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In consequence, any F1 hybrids produced were initially most likely to backcross into red deer, given their higher local population density compared to sika. The fact that there are mitochondrial hybrids which are pure sika at the nuclear markers but carry red deer mtDNA suggests that extensive backcrossing into sika also subsequently occurred. Regarding other parts of Scotland, only 0. There was no evidence for hybrids among phenotypically red deer sampled across the Central Highlands, even among 50 animals collected from Ralia, Abernethy, Inshriach, and Kinveachy on the western borders of the Cairngorm National Park into which sika are expanding.
Within the Lake District, only three of 2. A single putative modern F1 was identified at WLA which, if true, was the result of a sika male mating a red deer female. In the two areas with detected hybrids Counties Wicklow and Cork , no F1 hybrids were found. Overall, while introgression from wapiti is not currently a threat to red deer in Scotland, largely due to the modest numbers introduced and their failure to thrive, hybridization with sika is a threat, although it appears to be rather unpredictable as to where it occurs and more importantly, what the consequences are.
In some areas, hybridization is followed by a breakdown of assortative mating and the generation of a hybrid swarm e. For introgression to occur, a male of one species has to encounter, defend, and mate with a receptive female of the other species an event which all the evidence suggests has very low probability , and the hybrid offspring has to survive and itself reproduce, events which may also have modest probabilities, especially for males. Before making detailed management suggestions, there are a number of points with practical implications that are worth noting. Explanations for why hybrids do not occur in Killarney include that there is an unusually large physical difference in body size between red and sika in the area, and that there are sufficient females of both species present.
Second, and related, it is widely hypothesized that hybridization occurs when dispersing males of one species find themselves in an area without females of the other species. Third, identification of hybrids from phenotype is very difficult in the field, making efficient selective culling difficult. Fourth, nevertheless, culling by shooting is the most practical tool available by which the situation may be managed. Deer fencing is very expensive over the large distances and rugged landscapes involved in Scotland and has limited effectiveness over time and where snow accumulates, as in many parts of Scotland.
Moreover, in the area where we might be most concerned to contain hybrids by fencing, Kintyre, deer are quite frequently seen swimming between land masses Kevin McKillop. Live capture and testing followed by selective culling based on genotypes, as is suggested for the endangered bontebok Damaliscus pygargus pygargus being introgressed by common blesbok D. Although shooting is thus the most practical tool for deer management, it should also be noted there is probably an optimum level of culling that minimizes emigration—many deer managers believe that shooting at a population too hard can actually encourage dispersal, meaning that the spread of hybrids could be exacerbated.
Preventing further hybridization and introgression involves a number of steps 1 prevention of initial hybridization events 2 prevention of breeding by F1 and introgressed individuals and 3 prevention of the spread of heavily introgressed animals out of Kintyre and into the rest of mainland Scotland.
In addition 4 the red deer refugium in the Hebrides should be maintained free of sika introgression. To prevent initial hybridization, the best evidence suggests that selective culling of pioneering sika males, which have dispersed out of areas with sika females and into areas with only red females, is the best strategy. To prevent breeding by F1s and introgressed individuals, extreme vigilance for and culling of individuals with intermediate characteristics is required.
With regard to the hybrids in Kintyre, the swarm at WLA is the furthest north and in greatest danger of sending dispersing males into the rest of mainland Scotland. Deer managers in North Kintyre, the adjacent Cowal peninsula, and in Argyll in general should therefore be especially vigilant for sika and intermediate animals moving north and east from WLA. For all three of these suggested actions, improved training of deer managers on the phenotypic appearance of sika and hybrids compared with red deer would be highly desirable. The only way to reveal whether this is the case would be to use a large panel of markers, probably single nucleotide polymorphisms, SNPs , giving reliable estimates of an individual's genetic status after many more generations of backcrossing.
Bearing in mind the number of fixed markers required to define an individual as introgressed versus pure doubles with each generation of backcrossing, and that introgression may have been happening for several generations, several hundred or thousands of markers will be required. Second, although this study covers much of the Scottish Cervus deer range, there are gaps, of which the most notable is Dumfries and Galloway, in south west Scotland, where the red deer could have been introgressed by sika from the population that has expanded from the Dawyck, Peebleshire, introduction Ratcliffe, Thanks to Eleni Socratous, Eleanor A.
