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1.
Abstract.
  • 1 Wing form frequencies in 255 populations of 101 species of leafhoppers (Auchenorrhyncha) in temporary and permanent habitats were documented.
  • 2 The proportion of brachypterous specimens in the leafhopper assemblages on ruderal host plants in temporary habitats (median 2%, range 0–8%) was significantly lower than that on permanent host plants in undisturbed habitats (median 22%, range 0–94%).
  • 3 Leafhopper species typical of temporary habitats were either monomorphic, macropterous, or wing-dimorphic with macropterous forms prevailing in both sexes.
  • 4 Among the eighty-nine species recorded in permanent habitats, forty-five species were wing dimorphic. In forty-one dimorphic species, a brachypterous form prevailed. This prevalence was found for both sexes in thirty-one species, for only females in nine species and for only males in one species.
  • 5 The prevalence of brachypters in males, but not in females, found in Anoscopus flavostriatus, is probably the first such documented case in Auchenorrhyncha.
  • 6 The hypothesis is proposed that in temporary habitats, density-dependent production of macropters in wing dimorphic species is an adaptation to frequent habitat deterioration caused by factors independent of the density of the species.
  • 7 The predominance of brachypters in permanent habitats indicates that a density-dependent decrease in fitness usually does not offset the potential decrease in fitness connected with macroptery and dispersal. Because of this inability of leafhopper populations to decrease significantly the quality of their resources, a high population density cannot be used as a predictor of future quality of these resources, which is information essential for efficient dispersal behaviour.
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2.
Summary Habitat change by means of flight activities has been observed in the life cycle of the univoltine, macropterous species Amara plebeja Gyll. The hibernation- and reproduction habitats are quite different, viz. deciduous trees and grass vegetation, respectively. Young as well as old females and males are present in both habitats, but at different periods of the year. The flight muscles are generally autolysed but apparently only temporarily. After complete reconstruction of the flight muscles, habitats are changed by mean of flying both in spring and in autumn, if weather conditions are favourable. In spring, when the individuals are flying from hibernation to reproduction habitat they may be drifted by air currents, whereby migration becomes dispersal, and founding or refounding of populations in suitable localities may result. During the autumn, the insects fly from reproduction to hibernation habitats orientating towards distinct silhouettes in the landscape.Communication No. 185 of the Biological Station, Wijster  相似文献   

3.
Abstract. 1. A series of laboratory experiments was conducted to explore the effects of inter‐specific interactions, both direct interactive effects and those induced through previous feeding, on the dispersal capability (proportion of macropterous adults) and performance (development time and survival) of two wing‐dimorphic planthoppers, the whitebacked planthopper Sogatella furcifera and the brown planthopper Nilaparvata lugens, two pests of rice throughout Asia. 2. An asymmetric effect of inter‐specific crowding on dispersal capability was detected between the two planthoppers. With density controlled, the proportion of macropterous adults in N. lugens was higher when raised in mixed populations with S. furcifera than in pure cultures comprised of conspecifics, suggesting that interspecific effects on dispersal capability are stronger than intraspecific effects. In contrast, interspecific effects on macroptery (%) were weaker than intraspecific impacts for S. furcifera. 3. This trend was parallelled by adverse competitive effects on survival, with interspecific effects stronger than intraspecific impacts for N. lugens and intraspecific effects stronger for S. furcifera. 4. Crowding also affected the incidence of macroptery indirectly through feeding‐induced changes in plant physiology. Rearing N. lugens on plants fed on previously by heterospecifics resulted in a higher incidence of macroptery in females and protracted development than for N. lugens raised on plants fed on previously by conspecifics. By contrast, the intraspecific effect of previous feeding was stronger than the interspecific effect in S. furcifera, with higher macropter production and prolonged development occurring on plants exposed previously to conspecifics. 5. The results suggest that interspecific interactions between herbivorous insects, both direct and via feeding‐induced changes in plant physiology, can have negative consequences for performance and survival and promote the production of flight‐capable adults that can disperse.  相似文献   

