Seasonal modulation of flight speed among nocturnal passerine migrants: differences between short- and long-distance migrants

Migrating birds are expected to fly at higher airspeeds when minimizing time rather than energy costs of their migratory journeys. Spring migration has often been suggested to be more time selected than autumn migration, because of the advantage of early arrival at breeding sites. We have earlier demonstrated that nocturnal passerine migrants fly at higher airspeeds during spring compared to autumn, supporting time-selected spring migration. In this study, we test the hypothesis that seasonal airspeeds are modulated differently between short- and long-distance migrants, because of a stronger element of time selection for autumn migration over long distances. In support of this hypothesis, we demonstrate that the seasonal difference in airspeed is significantly larger (spring airspeed exceeding autumn airspeed by a factor of 1.16 after correcting for the influence of altitude, wind and climb/descent on airspeed) among short-distance compared to long-distance (factor 1.12) migrants. This result is based on a large sample of tracking radar data from 3 years at Falsterbo, South Sweden. Short-distance migrants also tend to fly with more favourable winds during autumn, indicating relaxed time constraints (being able to afford to wait for favourable winds) compared to long-distance migrants. These results indicate surprisingly fine-tuned seasonal modulation of airspeed and responses to wind, associated with behavioural strategies adapted to different levels of time selection pressures during spring and autumn migration.     

Evidence of obligate female promiscuity in a socially monogamous passerine

Extrapair paternity in birds has been the focus of an increasing number of studies over the last decades. Nevertheless, there is little knowledge about extrapair copulation (EPC) behavior. In this study, we investigate whether female bluethroats (Luscinia s. svecica) with no extrapair offspring (EPO) in their broods really have been sexually monogamous or if they have copulated with extrapair males. We used an experimental approach to prevent transfer of sperm from the social male during copulation by fitting a rubber tube around the cloaca of randomly chosen males. If the females mated to these males do not participate in EPCs, they will produce infertile eggs, whereas females that copulate with extrapair males will only produce EPO. We found that 87% (n=15) of the experimental pairs compared to only 36% (n=51) of unmanipulated pairs produced EPO. Our result therefore suggests that females having no EPO may still have copulated with extrapair males and that promiscuity may be an obligate sexual strategy among bluethroat females.Communicated by S. Pruett-Jones     

Exaggerated orientation scatter of nocturnal passerine migrants close to breeding grounds: comparisons between seasons and latitudes

Using tracking radars, we investigated the variability of flight directions of long-distance nocturnal passerine migrants across seasons (spring versus autumn migration) and sites at the southern (56° N) and northern (68° N) ends of the Scandinavian Peninsula (Lund versus Abisko). Whilst most migrants at Lund are on passage to and from breeding sites in Fennoscandia, the majority of the migrants at Abisko are close to their breeding sites, and migration at Abisko thus to a large degree reflects initial departure from breeding sites (autumn) or final approach to breeding destinations (spring). The radar data were used to test predictions about differences in orientation and wind drift effects between adult and juvenile birds (a large proportion of autumn migrants consists of juvenile birds on their first journey), between situations far away from or near the goals and between different phases of migration (initial departure, en route passage, final approach to goal). The concentrations (both total and within-night concentrations) of flight directions differed significantly between seasons as well as sites, with the highest concentration at Lund in spring (mean vector length of track directions, r = 0.79) and lowest at Abisko during spring (r = 0.35). Partial wind drift and partial compensation were recorded at Lund, with a similar effect size in spring and autumn, whilst possible wind drift effects at Abisko were obscured by the large directional scatter at this site. The results from Lund support the prediction that the high proportion of juveniles in autumn contributes to increase the directional scatter during this season, whilst there was no support for predictions of differential wind drift effects between seasons and situations with different goal distances. The most striking and surprising result was the exceedingly large scatter of flight directions at Abisko, particularly in spring. We suggest that such an exaggerated scatter may be associated with final approach orientation, where migrants reach their specific goals from all various directions by final navigation within a more wide-ranging goal region. The larger scatter of autumn flight directions at Abisko compared to Lund may be due to exploratory flights in variable directions being more common at initial departure from breeding sites than later during migratory passage. These surprising results highlight the importance of studying and analysing orientation during final approach to (and initial departure from) migratory goals for understanding the orientation systems of migratory birds.     

