Can a ‘wintering area effect’ explain population status of Swainson's hawks? A stable isotope approach

It has been suggested that declines in breeding populations of Swainson's hawks ( Buteo swainsoni ) in California, Oregon, and Nevada may be due to differential mortality of hawks on their wintering grounds. Although massive mortality incidents reported on the wintering grounds partially support this suggestion, there are no data showing differential use of wintering areas by breeding populations of Swainson's hawks. We used stable-hydrogen isotope analysis of feathers to determine whether large flocks of hawks wintering in Argentina consisted of a mixture of individuals from across the North American breeding range or consisted of individuals from discrete breeding populations. We found that flocks of wintering Swainson's hawks consisted of a mixture of individuals. The lack of connectivity between populations of breeding and wintering hawks suggests that high wintering mortality, either natural or human-induced, is unlikely to have direct consequences on a single breeding area in North America. The demographic effects of winter mortality should be 'diluted' across the entire breeding range of Swainson's hawks.     

Geese and their future fortune

Wild goose populations were probably limited by winter food supplies before Man modified their winter range and effectively removed that limitation on population growth. At the same time, reduction of hunting lowered Man-induced mortality and allowed most populations to increase both in Europe and in North America.
This paper considers likely future trends, and, using the population of Barnacle Geese breeding in Svalbard as an example, suggests how numbers might be regulated in a density related manner. The Svalbard Barnacle Goose population has shown a decline in recruitment rate and an increase in the natural mortality rate as numbers have increased. Both of these are reflections of competition on the breeding grounds, though there may be some effects at other times. We suggest that most populations will eventually be limited by resources on the breeding area.
Some of the implications of changes in numbers are that pair-bond stability and breeding area philopatry are less advantageous and in time will become less pronounced as the selective value of these traits are diminished.
We argue that both for applied reasons (conservation, crop damage prediction) and for academic investigations of population processes, long-term studies are vital in the management and understanding of goose populations. A number of features of their life history make geese very suitable subjects for such studies.     

Biogeographic models of gene flow in two waterfowl of the Australo‐Papuan tropics

There are many large, easy‐to‐observe anseriform birds (ducks, geese, and swans) in northern Australia and New Guinea and they often gather in large numbers. Yet, the structure of their populations and their regional movements are poorly understood. Lack of understanding of population structure limits our capacity to understand source‐sink dynamics relevant to their conservation or assess risks associated with avian‐borne pathogens, in particular, avian influenza for which waterfowl are the main reservoir species. We set out to assess present‐day genetic connectivity between populations of two widely distributed waterfowl in the Australo‐Papuan tropics, magpie goose Anseranas semipalmata (Latham, 1798) and wandering whistling‐duck Dendrocygna arcuata (Horsfield, 1824). Microsatellite data were obtained from 237 magpie geese and 64 wandering whistling‐duck. Samples were collected across northern Australia, and at one site each in New Guinea and Timor Leste. In the wandering whistling‐duck, genetic diversity was significantly apportioned by region and sampling location. For this species, the best model of population structure was New Guinea as the source population for all other populations. One remarkable result for this species was genetic separation of two flocks sampled contemporaneously on Cape York Peninsula only a few kilometers apart. In contrast, evidence for population structure was much weaker in the magpie goose, and Cape York as the source population provided the best fit to the observed structure. The fine scale genetic structure observed in wandering whistling‐duck and magpie goose is consistent with earlier suggestions that the west‐coast of Cape York Peninsula is a flyway for Australo‐Papuan anseriforms between Australia and New Guinea across Torres Strait.     

Lack of detection of host associated differences in Newcastle disease viruses of genotype VIId isolated from chickens and geese