Graham, and Guy N. David Coltman University of Alberta for the provision of wapiti samples. Introgression of exotic Cervus nippon and canadensis into red deer Cervus elaphus populations in Scotland and the English Lake District. National Center for Biotechnology Information , U. Journal List Ecol Evol v.
Published online Jan Smith , 1 Helen V. Wyman , 4 Elizabeth Heap , 1 and Josephine M. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
Open in a separate window. To assess the current extent of hybridization and introgression between introduced Cervus deer and red deer across Scotland and in the Lake District. To suggest future management actions that could prevent further hybridization and introgression. Data analysis The Bayesian clustering software structure 2. Mitochondrial introgression is restricted relative to nuclear markers in a water strider Hemiptera: The problems with hybrids: Reproductive performance of pubertal red deer Cervus elaphus hinds: Animal Reproduction Science , 90 , — Hybridisation between red and sika deer.
Zoologischer Anzeiger , , — Asymmetric and differential gene introgression at a contact zone between two highly divergent lineages of field voles Microtus agrestis. Journal of Evolutionary Biology , 25 , — Genetic analysis of hybrid zones: Numbers of markers and power of resolution. Ecology , 78 , — Red deer stocks in the Highlands of Scotland. Nature , , — A mitochondrial control region and cytochrome b phylogeny of sika deer Cervus nippon and report of tandem repeats in the control region.
Molecular Phylogenetics and Evolution , 12 , 47— Maintenance of species boundaries despite rampant hybridization between three species of reef fishes Hexagrammidae: Implications for the role of selection. Biological Journal of the Linnean Society , 91 , — Is there evidence of recent or past hybridization with red deer Cervus elaphus? Journal of Zoology , , — Molecular Ecology , 14 , — Inference of population structure using multilocus genotype data: Linked loci and correlated allele frequencies.
Genetics , , — Dominant markers and null alleles. Molecular Ecology Notes , 7 , — Introgression through rare hybridization: A genetic study of a hybrid zone between red and sika deer genus Cervus in Argyll, Scotland. Bottlenecks, drift and differentiation: The population structure and demographic history of sika deer Cervus nippon in the Japanese archipelago.
Molecular Ecology , 10 , — Hybridisation among deer and its implications for conservation. Irish Forestry , 30 , 64— In its native range, C.
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The red deer introduced to the New Zealand and Australia in the 19 th century came from Britain, while those introduced to South America came from western Europe; they were therefore all part of the western group of C. These source populations often came from managed herds on game estates and were themselves often of mixed provenance. In New Zealand, the wild C. Red deer farming began in New Zealand in the s Nugent and Fraser, and active hybridisation with wapiti to increase body size was encouraged Sika deer are not farmed in New Zealand.
The modern ability to import semen or embryos has also facilitated the mixing of these original British red deer populations with larger phenotypes from eastern Europe and with wapiti Pearse and Goosen, Despite legal standards for deer fencing in New Zealand, escapes are common Fraser et al.
This genetic mixing is also happening in South America -- a recent photograph of a wild deer in Argentina with clear C. When liberated in the southern hemisphere, seasonally breeding species such as C. The amount and quality of food per capita determines maternal body condition which determines age at first breeding.
When animals are in good condition a proportion of females may first breed as yearlings. Males attain sexual maturity as yearlings, but they continue growing until at least 6 years of age and cannot compete for females with other males until then IUCN, The natural lifespan of C. In Europe excluding Russia, the species numbered 1. In New Zealand, the density of C. The intensity of this commercial exploitation has declined in the 21st century, so populations are generally increasing Forsyth et al. There are no estimates of the population size or density of C.
In their introduced range, C. In New Zealand they have no non-human predators. At natural densities they are spillover hosts of tuberculosis and only maintain the disease when held at artificially high densities, e. Deer are also capable of being infected with foot and mouth virus, but as with tuberculosis they appear not to maintain the disease in the absence of infected livestock Davies, In general wild C.
In New Zealand, of 80 deer harvested from the wild between and and inspected at processing works, only 2. Natural dispersal rates of 1. In such case they may cause damage to crops e. However, as in other countries the actual costs of such impacts have not been assessed, perhaps because they are relatively small.