A multi-scale examination of stopover habitat use by birds

Most of our understanding of habitat use by migrating land birds comes from studies conducted at single, small spatial scales, which may overemphasize the importance of intrinsic habitat factors, such as food availability, in shaping migrant distributions. We believe that a multi-scale approach is essential to assess the influence of factors that control en route habitat use. We determined the relative importance of eight variables, each operating at a habitat-patch, landscape, or regional spatial scale, in explaining the differential use of hardwood forests by Nearctic-Neotropical land birds during migration. We estimated bird densities through transect surveys at sites near the Mississippi coast during spring and autumn migration within landscapes with variable amounts of hardwood forest cover. At a regional scale, migrant density increased with proximity to the coast, which was of moderate importance in explaining bird densities, probably due to constraints imposed on migrants when negotiating the Gulf of Mexico. The amount of hardwood forest cover at a landscape scale was positively correlated with arthropod abundance and had the greatest importance in explaining densities of all migrants, as a group, during spring, and of insectivorous migrants during autumn. Among landscape scales ranging from 500 m to 10 km radius, the densities of migrants were, on average, most strongly and positively related to the amount of hardwood forest cover within a 5 km radius. We suggest that hardwood forest cover at this scale may be an indicator of habitat quality that migrants use as a cue when landing at the end of a migratory flight. At the patch scale, direct measures of arthropod abundance and plant community composition were also important in explaining migrant densities, whereas habitat structure was of little importance. The relative amount of fleshy-fruited trees was positively related and was the most important variable explaining frugivorous migrant density during autumn. Although constraints extrinsic to habitat had a moderate role in explaining migrant distributions, our results are consistent with the view that food availability is the ultimate factor shaping the distributions of birds during stopover.     

Root plasticity buffers competition among plants: theory meets experimental data

Morphological plasticity is a striking characteristic of plants in natural communities. In the context of foraging behavior particularly, root plasticity has been documented for numerous species. Root plasticity is known to mitigate competitive interactions by reducing the overlap of the individuals' rhizospheres. But despite its obvious effect on resource acquisition, plasticity has been generally neglected in previous empirical and theoretical studies estimating interaction intensity among plants. In this study, we developed a semi-mechanistic model that addresses this shortcoming by introducing the idea of compensatory growth into the classical-zone-of influence (ZOI) and field-of-neighborhood (FON) approaches. The model parameters describing the belowground plastic sphere of influence (PSI) were parameterized using data from an accompanying field experiment. Measurements of the uptake of a stable nutrient analogue at distinct distances to the neighboring plants showed that the study species responded plastically to belowground competition by avoiding overlap of individuals' rhizospheres. An unexpected finding was that the sphere of influence of the study species Bromus hordeaceus could be best described by a unimodal function of distance to the plant's center and not with a continuously decreasing function as commonly assumed. We employed the parameterized model to investigate the interplay between plasticity and two other important factors determining the intensity of competitive interactions: overall plant density and the distribution of individuals in space. The simulation results confirm that the reduction of competition intensity due to morphological plasticity strongly depends on the spatial structure of the competitive environment. We advocate the use of semi-mechanistic simulations that explicitly consider morphological plasticity to improve our mechanistic understanding of plant interactions.     

Inter-litter competition and communal nursing among Norway rats: advantages of birth synchrony

Summary The effect of inter-litter competition on pup survival was investigated in pairs of female rats (Rattus norvegicus) living and breeding in the same environment. If a female gave birth when a 0- to 14-day-old litter was already present in the environment, her pups had a very high chance of surviving, similar to the situation in which no other litter was present. Moreover, the mother was likely to nurse communally with the mother of the 0- to 14-day-old litter. This communal nursing benefitted the newborn pups as evidenced by their being heavier at weaning than litters that were not nursed communally. In contrast, if a female gave birth when a 15-to 28-day-old litter was already present in the environment, her newborn pups were likely to die within 3 days postpartum, owing to the fact that they were often prevented from suckling at their mother's teats, resulting in milk deprivation, and were often beneath the older pups, resulting in physical trauma. These findings suggest that inter-litter competition is an important source of pup mortality when litters are born 15-28 days apart. The data are discussed in terms of the advantages of birth synchrony.     