ABSTRACT: BACKGROUND: The goose is usually considered to be resistant even to strains of Newcastle disease virus (NDV) that are markedly virulent for chickens. However, ND outbreaks have been frequently reported in goose flocks in China since the late 1990s with the concurrent emergence of genotype VIId NDV in chickens. Although the NDVs isolated from both chickens and geese in the past 15 years have been predominantly VIId viruses, published data comparing goose- and chicken-originated ND viruses are scarce and controversial. RESULTS: In this paper, we compared genotype VIId NDVs originated from geese and chickens genetically and pathologically. Ten entire genomic sequences and 329 complete coding sequences of individual genes from genotype VIId NDVs of both goose- and chicken-origin were analyzed. We then randomly selected two goose-originated and two chicken-originated VIId NDVs and compared their pathobiology in both geese and chickens in vivo and in vitro with genotype IV virus Herts/33 as a reference. The results showed that all the VIId NDVs either from geese or from chickens shared high sequence homology and characteristic amino acid substitutions and clustered together in phylogenetic trees. In addition, geese and chickens infected by goose or chicken VIId viruses manifested very similar pathological features distinct from those of birds infected with Herts/33. CONCLUSIONS: There is no genetic or phenotypic difference between genotype VIId NDVs originated from geese and chickens. Therefore, no species-preference exists for either goose or chicken viruses and more attention should be paid to the trans-species transmission of VIId NDVs between geese and chickens for the control and eradication of ND.     

Age bias in the bag of pink-footed geese <Emphasis Type="Italic">Anser brachyrhynchus</Emphasis>: influence of flocking behaviour on vulnerability

In pink-footed goose (Anser brachyrhynchus) wintering in Denmark, The Netherlands and Belgium, the proportion of juveniles in the hunting bag is consistently higher than that observed in the autumn population. Such juvenile bias in the bag is usually ascribed to young geese lacking experience with hunting or disruption of juveniles from families. An alternative explanation may be that flocking behaviour of families make juveniles more vulnerable. Observations of morning flights of pink-footed geese to the feeding grounds from two of the major autumn-staging areas showed that geese were distributed in many small flocks (median flock size = 9). This was not significantly different from the flock size distribution shot at by hunters (median = 8), suggesting that hunters targeted goose flock size in proportion to the general probability of encounter. The rate at which hunters downed geese was independent of flock size. The ratio between juveniles and adults in flocks decreased with flock size and flocks of <60 individuals primarily comprised family groups. The likelihood of being shot at was 2.4 times higher for juveniles and 3.4 times higher for older birds in small flocks (<10 individuals) compared to larger flocks. The observations suggest that both juveniles as well as successful adult breeding birds were more vulnerable than non-breeding/failed breeding birds as a result of flocking behaviour.     

Relationship between wild greylag and European domestic geese based on mitochondrial DNA

The origins of the European domestic goose are uncertain. The available information comes from archaeological findings and historical literature, but genetic evidence has hitherto been scarce. The domestic goose in Europe is derived from the greylag goose (Anser anser), but it is not known where the initial domestication took place and which of the two subspecies of greylag goose was ancestral. We aimed to determine the amount and geographical distribution of genetic diversity in modern populations of greylag geese as well as in different breeds of the domestic goose to make inferences about goose domestication. We studied DNA sequence variation in the mitochondrial control region of greylag geese from multiple populations across Europe and western Asia as well as specimens of domestic geese representing 18 modern breeds and individuals not belonging to any recognised breed. Our results show notable differences in genetic diversity between different greylag goose populations and the presence of six mitochondrial haplogroups which show a degree of geographical partitioning. The genetic diversity of the domestic goose is low, with 84% of sampled individuals having one of two major closely related haplotypes, suggesting that modern European domestic geese may derive from a narrow genetic base. The site of domestication remains unresolved, but domestic geese in Turkey were unusually diverse, indicating the importance of further sampling in the vicinity of the eastern Mediterranean and the Near East. There appears to be past or ongoing hybridisation between greylags and domestic geese in particular areas, consistent with field observations.     

Of fleas and geese: the impact of an increasing nest ectoparasite on reproductive success

Past studies on the relationship between nest ectoparasites and avian fitness have been primarily limited to altricial hosts. Life history strategies of precocial and altricial birds vary considerably, limiting our ability to infer the effect of nest parasites on fitness of precocial species. Ross's Chen rossii and lesser snow goose Chen caerulescens caerulescens populations have been growing at unprecedented high rates. New limiting factors on vital rates of these precocial birds may arise after populations have been released from previously regulating factors. The flea Ceratophyllus vagabundus vagabundus is an apparently newly emerging nest parasite in the arctic goose colony at Karrak Lake, Nunavut, Canada. We examined the relationship between flea abundance (measured by the proportion of goose eggs covered by blood in each nest) and goose reproductive success from 2001–2004. In three of four years of study, nest success was inversely related to flea abundance in nests. Despite the potential for high costs to individuals, the overall effects of fleas on goose nesting success have thus far been small. We demonstrated that nest parasites negatively influence reproductive success of precocial bird hosts despite host life history strategy of leaving the nest quickly after hatch, which results in minimal exposure to nest parasites compared to altricial birds that raise their young in the nest.     

