability, whereas adaptation of a future ex-situ breed line will be halted throughout the freezing period. Lack of adaptability also has relevance to learned behaviour that is passed between generations.

Environmentally Determined (Learned) Grazing Characteristics Erosion

Perhaps of equal importance to the need to conserve genetic grazing characteristics in semi-feral native ponies, is a less readily appreciated challenge of minimising and if possible halting the rapidly accelerating loss of environmentally determined grazing characteristics. For example semi-feral foals are hefted, i.e. learn from their mothers how and what to graze throughout different seasons. During the last two decades the disappearance of native Fell pony breed lines from their traditional commons, moorlands, mountains and marshes has become more common-place. When native ponies are removed from a defined grazing situation where learned behaviour is the consequence of many generations of experience, whether or not the original grazing behaviour can be quickly relearned by introducing a new generation later on to that regime is uncertain. Little research has attempted to assess the importance of learned behaviour upon grazing effects. Neither do we understand possible consequences upon vegetation, habitat and wildlife, of introducing other British or foreign breeds, for example, the Polish Konik pony, into geographical areas previously managed by an endemic population of a local indigenous breed. Recently, for example, Exmoor ponies have been introduced into northern England, particularly into Cumbria. The trend of not using the pony of local origin, which may have been indigenous to an area for centuries, must be resisted and reversed. I propose that environmentally determined or learned erosion, as defined in this paper, should be clearly identified in future discussion and study of conservation grazing as a distinctive form of biological diversity loss called Learned Grazing Characteristics Erosion. Controlled studies are urgently needed to assess the effects of losing extensively grazing breed-lines from specific habitats which they have grazed, in some instances, for at least several centuries. Put another way, will native herds or individuals be able to adapt easily, quickly enough or sufficiently to the full spectrum of unfamiliar grazing conditions during any future agricultural crisis? Such a crisis might be triggered by a consequence of mixed grazing involving only cattle and sheep or by accelerated local climate change due to, for example, localised seasonal warming or increased precipitation. An assessment of the extent to which learned grazing characteristics may be affecting observed grazing outcomes involving extensively grazing ponies is urgently needed.

The Fell pony: a breed only AT RISK?

The current Rare Breeds Survival Trust classifies the Fell pony breed on its 2011 Watchlist (4) as At Risk based upon the total number of registered breeding females, because there are at least 900 registered Fell ponies throughout the UK, most of which cannot be considered to be semi-feral and which are not grazing extensively. However, only 135 foals were born to semi-feral Fell ponies this year and this number may be significantly reduced if a test of whether these foals are extensively grazing is made. On this basis alone, and taking into consideration the perceived importance of environmental characteristics in addition to genetic characteristics to conservation management of habitats, an additional classification is urgently needed for extensively grazing sub-populations of the Fell pony (and all other native pony breeds) which is based upon appropriate empirical and anecdotal evidence and which takes into account learned grazing characteristics. Additionally, if an extensively grazing sub-population is proven to be genetically distinct, this sub-population should receive a special classification. Should the Fell pony extensively grazing subpopulation be regarded as Critical (there are many fewer than 300 registered breeding females) based upon current RBST Watchlist criteria?

References
1. College of Veterinary Medicine, University of Minnesota, St Paul, MN 55108 USA, Personal Communication, 2011, 1 page. 2. MURRAY, D. A. 2005. The Fell pony: grazing characteristics and breed profile a preliminary assessment. A feasibility study on the potential role of Fell ponies in conservation grazing post Common Agricultural Policy reform. Findings of surveys conducted in 2003 and 2004. 2005. A report prepared for Defra. British Library Catalogue No. 636.16, 399 pages. Published by David Anthony Murray, 31-05-05, 399 pages. 3. MURRAY, D. A. 2006. Current status, with reference to conservation grazing, of the Welsh Mountain (Section A) semi-feral pony following decoupling of farm subsidies through CAP reform in Wales a scoping study. Findings of a survey conducted in 2006. Catalogued with the British Library. A report prepared for The Countryside Council for Wales. Published by David Anthony Murray, 31-10-06, 380 pages. 4. Rare Breeds Survival Trust, 2011, Guidelines for acceptance on the RBST Watchlist, www.rbst.org.uk, 3 pages. 5. http://www.intl-pag.org/19/abstracts/P05o_PAGXIX_617.html, Plant and Animal Genomes XIX Conference, 15-19 January, 2011, 2 pages. 6. http://www.defra.gov.uk/fangr/files/definitions-breeds-2011.pdf. Definition of a breed for the purpose of the UK National Inventory D A Murray, 2011