Apparent resource competition among embryos in the brood pouch of a male pipefish

Embryo success was studied in the paternally brooding pipefish Syngnathus typhle. During brooding, which lasts about a month, males provide embryos in their brood pouch with nutrients and oxygen via a placenta-like structure. Egg size depends on female size. In aquaria, males were mated with differently sized females to give the following treatments: M, mixed-egg-size broods of approximately half large and half small eggs; L, single-egg-size broods of large eggs; S, single-egg-size broods of small eggs; and F, field mated males. All males were kept in aquaria for a full brooding period. For each egg-size category, the number of newborn was compared with the number of eggs the male initially fertilized in his brood pouch. Within mixed-egg-size broods, a higher proportion of large eggs survived and large eggs resulted in heavier newborn than small eggs. Indeed, small eggs from a mixed-egg-size brood had significantly lower relative success (proportion of embryos surviving to birth) than those from a brood entirely composed of small eggs. The implication is that embryos compete for resources within the brood pouch, and that competitive success depends on egg size. Given that females produce eggs corresponding in size to their body size, and that females are known to compete indirectly for access to mates (i.e., the sex-roles are reversed), this intrabrood competition could be seen as an extension of female-female competition, but alternative explanations are discussed. Received: 28 April 1995/Accepted after revision: 28 October 1995     

非稳态条件下藻类种间非生物资源竞争理论及研究进展

从理论及实验研究的角度论述了藻类非稳态种间资源竞争研究进展。实验室内稳态条件下的竞争理论得到了大量实验验证,由于自然水体处于非稳状态,非稳态条件下藻类种间资源竞争日益成为藻类水华形成机制研究的重点。主要理论包括Tilman的R*法则和资源比假说,Grover的拾遗者-机会主义者交替竞争假说(VIS),中度干扰假说(IDH)及关键光强(CLIH)假说。这些假说的提出丰富了藻类资源竞争的理论体系,为研究自然状态下藻类增殖和竞争行为奠定理论基础。今后的藻类种间资源竞争的研究重点为非稳态下物理条件变化以及物理、化学和生物资源共同变化下藻类种间关系及生物多样性的研究。     

Proximate mechanisms of contest competition among female Bwindi mountain gorillas (Gorilla beringei beringei)

Socioecological models provide a framework for predicting how animals respond competitively to the abundance and distribution of food resources. Testing predictions of socioecological models requires analysis of relationships among food resource characteristics, competitive behaviors, and measures of rank-related skew in energy balance or reproductive success. A positive relationship between dominance rank and energy balance has been observed among female mountain gorillas in Bwindi Impenetrable National Park, Uganda. This study examines the proximate mechanisms underlying feeding competition among those females. To assess the contestability of food resources, we measured the time a female spent feeding at a food site (food site residence time). We also examined the relationship between dominance rank and the access to resources, as well as the rate, context, and direction of aggression, and the number of neighbors in close proximity. As predicted, females had longer food site residence times and higher aggression rates with fruit and decaying wood than with herbaceous vegetation, suggesting that those resources may be contestable. Aggression was predominantly directed down the dominance hierarchy, although against expectation, rank was not significantly correlated with aggression rates or the time spent feeding on contestable foods. Higher-ranking females had significantly fewer neighbors, suggesting that lower-ranking females avoid higher-ranking ones. This study provides additional support for the claim that there is variability in how primates respond to the quality and distribution of food resources and that avoidance as a strategy to cope with feeding competition may result in similar skew in energy balance as rank-related aggression.     