4.
The trade-offs associated with macroptery in males were investigated in a wing-dimorphic planthopper, Nilaparvata lugens (Stål) (Delphacidae, Hemiptera). Developmental time of brachypters was significantly shorter than that of macropters across a wide range of population densities, while no within-population differences between wing morphs in body weight were found. In mating experiments, brachypterous males 12 to 24 h old were almost three times more successful in competition for females than macropterous males of the same age, while in five days old males, macropters were twice as successful as brachypters. In competition for females involving direct body contact between males, brachypterous and macropterous males were equally successful. It is suggested that macroptery and delayed peak in sexual activity in males may both be influenced by the titre of juvenile hormone, as is known from females. The combination of prolonged development and delayed maximum competitive ability for mates, associated with macroptery, is likely to be disad vantageous, representing probably a penalty associated with macroptery in males in undisturbed habitats.  相似文献   

5.
ABSTRACT Greater Roadrunners (Geococcyx californianus) are common, poorly studied birds of arid and semi‐arid ecosystems in the southwestern United States. Conservation of this avian predator requires a detailed understanding of their movements and spatial requirements that is currently lacking. From 2006 to 2009, we quantified home‐range and core area sizes and overlap, habitat selection, and survival of roadrunners (N= 14 males and 20 females) in north‐central Texas using radio‐telemetry and fixed kernel estimators. Median home‐range and core‐area sizes were 90.4 ha and 19.2 ha for males and 80.1 ha and 16.7 ha for females, respectively. The size of home range and core areas did not differ significantly by either sex or season. Our home range estimates were twice as large (x?= 108.9 ha) as earlier published estimates based on visual observations (x?= 28–50 ha). Mean percent overlap was 38.4% for home ranges and 13.7% for core areas. Male roadrunners preferred mesquite woodland and mesquite savanna cover types, and avoided the grass‐forb cover type. Female roadrunners preferred mesquite savanna and riparian woodland cover types, and avoided grass‐forb habitat. Kaplan‐Meier annual survival probabilities for females (0.452 ± 0.118[SE]) were twice that estimated for males (0.210 ± 0.108), but this difference was not significant. Mortality rates of male roadrunners were higher than those of females during the spring when males call from elevated perches, court females, and chase competing males. Current land use practices that target woody‐shrub removal to enhance livestock forage production could be detrimental to roadrunner populations by reducing availability of mesquite woodland and mesquite savanna habitat required for nesting and roosting and increasing the amount of grass‐forb habitat that roadrunners avoid.  相似文献   

6.
Abstract Forest clearance results in a marked change in the dispersion of resources such as food and shelter: from a relatively continuous distribution, to patches and strips of remnant habitat set in a more or less inhospitable matrix. Such changes in the patterns of resource dispersion have the potential to strongly influence the dynamics of social and mating systems of resident animal populations. We studied the den‐use patterns, home range characteristics and mating system of a population of bobucks, Trichosurus cunninghami (Marsupialia: Phalangeridae), permanently resident in linear roadside strips of vegetation in north‐eastern Victoria, Australia in order to gain a greater understanding of the impacts of occupying linear roadside habitats on the behavioural ecology of arboreal marsupials. We radio‐tracked 11 adult females and six adult males and collected 2126 diurnal fixes (mean 125/individual) and 1044 nocturnal fixes (mean 61/individual). Males used significantly more den‐trees (mean 16.5 ± 1.5 den‐trees/individual) than females (mean 7.4 ± 0.6 den‐trees/individual) and had home ranges more than twice the size of those of females (male mean 5.1 ± 0.8 ha, female mean 2.1 ± 0.3 ha). On average, each male's home range overlapped with those of three females; there was little intrasexual home range overlap in either sex. Genetic parentage analysis of all young sampled during the study (n = 22) established that only males that overlapped in home range with females had sired those females' young. All but one male in the study sired multiple young in each of the years they were recorded breeding. These behavioural and genetic data indicate that the roadside population was polygynous, in contrast to the socially monogamous bobuck population we studied in a neighbouring forest patch. These differences in behaviour may reflect patterns of resource distribution in the two habitats.  相似文献   