Nicarbazin OvoControl G Bait Reduces Hatchability of Eggs Laid by Resident Canada Geese in Oregon

Abstract: Expanding populations of resident Canada geese (Branta canadensis) are resulting in increased conflicts with humans. Nonlethal and humane means are needed for managing Canada goose flocks at a variety of sites, including golf courses, industrial parks, government sites, and city parks. Decreased egg production and hatching are side effects of nicarbazin, a veterinary drug used to treat coccidiosis in chickens. Capitalizing on these effects, we developed nicarbazin as a reproductive inhibitor for Canada geese and conducted a field efficacy study. We recruited study sites in 2002 and 2003. Following laboratory testing, we conducted a field efficacy trial of nicarbazin for reducing the hatchability of Canada goose eggs in spring 2004 in Oregon, USA. The study began in February 2004 at 10 sites in Oregon, with 2 control and 3 treated sites on each side of the Cascades. We fed bait daily to resident Canada geese for approximately 6 weeks. We located and monitored nests until hatching or ≥5 days beyond the expected hatching date to determine hatchability. We completed data collection in May 2004. Geese consumed 8,000 kg of bait, with 5,100 kg of OvoControl G® (Innolytics, LLC, Rancho Santa Fe, CA) 2,500-ppm nicarbazin bait consumed among 6 treated sites and 2,900 kg of untreated bait consumed among 4 control sites. We monitored 63 nests at treated sites and 46 nests at control sites to determine hatching success of eggs. There was a 62% reduction in the percentage of nests with 100% hatchability at treated sites as compared to controls. There was a 93% increase in the percentage of nests at treated sites with 0% hatchability as compared to nests with no eggs hatching at control sites. Hatchability from treated sites versus control sites was reduced 36% (F = 5.72, P = 0.0622). We submitted results from this study to support Environmental Protection Agency registration of nicarbazin as a reproductive inhibitor for use in Canada geese. We have shown that treatment of resident Canada geese with OvoControl G 2,500-ppm nicarbazin bait by licensed, trained applicators immediately prior to and during the breeding season can reduce hatchability of eggs laid by treated geese, thereby reducing recruitment of goslings into problem resident Canada goose populations.     

Flyway structure in the circumpolar greater white‐fronted goose

Dispersal and migratory behavior are influential factors in determining how genetic diversity is distributed across the landscape. In migratory species, genetic structure can be promoted via several mechanisms including fidelity to distinct migratory routes. Particularly within North America, waterfowl management units have been delineated according to distinct longitudinal migratory flyways supported by banding data and other direct evidence. The greater white‐fronted goose (Anser albifrons) is a migratory waterfowl species with a largely circumpolar distribution consisting of up to six subspecies roughly corresponding to phenotypic variation. We examined the rangewide population genetic structure of greater white‐fronted geese using mtDNA control region sequence data and microsatellite loci from 23 locales across North America and Eurasia. We found significant differentiation in mtDNA between sampling locales with flyway delineation explaining a significant portion of the observed genetic variation (~12%). This is concordant with band recovery data which shows little interflyway or intercontinental movements. However, microsatellite loci revealed little genetic structure suggesting a panmictic population across most of the Arctic. As with many high‐latitude species, Beringia appears to have played a role in the diversification of this species. A common Beringian origin of North America and Asian populations and a recent divergence could at least partly explain the general lack of structure at nuclear markers. Further, our results do not provide strong support for the various taxonomic proposals for this species except for supporting the distinctness of two isolated breeding populations within Cook Inlet, Alaska (A. a. elgasi) and Greenland (A. a. flavirostris), consistent with their subspecies status.     