Cervus elaphus (red deer)
It is possible that the route of infection of bovine tuberculosis from cattle to the wild maintenance host in New Zealand the brushtail possum, Trichosurus vulpecula was via carcasses of infected deer, which were prevalent in many areas during the commercial harvesting boom of wild deer during the s and s, and scavenged by possums Nugent et al. In the native range of C. Gill and Fuller, The major adverse impact of introduced red deer is on native vegetation; in turn, changes in the structure and composition of vegetation affect native animals.
Most of the research on impacts on vegetation has been conducted in New Zealand where the native vegetation evolved in the absence of mammalian herbivores. However, the vulnerability of deer to helicopter hunting in these grassland areas enabled their virtual elimination deer by the s, allowing significant recovery of the vegetation Rose and Platt, Deer grazing in these alpine grasslands is thought to be a significant threat via competition for food to the endangered, flightless gallinule, the takahe Porphyrio hochstetteri Mills and Mark, The impact of deer in forest ecosystems in New Zealand is complex and may have long-term consequences which may not always be reversible even if deer numbers are reduced Coomes et al.
For example, Nugent et al. The deer then rely to a large extent on leaves of palatable species that fall from the canopy. This buffers the expected feedback of declining food availability in the browse tier on deer density and so inhibits the recovery of these understorey palatable plants. Of course the new balance between deer and their food can last only as long as the canopy trees live — and in New Zealand this is unnaturally shortened for palatable species because of the impacts of another introduced herbivore, the arboreal brushtail possum Trichosurus vulpecula.
A recent study modelled the long-term consequences of deer and introduced mice as granivores in New Zealand podocarp forests, suggesting that the changes in structure and composition will take centuries to resolve Forsyth et al. Determining the impact of deer on these communities is difficult as they are sympatric with native macropod herbivores, and fire and rainfall impose significant extrinsic influences on such plant communities.
The only study of C. They showed that deer had nearly eliminated one common subcanopy tree Aristotelia chilensis , had reduced the abundance of many other woody and herbaceous species, and impeded the regeneration of the dominant canopy trees after canopy gaps are created. It has been suggested that C. Unlike some cervids e. Donne, , where such hunting is the main contributor to the annual harvests of deer. Introduced red deer are exploited for productive purposes in three ways. The harvest size is determined largely by the price of venison in the European market and the costs of harvesting.
The annual harvest is currently about 17 Warburton et al. Red deer are also held in semi-natural states on large private ranches where trophy hunters pay for access to large-antlered stags. Genetic manipulation of some of these herds to increase body and antler size is common in New Zealand, Argentina and Mexico, including the import of stags from larger subspecies or wapiti, or imported semen or embryos of such animals Pearse and Goosen, The third productive use of red deer is as farmed animals for venison or velvet production Gilbey and Perezgonzalez, Currently about deer are held with an annual export of about 14 tonnes of venison Stats NZ, Red deer are also farmed in other countries outside their native range Australia, Canada and the USA , as well as inside it.
The genus Cervus forms a ring of sub-species and species around the Northern Hemisphere. Deer at the western end of this ring red deer, C. Generally, there is an increase in body size and lighter pelage across the genus from west to east. Wapiti are typically the largest species in the genus. Both sexes have a lighter coloured body and larger cream-coloured rump patch but with dark head, neck and legs than red and sika deer. Wapiti calves are brown with a dark dorsal strip and may have a pattern of indistinct light spots for the first few months — red deer fawns also have white spots with a distinct double row along their back that are more distinct than those in wapiti.
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There are differences in antler conformation between the three species, with wapiti sur-royals the top tines in adults sweeping backward, while those in red deer stay in the same plane as the lower tines, as do those for sika deer. See King for more detailed descriptions of the three species. Their management in New Zealand, Australia, Argentina and Chile is confounded by their mixed status as farmed assets, game animals and biodiversity pests e. Nugent and Fraser, It is possible that introduced C. Most countries either prohibit all legal introductions of exotic species unless they pass a risk analysis, or maintain a black list of banned species.
Established populations of C. They can swim, so the current ongoing attempt to remove them from Resolution Island 20 ha in Fiordland National Park in New Zealand, and the completed removal programme on Secretary Island ha Crouchley et al. See also the case study below.