7.
Abstract: We assessed dispersal behavior of 78 radiotagged adult spotted salamanders (Ambystoma maculatum) at a 36-hole golf course in southeastern Connecticut, USA. Lake of Isles Golf Course is atypical of most golf courses in North America because it is much larger (461 ha) than average 18-hole golf courses (54 ha) and deciduous forests accounted for 70% landscape composition on the course. We tracked individuals an average of 164 days as they emigrated from 3 seasonal pools surrounded by golf course fairways and one pool located in contiguous forest (control pool) from March through December 2004. Males and females dispersed similar distances at the control pool, averaging 71 ± 10 m. However, females migrating across the golf course dispersed twice as far (214 ± 25 m) as males on the golf course (102 ± 15 m) and both genders at the control pool. Over 40% the salamanders at the golf course crossed fairways; thus, fairways were not a dispersal barrier to adult spotted salamanders. Previous researchers have suggested establishing a 164-m life zone around breeding ponds to protect pond-breeding amphibian populations. Our results suggest that strategies that protect core upland habitat within 164 m of wetland boundaries would include 82% of adult males and only 50% of adult females. Empirical estimates based on our telemetry study suggest that core terrestrial habitat would have to extend 370 m to protect 95% of adult females, which is much farther than previous estimates.  相似文献   

8.
Wing polymorphism in insects provides a good model system for investigating evolutionary dynamics and population divergence in dispersal‐enhancing traits. This study investigates the contribution of divergent selection, trade‐offs, behaviour and spatial sorting to the evolutionary dynamics of wing polymorphism in the pygmy grasshopper Tetrix subulata (Tetrigidae: Orthoptera). We use data for > 2800 wild‐caught individuals from 13 populations and demonstrate that the incidence of the long‐winged (macropterous) morph is higher and changes faster between years in disturbed habitats characterized by succession than in stable habitats. Common garden and mother‐offspring resemblance studies indicate that variation among populations and families is genetically determined and not influenced to any important degree by developmental plasticity in response to maternal condition, rearing density or individual growth rate. Performance trials show that only the macropterous morph is capable of flight and that propensity to fly differs according to environment. Mark–recapture data reveal no difference in the distance moved between free‐ranging long‐ and short‐winged individuals. There is no consistent difference across populations and years in number of hatchlings produced by long‐ and shorter‐winged females. Our findings suggest that the variable frequency of the long‐winged morph among and within pygmy grasshopper populations may reflect evolutionary modifications driven by spatial sorting due to phenotype‐ and habitat type–dependent emigration and immigration.  相似文献   

9.
ABSTRACT.
  • 1 Gerris remigis Say (Hemiptera; Gerridae) is primarily apterous, but populations with up to 33% macropters have been reported. The macropters seldom fly, and field studies have revealed no detectable differences between wing morphs in movement or survival at any time of year.
  • 2 In this paper, life history traits of macropterous and apterous G. remigis are compared in an attempt to determine the mechanisms responsible for the maintenance of macroptery in this species in spite of the very low flight capacity and infrequent flight of macropters.
  • 3 Development time, proportion breeding without diapause, and overwinter survival do not differ between morphs. However, pre-diapause macropterous females have a significantly shorter pre-oviposition period than apterous females. In contrast, post-diapause macropters begin reproducing later than apters, and have a lower cumulative fecundity.
  • 4 These results suggest that macropters may be at a selective advantage in warm habitats which favour pre-diapause reproduction, but that apters should be favoured in the preferred, cool, lotic habitats.
  • 5 However, crossing and rearing experiments indicate that wing morphology is primarily environmentally controlled in this species, and that the heritability of wing morphology is low, at best. In light of this, the relative impacts of purely phenotypic (environmental) variation, random effects, and the observed fitness differences on the maintenance of macroptery in this species are discussed.
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10.
Aims To test a new concept for island ecology and metapopulation studies: if population persistence is a result of colonization and extinction processes, one would expect remarkable variability in the age of neighbouring populations – there would be both old and young (recently established) populations. Location The lake Mamry archipelago, Poland. Methods Forty trap transects with a total of 114 Barber traps were operating from June to September in 1997 and in 1998. The traps (0.5‐L plastic beakers, mouth diameter 120 mm, with a 20 × 20 cm wooden roof) were placed by applying a stratified sampling design into 13 habitat types on the 16 islands and two mainland sites. The frequency of macropterous individuals in wing‐dimorphic populations of three carabid species (Pterostichus melanarius, P. anthracinus and Carabus granulatus) was used as an indicator of population ‘age’. Results The frequency of macropterous individuals in the populations varied from 0% to 100% for P. melanarius (18.5% on average), from 0% to 91% for P. anthracinus (12.6%) and from 0% to 29% for C. granulatus (8.9%). Populations hosted more long‐winged individuals (and were therefore interpreted as being younger) on smaller islands, compared with those inhabiting large islands and the mainland sites. The results also revealed that the viability of the populations of the autumn‐breeding P. melanarius might be more affected by population size than that of the two studied spring breeders. Island connectivity did not have a significant effect on the frequency of macropterous individuals in the studied populations. Main conclusions The inverse relationship between the proportion of macropterous individuals and island size contrasts with the accepted theory of dispersal, which assumes that there is selection against dispersal on small islands. A regression analysis for population age and habitat characteristics reveals the extinction probability of a given population. I suggest that investigations based on variability in population age can help in studies of colonization–extinction processes that would otherwise face logistic and methodological obstacles.  相似文献   