Monitoring exposure to avian influenza viruses in wild mammals

• 1 Avian influenza (AI) viruses primarily circulate in wild waterfowl populations and are occasionally transmitted to domestic poultry flocks. However, the possible roles of other wildlife species, such as wild mammals, in AI virus ecology have not been adequately addressed.
• 2 Due to their habitat and behaviour, many wild mammals may be capable of transmitting pathogens among wild and domestic populations. Exposure to AI viruses has been reported in an array of wild and domestic animals. The presence of wild mammals on farms has been identified as a risk factor for at least one poultry AI outbreak in North America. These reports suggest the need for seroprevalence studies examining the exposure of wild mammals to AI viruses.
• 3 Serological tests are routinely used to assess domestic poultry, domestic swine and human exposure to influenza A viruses, but these tests have not been validated for use in wild mammals. As such, some of these protocols may require adjustments or may be inappropriate for use in serology testing of wild mammals. Herein, we review these serological techniques and evaluate their potential usefulness in AI surveillance of wild mammals. We call for care to be taken when applying serological tests outside their original area of validation, and for continued assay verification for multiple species and virus strains.

     

Seagrass (Zostera spp.) as food for brent geese (Branta bernicla): an overview

Brent geese (called brant in North America) are among the smallest and the most marine of all goose species, and they have very long migration routes between high Arctic breeding grounds and temperate wintering grounds. Like all other geese, brent geese are almost entirely herbivorous. Because of these ecological characteristics they have a high food demand and are strongly dependent on stopover sites to ”refuel” during the migration period. Three subspecies of brent geese are distributed around the Holarctic, forming seven populations with distinct migration routes. Most or all of these populations make heavy use of Zostera spp. during migratory stopovers on spring and/or autumn migration. Examples of Zostera stopover areas being used by large numbers of brent geese for several weeks each year are Izembek Lagoon (Alaska), lagoons in Baja California, the German/Danish Wadden Sea, the Golfe du Morbihan (France), British estuaries, and the White Sea (Western Russian Arctic). Brent geese feed on Zostera wherever they can, but they can only reach the plants at low tide or in shallow water. Changes in Zostera abundance affect brent goose distribution, and the ”wasting disease” affecting Atlantic Zostera stocks during the 1930s was at least partly responsible for a steep decline in brent goose population sizes on both sides of the Atlantic. While Zostera is of outstanding importance as food for brent geese, the impact of the geese on Zostera stocks seems to be less important – at many sites, the geese consume only a small amount of the available Zostera, or, if they consume more, the seagrass can regenerate fully until the following season. Received: 6 December 1998 / Received in revised form: 6 August 1999 / Accepted: 9 August 1999     

西藏南部羊卓雍错水鸟群落及斑头雁活动区域特征

2009年4月至2010年1月,对西藏南部羊卓雍错的水鸟资源状况进行了调查。采用定点观察的方法,沿湖选择了24个观察点,分别在繁殖前期、中期和后期,以及秋季和冬季进行了6次调查。采用核密度分析(Kernel analysis)的方法,对两只卫星跟踪斑头雁(Anser indicus)的活动区进行了分析。调查期间,记录到水鸟32种31044只,隶属于6目10科。雁鸭类和鸥类分别占水鸟总数73.9%和19.1%,主要是斑头雁、赤嘴潜鸭(Rhodonessa rufina)、赤麻鸭(Tadorna ferruginea)、棕头鸥(Larus brunnicephalus)等。水鸟多样性较高的季节是春秋迁徙季节。羊卓雍错夏季主要的繁殖种群是斑头雁和棕头鸥,也有少量黑颈鹤(Grus nigricollis)的繁殖个体;冬季主要物种是赤嘴潜鸭,经常聚集在融化的冰面上。春季斑头雁的数量增加趋势较为明显;进入繁殖期后,斑头雁处于孵卵阶段,繁殖种群的数量达到2000余只;繁殖后期,斑头雁换羽结束,成鸟带领幼鸟在鸟岛附近的湖边取食,此时观察到斑头雁的数量又有明显的增加;秋季斑头雁的南迁致使种群数量呈下降趋势;冬季许多斑头雁从北方如青海湖等地迁来越冬使得种群数量有所增加,多分布于湖西浪卡子县城附近的沼泽湿地和湖南部的绒波臧布河流的入口处。卫星跟踪结果表明,羊卓雍错是青海湖繁殖的斑头雁重要的越冬地之一,湖西部沼泽湿地和湖南部的河流入口处是其主要活动区域,而且该湖与雅鲁藏布江河谷之间通过斑头雁的往来移动存在着联系,因而是西藏南部禽流感监测的重要地点。     