Current distributions of free-living C. Preventing further spread is not a trivial option as effective management depends on the way the deer are likely to spread — by natural dispersal, by deliberate introduction, or by escape from farms. In most cases all three mechanisms are involved. Natural dispersal is the most difficult mechanism to manage, as seen in the spread of the species across the border between Argentina and Chile Nugent et al.
Although illegal, deliberate liberations appear to be a major cause of spread in Australia Moriarty, and to date has not been effectively managed. Escape from deer farms is inevitable, so management either requires a zonation system where deer farming can be prohibited or an early detection-rapid response approach. A study of deer farmed in areas of New Zealand where no wild deer were present showed that over seven years 27 escape events were reported for 58 farms.
Inadequate fences, human error and storm damage to fences each accounted for a third of escape events, suggesting a reactive approach to inevitable events is required. The project has a telephone hotline for deer farmers to use to call for assistance when they have an escape. Being more proactive e. The principles of sustained control are that the target population must be reduced enough to mitigate the impacts on some asset they adversely affect Choquenot and Parkes, Most introduced deer populations are harvested by recreational hunters who see the animals as a resource and asset.
A question faced by land managers who see the deer as pests is whether this harvest is sufficient by itself or in combination with other tools to act as a control on the deer numbers. They may be legally hunted with no restrictions on the numbers, age or sex by anyone with a firearms licence who has the permission of the land owner. It generally views the animals as pests and where it does not itself control deer it encourages recreational and commercial hunters to kill deer.
Nugent and Choquenot explored the relative cost-effectiveness of commercial harvesting, recreational hunting and state-funded culling as red deer control tools in New Zealand, where recreational hunting of deer for trophies or for meat is a popular activity with about 31 hunters taking about 41 C. They concluded that payment of incentives to commercial aerial hunters would usually be more cost-effective than state-funded culling, at least in non-forested habitats, but payment to recreational hunters would not be effective.
In general, even when there are no restrictions on how many deer can be taken by recreational hunters as in New Zealand it appears that the harvest is not large enough to reduce deer densities by enough to protect more sensitive biodiversity assets Nugent and Fraser, In one area of New Zealand occupied by wapiti, C. Commercial harvesting of C. The important factors are competitive access, the presence or absence of refugia for the deer e.
Commercial exploitation of C. This ceased when a market for venison was developed in the late s. In the late s this harvest was largely taken by hunters shooting from helicopters.
Currently the economics of the industry means about 17 deer are shot each year Warburton et al. The final strategy to sustain control of deer is via official government culling, again only used to any extent in New Zealand. Large-scale culling of deer was conducted by government agencies first across all herds, and then in priority water catchments Caughley Since the advent of the commercial harvest, government culling is limited to deer-free areas to manage escapees Fraser et al. These projects account for a few hundred deer each year.
In the mids the growing farmed deer industry in New Zealand led to large numbers of wild C. In Australia, the status of C.
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They are classed as pests in South Australia, the Australian Capital Territory and Queensland as well as the States in which they do not occur , but as game that can be managed as pests if justified in New South Wales, and simultaneously as pests, protected wildlife and game in Victoria Forsyth et al. There is some limited government control of red deer in National Parks in some states Natural Resource Council, Most States allow recreational hunting under permit i.
In Argentina, the policy to manage C. This appears to have failed, as the range of deer expanded by about a third between and The hunters focus on adult stags as trophies so are unlikely to be an effective control tool by themselves Nugent et al. In Chile, recreational hunting is not allowed on public lands and that on private land is mostly paid trophy hunting on ranches Nugent et al.
This section consists of three case studies from New Zealand. A more detailed account of deer control in two regions of New Zealand is provided by Fraser et al. The development of farming or ranching of C. Where these farms are developed in areas without wild populations there is an obvious risk that new populations can establish if animals escape.
In some areas of New Zealand there are no wild C. A government-funded programme aims to detect and manage any deer that escape from these farms; the results were assessed between and Fraser et al. Results of a survey of deer farm escapes from Northland and Taranaki regions, — after Parkes, Butterflies of the British Isles. In Pursuit of Butterflies. Outdoor Sports In Scotland: Me Father was the Keeper: John Smeaton and the Eddystone Light. Nooks and Corners of Cornwall. Old Glass and How to Collect it. The Bronze Age in Ireland.
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