11.
12.
Wing-dimorphic, delphacid planthoppers were used to test hypotheses concerning the effects of habitat persistence and architectural complexity on the occurrence of dispersal. For reasons concerning both the durational stability of the habitat and the reduced availability of mates, selection has favored high levels of dispersal in species occupying temporary habitats. Flightlessness predominates in species occupying persistent habitats, and is promoted by a phenotypic trade-off between reproductive success and flight capability. Wings are retained in tree-inhabiting species, probably for reasons concerning the more effective negotiation of three-dimensional habitats. In contrast, flightlessness is characteristic of those species inhabiting low profile host plants. For several delphacid genera, migratory species are larger than their sedentary congeners. Because body size and fecundity are positively related in planthoppers, the large body size observed in migratory taxa may result from selection for increased fecundity in colonizing species.  相似文献   

13.
1. In fragmented landscapes many insect species depend on a regular exchange of individuals between subpopulations to ensure the persistence of the population. Thus, the ability to disperse is of particular relevance. 2. However, in some insect species mobility is not a fixed trait. Hence, knowing the causes of phenotypic plasticity is of great importance when evaluating whether a species is able to survive in fragmented landscapes or not. 3. A multi‐year field study was conducted to identify possible causes of macroptery in the wing‐dimorphic habitat specialist Metrioptera brachyptera L. and to quantify its dispersal capability (% macropters). Therefore, 746 individuals of the species were caught on 135 plots. Additionally, environmental variables that possibly induce the development of macropters (population density and habitat moisture) were recorded. 4. Dispersal capability of M. brachyptera was very low. Less than 3% were long‐winged. The statistical analysis revealed that the proportion of long‐winged M. brachyptera was strongly correlated with high bush‐cricket densities and not with habitat moisture. 5. The low dispersal capability of M. brachyptera leads to the conclusion that individual exchange between isolated populations is limited or even impossible. Habitat specialists, like M. brachyptera, may thus be unable to respond to rapid changes in the availability of suitable habitats by dispersing, and hence may be especially dependent on habitat management activities that promote the long‐term stability of existing habitat patches.  相似文献   

14.
In the highly fragmented landscape of central Europe, dispersal is of particular importance as it determines the long‐term survival of animal populations. Dispersal not only secures the recolonization of patches where populations went extinct, it may also rescue small populations and thus prevent local extinction events. As dispersal involves different individual fitness costs, the decision to disperse should not be random but context‐dependent and often will be biased toward a certain group of individuals (e.g., sex‐ and wing morph‐biased dispersal). Although biased dispersal has far‐reaching consequences for animal populations, immediate studies of sex‐ and wing morph‐biased dispersal in orthopterans are very rare. Here, we used a combined approach of morphological and genetic analyses to investigate biased dispersal of Metrioptera bicolor, a wing dimorphic bush‐cricket. Our results clearly show wing morph‐biased dispersal for both sexes of M. bicolor. In addition, we found sex‐biased dispersal for macropterous individuals, but not for micropters. Both, morphological and genetic data, favor macropterous males as dispersal unit of this bush‐cricket species. To get an idea of the flight ability of M. bicolor, we compared our morphological data with that of Locusta migratoria and Schistocerca gregaria, which are very good flyers. Based on our morphological data, we suggest a good flight ability for macropters of M. bicolor, although flying individuals of this species are seldom observed.  相似文献   