RAPD-PCR analysis of various goose populations

The aim of this study was to genetically analyse by the RAPD-PCR method four indigenous Polish goose breeds, Kartuska (Ka), Lubelska (Lu), Kielecka (Ki) and Podkarpacka (Pd), in order to determine the band-sharing frequency as well as bands characteristic of the evaluated breeds. The birds were maintained as conservative flocks, accounting for a reserve of genetic resources. A total of 102 scorable bands were obtained, their number ranging from 0 to 8, depending on one of seven primers used and the group of birds analyzed, within a mean of 3.64. For each genetic group specific bands with given primers were obtained, suggesting their potential for use as population-specific markers, especially in ex-situ conservation methods. The results also suggest that keeping endangered geese as separate flocks is relevant for their preservation.     

Benzimidazole resistance of sheep nematodes in Norway confirmed through controlled efficacy test

ABSTRACT: BACKGROUND: Resistance against benzimidazoles (BZ) has recently been detected in Norwegian sheep flocks through a large scale prevalence survey based on the faecal egg count reduction test (FECRT). The use of this test in combination with bulk larval culture only gives an indication of which gastrointestinal nematodes genera that are involved and these results have to be confirmed by a controlled efficacy test (CET) to get accurate information about resistant nematodes populations at species level. A CET was therefore performed with larvae from two flocks where BZ resistance was previously detected through FECRT. RESULTS: The latter test confirmed the previous results in both flocks. In flock A, the BZ resistant nematode population consisted solely of Haemonchus contortus, whereas H. contortus and Teladorsagia circumcincta comprised the resistant worm population in flock B. CONCLUSIONS: Some discrepancies that have been recorded between FECRT and CET results regarding time for post-treatment coproscopical examination and a temporary suppression of faecal egg excretion are discussed.     

Tug of war between continental gene flow and rearing site philopatry in a migratory bird: the sex-biased dispersal paradigm reconsidered

Nonrandom dispersal has been recently advanced as a mechanism promoting fine-scale genetic differentiation in resident populations, yet how this applies to species with high rates of dispersal is still unclear. Using a migratory species considered a classical example of male-biased dispersal (the greater snow goose, Chen caerulescens atlantica ), we documented a temporally stable fine-scale genetic clustering between spatially distinct rearing sites (5–30 km apart), where family aggregates shortly after hatching. Such genetic differentiation can only arise if, in both sexes, dispersal is restricted and nonrandom, a surprising result considering that pairing occurs among mixed flocks of birds more than 3000 km away from the breeding grounds. Fine-scale genetic structure may thus occur even in migratory species with high gene flow. We further show that looking for genetic structure based on nesting sites only may be misleading. Genetically distinct individuals that segregated into different rearing sites were in fact spatially mixed during nesting. These findings provide new, scale-dependent links between genetic structure, pairing, and dispersal and show the importance of sampling different stages of the breeding cycle in order to detect a spatial genetic structure.     

Genetic population structure indicates sympatric speciation of Lake Malawi pelagic cichlids.

Allopatric processes of speciation have routinely been presented to explain the extraordinary radiation of the East African Great Lakes cichlid fish species flocks. The 21 or more species of pelagic cichlids within the Lake Malawi flock appear to have lake-wide distributions that challenge such a concept. Data from six microsatellite DNA loci indicate single, panmictic populations across the lake of three Diplotaxodon species. Levels of variability at these loci suggest that populations have been large and stable. Mitochondrial DNA sequence data (872 bp of control region + 981 bp of the NADH-2) from 90 species, representing all major clades within the Lake Malawi flock, indicate reciprocal monophyly of the pelagic clade. We suggest that these data support a hypothesis that speciation in sympatry is more plausible (and widespread) within the cichlid species flocks than previously thought.     

