Migration is the regular seasonal movement of animals from one place to another, often from a breeding site to a nonbreeding site and back. Because the act of migration makes it difficult to follow individuals and populations year round, our understanding of the ecology and evolution of migrating organisms, particularly birds, has been severely impeded. Exciting new advances in satellite telemetry, genetic analyses and stable isotope chemistry are now making it possible to determine the population and geographical origin of individual birds. Here, we review these new approaches and consider the relevance of understanding migratory connectivity to ecological, evolutionary and conservation issues.

Links between worlds: unraveling migratory connectivity

There is growing evidence that inbreeding can negatively affect small, isolated populations. This contrasts with the perception in New Zealand, where it has been claimed that native birds are less affected by inbreeding depression than threatened species from continental regions. It has been argued that New Zealand's terrestrial birds have had a long history of small population size with frequent inbreeding and that this has "purged" deleterious alleles. The rapid recovery of many tiny and inbred populations after introduced predators have been controlled, and without input from more genetically diverse populations, has further supported the view that inbreeding is not a problem. This has led to a general neglect of inbreeding as a factor in recovery programs for highly endangered species such as the Black Robin ( Petroica traversi) and Kakapo ( Strigops habroptilis). We examined the reasons for this situation and review the New Zealand evidence for genetic purging. Complete purging of the genetic load and elimination of inbreeding depression are unlikely to occur in natural populations, although partial purging may be more likely where small populations have become inbred over an extended period of time, such as on small isolated islands. Recent molecular data are consistent with the view that island endemics, including New Zealand's threatened birds, have low genetic variation and hence have possibly gone through longer periods of inbreeding than threatened species from continental regions. Nevertheless, results from recent field studies in New Zealand indicate that, despite the opportunity for purging, inbreeding depression is evident in many threatened species. Although inbreeding depression has not prevented some populations from recovering from severe bottlenecks, the long-term consequences of inbreeding and small population size—the loss of genetic variation—are potentially much more insidious. The degrees to which genetic factors reduce population viability generally remain unquantified in New Zealand. Although minimizing ecological risks (e.g., preventing reinvasion of islands by mammalian predators) will continue to receive high priority in New Zealand because of their much larger impacts, we advocate that genetic considerations be better integrated into recovery plans.

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Inbreeding and endangered species management: is New Zealand out of step with the rest of the world?

To ask whether natural selection occurs at the level of the community is to consider whether communities represent a major transition in evolution – can particular community configurations evolve and maintain their integrity in the face of disruption arising from the self-interest of component individuals? This requires heritable variation among subcommunities in a landscape, and that alternative subcommunities maintain a degree of individuality in both time and space. Recently developed models show that spatial self-structuring in multispecies systems can meet both criteria and provide a rich substrate for community-level selection and a major transition in evolution.

Selection at the level of the community: the importance of spatial structure

Microsatellite as well as sequence analysis of the mitochondrial control region were applied to infer phylogeography and population genetic structure of the saker falcon (Falco cherrug). Furthermore, we compared the patterns of mitochondrial haplotypes with the variation of microsatellite alleles among the species of the hierofalcon complex (F. cherrug, Falco rusticolus, Falco biarmicus, Falco jugger) to test hypotheses on population history. Historical samples from museum specimens of F. cherrug were analysed together with samples from contemporary populations to investigate possible influences of hybrid falcons escaped from falconry on the genetic composition. In the mitochondrial DNA analysis, none of the four species represents a monophyletic group. Moreover, there are no clearly defined groups of haplotypes corresponding to taxonomic entities. In the microsatellite analysis most of the variation is shared between species and no clear differentiation by private alleles is found. Yet, with a Bayesian clustering method based on allele frequencies, a differentiation of F. cherrug, F. rusticolus and two geographic groups of F. biarmicus was detected. Results from both nuclear and mitochondrial markers are compatible with the previously postulated 'Out of Africa' hypothesis assuming an African origin of the hierofalcons. From an ancestral African population, F. cherrug, F. rusticolus and F. jugger split off in separate waves of immigration into Eurasia and South Asia. A combination of evolutionary processes, including incomplete lineage sorting as well as hybridization, may be responsible for the currently observed genetic patterns in hierofalcons.

http://www.globalraptors.org/grin/researchers/uploads/350/nittinger_et_al_2007_molecol.pdf

Phylogeography and population structure of the saker falcon (Falco cherrug) and the influence of hybridization: mitochondrial and microsatellite data.

Peregrine falcon populations underwent devastating declines in the mid-20th century due to the bioaccumulation of organochlorine contaminants, becoming essentially extirpated east of the Great Plains and significantly reduced elsewhere in North America. Extensive re-introduction programs and restrictions on pesticide use in Canada and the United States have returned many populations to predecline sizes. A proper population genetic appraisal of the consequences of this decline requires an appropriate context defined by (i) meaningful demographic entities; and (ii) suitable reference populations. Here we explore the validity of currently recognized subspecies designations using data from the mitochondrial control region and 11 polymorphic microsatellite loci taken from 184 contemporary individuals from across the breeding range, and compare patterns of population genetic structure with historical patterns inferred from 95 museum specimens. Of the three North American subspecies, the west coast marine subspecies Falco peregrinus pealei is well differentiated genetically in both time periods using nuclear loci. In contrast, the partitioning of continental Falco peregrinus anatum and arctic Falco peregrinus tundrius subspecies is not substantiated, as individuals from these subspecies are historically indistinguishable genetically. Bayesian clustering analyses demonstrate that contemporary genetic differentiation between these two subspecies is mainly due to changes within F. p. anatum (specifically the southern F. p. anatum populations). Despite expectations and a variety of tests, no genetic bottleneck signature is found in the identified populations; in fact, many contemporary indices of diversity are higher than historical values. These results are rationalized by the promptness of the recovery and the possible introduction of new genetic material.

Appraisal of the consequences of the DDT-induced bottleneck on the level and geographic distribution of neutral genetic variation in Canadian peregrine falcons, Falco peregrinus.

Genetic variability and population structure in the endangered peregrine falcon (Falco peregrinus) were studied using DNA microsatellite markers. Special emphasis was placed on the subspecies F. p. peregrinus living in Scandinavia and Scotland. For comparative purposes, three North American peregrine subspecies (F. p. tundrius, F. p. pealei and F. p. anatum) and one Tasmanian subspecies (F. p. macropus) were included in this analysis. Twelve DNA microsatellite loci were analysed across a total of 146 individuals. The amount of genetic variation did not differ in peregrine populations for which adequate sample sizes were obtained. Within F. p. peregrinus, significant genetic differentiation was found between populations in northern and southern Scandinavia and between the Scandinavian and Scottish populations, while the populations in south-eastern Norway and south-western Sweden did not differ significantly. Analysis of cliff-nesting peregrines in northern Sweden and bog-nesting peregrines in northern Finland/Sweden suggests that the difference in nesting habitat is not associated with genetic differentiation. Population structuring in F. p. peregrinus is further supported by an assignment test, wherein simulated genotypes are correctly assigned to the northern and southern Scandinavian and Scottish populations with relatively high probabilities, and by analysis of allele-sharing among individuals. Cluster analysis of genetic distances grouped populations of peregrines in accordance with their subspecific designation, and peregrines on either side of the Atlantic Ocean were clearly separated.

D. A. Bell

Papers

Microsatellite analysis of population structure and genetic variability in peregrine falcons (Falco peregrinus)