15.
In Europe, the consequences of commercial plantation management for birds of conservation concern are poorly understood. The European Nightjar Caprimulgus europaeus is a species of conservation concern across Europe due to population depletion through habitat loss. Pine plantation‐forest is now a key Nightjar nesting habitat, particularly in northwestern Europe, and increased understanding of foraging habitat selection is required. We radiotracked 31 Nightjars in an extensive (185‐km2) complex conifer plantation landscape in 2009 and 2010. Home‐range 95% kernels for females, paired males and unpaired males were an order of magnitude larger than song territories of paired males, emphasizing the importance of habitats beyond the song territory. Nightjars travelled a mean maximum distance of 747 m from the territory centre each night. Home‐range placement relative to landscape composition was examined by compositional analysis. Pre‐closure canopy forest (aged 5–10 years) was selected at all scales (MCP, 95% and 50% kernels), with newly planted forest (aged 0–4 years) also selected within 50% kernels. For telemetry fixes relative to habitat composition within 2 km of their territory centre, individuals again selected pre‐closure and newly planted forest, and also grazed grass heath. Open ungrazed habitat was not selected, with implications for open habitat planning for biodiversity conservation within public‐owned forests. Despite the Nightjars’ selection for younger growth, moth biomass was greater in older forest stands, suggesting that foraging site selection reflects ease of prey capture rather than prey abundance. Within large plantation‐forest landscapes, a variety of growth stages is important for this species and our results suggest that grazing of open habitats within and adjacent to forest will additionally benefit the European Nightjar.  相似文献   

16.
Gene flow through dispersal has traditionally been thought to function as a force opposing evolutionary differentiation. However, directional gene flow may actually reinforce divergence of populations in close proximity. This study documents the phenotypic differentiation over more than two decades in body size (tarsus length) at a very short spatial scale (1.1 km) within a population of pied flycatchers Ficedula hypoleuca inhabiting deciduous and coniferous habitats. Unlike females, males breeding in the deciduous forest were consistently larger than those from the managed coniferous forest. This assortment by size is likely explained by preset habitat preferences leading to dominance of the largest males and exclusion of the smallest ones toward the nonpreferred coniferous forest coupled with directional dispersal. Movements of males between forests were nonrandom with respect to body size and flow rate, which might function to maintain the phenotypic variation in this heritable trait at such a small spatial scale. However, a deeply rooted preference for the deciduous habitat might not be in line with its quality due to the increased levels of breeding density of hole‐nesting competitors therein. These results illustrate how eco‐evolutionary scenarios can develop under directional gene flow over surprisingly small spatial scales. Our findings come on top of recent studies concerning new ways in which dispersal and gene flow can influence microevolution.  相似文献   

17.
Cipo Canasteros (Asthenes luizae, Furnariidae) have a fragmented and limited range restricted to the campos rupestres (rupestrian grasslands) habitat in the Brazilian highlands of the Espinhaço Range, and little is known about their behavior, ecology, and population biology. From March 2009 to November 2010, we monitored birds (24 banded and 22 radio‐tracked) at two study sites at Serra do Cipó in the state of Minas Gerais, Brazil, to estimate their home‐range sizes and population density, and describe their habitat use, natal dispersal behavior, and vocal behavior. We found an average density of 8.7 paired adults/km² in our study areas or 22.9 paired adults/km2 when considering only used habitats. The sex ratio was male‐biased (males/total adults = 0.68), adults exhibited high site fidelity, home‐ranges averaged 4.0 ha (fixed kernel 95%) or 3.5 ha (95% minimum convex polygon) in size, and both sexes defended territories year‐round. We recorded four main types of songs, including two uttered more often during the breeding season. We monitored the natal dispersal of two males and one female who moved maximum distances of 1238 m, 780 m, and 1056 m, respectively, from natal areas. Our results confirm that Cipo Canasteros are restricted to the rocky‐outcrop habitat of the campos rupestres. In part due to their habitat specialization, Cipo Canasteros are considered Near Threatened, but other factors contributing to their demographic fragility include the small number, and probably low survival, of females and low reproductive success due to predation and brood parasitism by cowbirds. Given these threats, along with their specialized habitat and restricted range, the future conservation of Cipo Canasteros will likely depend on the extent to which their campos rupestres habitat can be conserved.  相似文献   