Parent-offspring resemblance in degree of sociality in a passerine bird

The tendency to aggregate with conspecifics is a common type of social behavior, and interspecific differences in degree of sociality may indicate that evolutionary changes in sociality have been shaped by past selection pressures. A pre-requisite for such evolutionary change is that the behavior in question has an additive genetic basis. I investigated parent-offspring resemblance in two characteristics of sociality (colony size and nearest neighbor distance) in the semi-colonial barn swallow Hirundo rustica. Heritability estimates of these phenotypic characters were statistically significant. Offspring that were transferred to foster nests as part of brood size manipulation or cross-fostering experiments resembled their original parents more than their foster parents with respect to sociality. There was little evidence of phenotypic characteristics of mothers being significantly related to colony size or nearest neighbor distance of their offspring. Maintenance of genetic variation in sociality in this species may be related to the facts that the relationship between reproductive success and colony size differs among years, and that individuals differing in phenotypic quality have different optimal colony sizes.     

Evidence of large-scale source-sink dynamics and long-distance dispersal among Wood Thrush populations

Source-sink dynamics are commonly thought to occur among Wood Thrush (Hylocichla mustelina) and other songbird populations, allowing for the persistence of populations with negative growth rates ("sinks") through immigration from populations with positive growth rates ("sources"). Knowledge of source-sink dynamics is important for management and conservation because the removal of source habitat should result in the extinction of dependent sinks. However, since research has focused on identifying individual sources/sink populations, not source-sink pairs, we cannot predict these effects or the scale over which they occur. We posit that, when dispersal occurs from a source to a sink year after year, there will be a one-year time-lagged correlation in abundance between the two populations. This should occur for populations separated by distances over which juveniles disperse. Using the North American Breeding Bird Survey data, we tested for such time-lagged correlations between paired Wood Thrush populations from 10 to 200 km apart. Populations were linked with a one-year time lag over distances from 60 to 80 km, indicating that dispersal and source-sink dynamics may occur over these long distances. There was also a declining trend in forest cover from sources to sinks. Conservation and management strategies should therefore be designed at large scales, with consideration for source-sink dynamics and forest cover.     

Variation in sperm morphometry and sperm competition among barn swallow (Hirundo rustica) populations

Spermatozoa vary greatly in size and shape among species across the animal kingdom. Postcopulatory sexual selection is thought to be the major evolutionary force driving this diversity. In contrast, less is known about how sperm size varies among populations of the same species. Here, we investigate geographic variation in sperm size in barn swallows Hirundo rustica, a socially monogamous passerine with a wide Holarctic breeding distribution. We included samples from seven populations and three subspecies: five populations of ssp. rustica in Europe (Czech, Italy, Norway, Spain, and Ukraine), one population of ssp. transitiva in Israel, and one population of ssp. erythrogaster in Canada. All sperm traits (head length, midpiece length, tail length, and total length) varied significantly among populations. The variation among the European rustica populations was much lower than the differences among subspecies, indicating that sperm traits reflect phylogenetic distance. We also performed a test of the relationship between the coefficient of between-male variation in total sperm length and extrapair paternity levels across different populations within a species. Recent studies have found a strong negative relationship between sperm size variation and extrapair paternity among species. Here, we show a similar negative relationship among six barn swallow populations, which suggests that the variance in male sperm length in a population is shaped by the strength of stabilizing postcopulatory sexual selection.     

The good and the bad stopover: behaviours of migrant reed warblers at two contrasting sites

Successful migration for passerine birds depends largely on the quality of stopover habitats, but we still lack complete knowledge of how migrants search for habitats en route and how they behave when landing at poor quality stopover sites. We compared the distance of exploratory movements and stopover durations of the reed warbler Acrocephalus scirpaceus, a reedbed habitat specialist, released at suitable (reed bed) and unsuitable (sand dune) stopover sites. Birds tape-lured during nocturnal migration to a sand dune were captured, radio-tagged, released and tracked at two sites of contrasting habitat quality. Lean birds were found to move further in the dunes (max. 300 m) than in reeds (max. 200 m), whereas ‘fat’ individuals at both sites remained stationary. Birds spent just 1 day in the dunes and up to 13 days in the reeds. Our results suggest that some nocturnal migrants with restricted diurnal exploratory movements depend on stopover site selection when ceasing nocturnal flight.     