Genetic structure among greater white‐fronted goose populations of the Pacific Flyway

An understanding of the genetic structure of populations in the wild is essential for long‐term conservation and stewardship in the face of environmental change. Knowledge of the present‐day distribution of genetic lineages (phylogeography) of a species is especially important for organisms that are exploited or utilize habitats that may be jeopardized by human intervention, including climate change. Here, we describe mitochondrial (mtDNA) and nuclear genetic (microsatellite) diversity among three populations of a migratory bird, the greater white‐fronted goose (Anser albifrons), which breeds discontinuously in western and southwestern Alaska and winters in the Pacific Flyway of North America. Significant genetic structure was evident at both marker types. All three populations were differentiated for mtDNA, whereas microsatellite analysis only differentiated geese from the Cook Inlet Basin. In sexual reproducing species, nonrandom mate selection, when occurring in concert with fine‐scale resource partitioning, can lead to phenotypic and genetic divergence as we observed in our study. If mate selection does not occur at the time of reproduction, which is not uncommon in long‐lived organisms, then mechanisms influencing the true availability of potential mates may be obscured, and the degree of genetic and phenotypic diversity may appear incongruous with presumed patterns of gene flow. Previous investigations revealed population‐specific behavioral, temporal, and spatial mechanisms that likely influence the amount of gene flow measured among greater white‐fronted goose populations. The degree of observed genetic structuring aligns well with our current understanding of population differences pertaining to seasonal movements, social structure, pairing behavior, and resource partitioning.     

Goose hybrids, captive breeding and restocking of the Fennoscandian populations of the Lesser White-fronted goose (Anser erythropus)

The lesser white-fronted goose (Anser erythropus) isthe most threatened of the Palearctic goose species with a decliningpopulation trend throughout its distributional range. The currentestimate of the Fennoscandian subpopulation size is 30–50 breedingpairs, whereas it still numbered more than 10000 individuals at thebeginning of the last century. Reintroduction and restocking have beencarried out in Sweden and Finland using captive lesser white-frontedgoose stock with unknown origins. We have carried out a study of thegenetic composition of captive-bred stock by sequencing a 221 bphypervariable fragment of the mitochondrial DNA (mtDNA) control regionfrom 15 individuals from the Hailuoto farm, Finland. Two out of thethree maternal lineages detected in the captive stock are also presentin wild populations. The third maternal lineage among the captive lesserwhite-fronted geese originates from the closely related greaterwhite-fronted goose (Anser albifrons). None of the investigatedwild lesser white-fronted goose individuals carried the mtDNA of thegreater white-fronted goose. The presence of greater white-fronted goosemtDNA in the lesser white-fronted goose captive stock suggests thathybridization has occurred during captive propagation.     

Winter Fidelity and Apparent Survival of Lesser Snow Goose Populations in the Pacific Flyway

Abstract The Beringia region of the Arctic contains 2 colonies of lesser snow geese (Chen caerulescens caerulescens) breeding on Wrangel Island, Russia, and Banks Island, Canada, and wintering in North America. The Wrangel Island population is composed of 2 subpopulations from a sympatric breeding colony but separate wintering areas, whereas the Banks Island population shares a sympatric wintering area in California, USA, with one of the Wrangel Island subpopulations. The Wrangel Island colony represents the last major snow goose population in Russia and has fluctuated considerably since 1970, whereas the Banks Island population has more than doubled. The reasons for these changes are unclear, but hypotheses include independent population demographics (survival and recruitment) and immigration and emigration among breeding or wintering populations. These demographic and movement patterns have important ecological and management implications for understanding goose population structure, harvest of admixed populations, and gene flow among populations with separate breeding or wintering areas. From 1993 to 1996, we neckbanded molting birds at their breeding colonies and resighted birds on the wintering grounds. We used multistate mark-recapture models to evaluate apparent survival rates, resighting rates, winter fidelity, and potential exchange among these populations. We also compared the utility of face stain in Wrangel Island breeding geese as a predictor of their wintering area. Our results showed similar apparent survival rates between subpopulations of Wrangel Island snow geese and lower apparent survival, but higher emigration, for the Banks Island birds. Males had lower apparent survival than females, most likely due to differences in neckband loss. Transition between wintering areas was low (<3%), with equal movement between northern and southern wintering areas for Wrangel Island birds and little evidence of exchange between the Banks and northern Wrangel Island populations. Face staining was an unreliable indicator of wintering area. Our findings suggest that northern and southern Wrangel Island subpopulations should be considered a metapopulation in better understanding and managing Pacific Flyway lesser snow geese. Yet the absence of a strong population connection between Banks Island and Wrangel Island geese suggests that these breeding colonies can be managed as separate but overlapping populations. Additionally, winter population fidelity may be more important in lesser snow geese than in other species, and both breeding and wintering areas are important components of population management for sympatric wintering populations.