18.
Salinity interacts with many physiological functions and therefore probably influences the distribution of terrestrial fauna in tidal flooded salt marshes. The present study tests the hypothesis that the physiological tolerance of stenotopic wolf spiders for saline conditions at least partially determines their occurrence throughout salt‐marsh and nonsaline habitats. The duration of survival of three stenotopic wolf spider species (Araneae: Lycosidae) with different habitat preferences is compared in a controlled laboratory experiment. The forest‐dwelling Pardosa saltans, the salt‐marsh resident Pardosa purbeckensis and its sister species the inland‐living Pardosa agrestis are exposed to experimental conditions with different levels of salinity. Individuals (45 males and 20–45 females per treatment) are placed in individual air‐tight boxes filled with water‐saturated sand. Three levels of salinity are tested: nonsaline (0‰), medium saline (33–35‰) and highly saline (66–70‰). Contents of carbon, hydrogen and nitrogen and the molar ration carbon/nitrogen remain constant over time and do not differ among salinity treatments, indicating that starvation effects on survival, if any, are similar for all treatments. Conversely, body water significantly decreases over time and differs among salinity treatments, in accordance with patterns of survival. Conforming to their habitat preference, the survival of P. saltans and P. agrestis decreases quickly under highly saline conditions. Pardosa purbeckensis, however, has a high survival under both saline and nonsaline conditions. The duration of survival of females is significantly lower than that of males of P. saltans and P. purbeckensis. Durations of survival of ground‐living wolf spiders exposed to salinity partly match their habitat distribution but do not explain the restriction of salt‐marsh species to saline habitats.  相似文献   

19.
A detailed knowledge of the habitat requirements of steppe birds living in farmland habitats is necessary to identify agricultural practices compatible with their conservation. The globally threatened Great Bustard Otis tarda is a partial migrant in central Iberia, but factors affecting its winter habitat use have not been identified. We assessed habitat differences between breeding and wintering areas and winter habitat selection of radiotagged migrant female Great Bustards in central Spain. Of 68 tagged females, 35% moved to wintering areas located 64.3 ± 24.0 km south of their breeding areas, and 80% wintered in a single area of c. 236 km2. A census of the population in this area identified it as one of the most important wintering areas of this species in the world, holding c. 1500 individuals. There were significant differences between breeding and wintering habitats of individually marked migrant females. Compared with breeding areas, wintering areas of migrant females were located further from roads and urban nuclei, had lower human population densities and area of urban developments, and a higher diversity of land‐use types, with less cover of cereals and more vineyards and olive groves. Within this area, radiotracked migrant females preferred sites with more vineyards and a lower land‐use diversity. Our results highlight the importance of traditional Mediterranean dry farmland mosaics, and suggest that different conservation strategies are needed for migrant and resident populations in winter to secure the conservation of suitable wintering habitat for Great Bustards in the Iberian Peninsula.  相似文献   

20.
Knowledge of survival rates is critical for advancing our understanding of the dynamics of populations and here we report apparent annual survival and breeding dispersal of Scissor‐tailed Flycatchers (Tyrannus forficatus) breeding at two sites in southwest Oklahoma (Ft. Sill and Wichita Mountain Wildlife Refuge [WMWR]). Our Cormack‐Jolly‐Seber estimate of apparent adult survival for the period from 2008 to 2105 was relatively low (0.514) compared to estimates for 36 other migratory and socially monogamous passerines breeding in North America, and was independent of sex (males: = 151; females: = 119), breeding status (territory holder or floater), body mass, site, year, and precipitation during the year prior to breeding. Although apparent survival did not differ between sites, dispersal (= 66 individuals) was more common and breeding dispersal distance (BDD) was greater for Scissor‐tailed Flycatchers at Ft. Sill where anthropogenic disturbance was more frequent. BDD also increased with body mass at Ft. Sill (but not at WMWR) and, after accounting for it, BDD at Ft. Sill tended to be greater for birds that failed to breed successfully in the past year. Older birds and males had the longest BDDs at WMWR, and males exhibited a similar tendency at Ft. Sill. We contend that our estimate of apparent survival is low, not because of inherently low survivorship, but, instead, as a consequence of frequent permanent emigration from our population. We also suggest that the greater BDD of older birds (WMWR) and males (both sites) reflects a history of selection for dispersal in response to frequent habitat disturbance. Frequent habitat disturbance, in addition to the opportunity to prospect for territories both before and after breeding, probably enable the earliest spring arrivals (typically older birds and males) to often relocate between years.  相似文献   

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