Phylogenetic Comparative Methods Strengthen Evidence for Reduced Genetic Diversity among Endangered Tetrapods

Abstract: The fitness of species with little genetic diversity is expected to be affected by inbreeding and an inability to respond to environmental change. Conservation theory suggests that endangered species will generally demonstrate lower genetic diversity than taxa that are not threatened. This hypothesis has been challenged because the time frame of anthropogenic extinction may be too fast to expect genetic factors to significantly contribute. I conducted a meta‐analysis to examine how genetic diversity in 894 tetrapods correlates with extinction threat level. Because species are not evolutionarily independent, I used a phylogenetic regression framework to address this issue. Mean genetic diversity of tetrapods, as assessed by protein heterozygosity, was 29.7–31.5% lower on average in threatened species than in their nonthreatened relatives, a highly significant reduction. Within amphibians as diversity decreased extinction risk increased in phylogenetic models, but not in nonphylogenetic regressions. The effects of threatened status on diversity also remained significant after accounting for body size in mammals. These results support the hypothesis that genetic effects on population fitness are important in the extinction process.     

Context-dependent genetic benefits of extra-pair mate choice in a socially monogamous passerine

Extra-pair paternity is common in socially monogamous passerines; however, despite considerable research attention, consistent differences in fitness between within-pair offspring (WPO) and extra-pair offspring (EPO) have not been demonstrated. Recent evidence indicates that differences between maternal half-siblings may depend on environmental conditions, but it is unclear whether the influence of paternal genetic contribution should be most apparent under comparatively poor or favourable conditions. We compared phenotypic characteristics of WPO and EPO in 30 mixed-paternity broods of the tree swallow (Tachycineta bicolor) in relation to experimentally increased nest temperature (n = 13 heated nests; 17 control nests) and natural abundance of haematophagous parasites (Protocalliphora spp.). This allowed us to test the hypothesis that genetic benefits of extra-pair mating are environment dependent. EPO grew their ninth primary feathers faster than WPO regardless of nest temperature or parasite load and had significantly longer ninth primary feathers at fledging when parasite abundance was low, and when they were positioned early in the hatching sequence relative to WPO. In contrast, WPO under similar conditions did not differ from EPO in any phenotypic trait measured. These results indicate that the fitness benefits of extra-pair mating are likely to be context dependent, and that genetic effects on some phenotypic traits may be more apparent when conditions are relatively favourable.     

Fitness consequences of pre- and post-fledging timing decisions in a double-brooded passerine

The fitness consequences of a delayed timing of breeding are expected to affect the temporal characteristics of the whole annual breeding system. One major problem in quantifying the fitness relevance of timing is that individual differences between pairs may cause the seasonal trend. Differentials in juvenile survival due to pre-fledging timing decisions often only appear after fledging of the chicks. Therefore, timing decisions in the post-fledging period, i.e., the duration of parental care, might additionally influence juvenile survival. We tested the effects of timing and parental competence on the post-fledging survival of second-brood juvenile Barn Swallows (Hirundo rustica L.) by swapping earlier and later hatching clutches and radio-tracking the juvenile subjects. The mark-recapture models controlled for the effects of duration of post-fledging care and food availability. There was an annually varying negative seasonal trend in offspring survival that was associated with environmental conditions. Directional selection for early breeding occurred in the two years with scarce autumnal food supply. Furthermore, we found strong selection for long post-fledging parental care. The duration of care neither declined seasonally, nor did longer care compensate for the seasonal decline of juvenile survival. Hence, the reproductive output three weeks after fledging was determined by two parental timing decisions: the timing of breeding and the timing of family breakup. We suggest that differential survival of second-brood fledglings in relation to these decisions is an important part of the selective mechanisms shaping the reproductive system of Barn Swallows.     

The impacts of migrants on environmental degradation in developing countries

Environmental and Ecological Statistics - The prolonged issue of environmental degradation, especially in developing countries, has urgently called for a solution by first identifying the source of...