Insect migration: Variability and success in a capricious environment

Stochastic effects of climate and weather have a pervasive influence on the induction, performance and evolution of migration. In wing-dimorphic species, their influence on habitat quality, and on rates of development of the migrant itself, maintains variation in responses to environmental cues determining wing-form and migratory behaviour. Migrants flying above their flight boundary layer rely on winds to disperse them across landscapes in which their habitats are distributed. Patterns of distribution of habitat patches, and the influence of changing windspeeds and direction on the displacements of migrants, result in selection for variation in migratory potential at each migration. In subsequent migrations, this variation and stochastic effects of the winds on groundtracks of individual migrants ensure that their destinations ‘sample’ the landscapes they travel over. The extent and resolution of this sampling, by which migrants reach favourable habitats, depend on the components of migratory potential, their mode of inheritance, and genetic correlations between them, as well as on the characteristics of the winds on which they travel.     

A long winter for the Red Queen: rethinking the evolution of seasonal migration

This paper advances an hypothesis that the primary adaptive driver of seasonal migration is maintenance of site fidelity to familiar breeding locations. We argue that seasonal migration is therefore principally an adaptation for geographic persistence when confronted with seasonality – analogous to hibernation, freeze tolerance, or other organismal adaptations to cyclically fluctuating environments. These ideas stand in contrast to traditional views that bird migration evolved as an adaptive dispersal strategy for exploiting new breeding areas and avoiding competitors. Our synthesis is supported by a large body of research on avian breeding biology that demonstrates the reproductive benefits of breeding‐site fidelity. Conceptualizing migration as an adaptation for persistence places new emphasis on understanding the evolutionary trade‐offs between migratory behaviour and other adaptations to fluctuating environments both within and across species. Seasonality‐induced departures from breeding areas, coupled with the reproductive benefits of maintaining breeding‐site fidelity, also provide a mechanism for explaining the evolution of migration that is agnostic to the geographic origin of migratory lineages (i.e. temperate or tropical). Thus, our framework reconciles much of the conflict in previous research on the historical biogeography of migratory species. Although migratory behaviour and geographic range change fluidly and rapidly in many populations, we argue that the loss of plasticity for migration via canalization is an overlooked aspect of the evolutionary dynamics of migration and helps explain the idiosyncratic distributions and migratory routes of long‐distance migrants. Our synthesis, which revolves around the insight that migratory organisms travel long distances simply to stay in the same place, provides a necessary evolutionary context for understanding historical biogeographic patterns in migratory lineages as well as the ecological dynamics of migratory connectivity between breeding and non‐breeding locations.     

Ancestry and evolution of seasonal migration in the Parulidae

Seasonal migration in birds is known to be highly labile and subject to rapid change in response to selection, such that researchers have hypothesized that phylogenetic relationships should neither predict nor constrain the migratory behaviour of a species. Many theories on the evolution of bird migration assume a framework that extant migratory species have evolved repeatedly and relatively recently from sedentary tropical or subtropical ancestors. We performed ancestral state reconstructions of migratory behaviour using a comprehensive, well-supported phylogeny of the Parulidae (the 'wood-warblers'), a large family of Neotropical and Nearctic migratory and sedentary songbirds, and examined the rates of gain and loss of migration throughout the Parulidae. Counter to traditional hypotheses, our results suggest that the ancestral wood-warbler was migratory and that losses of migration have been at least as prevalent as gains throughout the history of Parulidae. Therefore, extant sedentary tropical radiations in the Parulidae represent losses of latitudinal migration and colonization of the tropics from temperate regions. We also tested for phylogenetic signal in migratory behaviour, and our results indicate that although migratory behaviour is variable within some wood-warbler species and clades, phylogeny significantly predicts the migratory distance of species in the Parulidae.     

Phenotypic divergence during speciation is inversely associated with differences in seasonal migration

Differences in seasonal migration might promote reproductive isolation and differentiation by causing populations in migratory divides to arrive on the breeding grounds at different times and/or produce hybrids that take inferior migratory routes. We examined this question by quantifying divergence in song, colour, and morphology between sister pairs of North American migratory birds. We predicted that apparent rates of phenotypic differentiation would differ between pairs that do and do not form migratory divides. Consistent with this prediction, results from mixed effects models and Ornstein–Uhlenbeck models of evolution showed different rates of divergence between these groups; surprisingly, differentiation was greater among non-divide pairs. We interpret this finding as a result of variable rates of population blending and fusion between partially diverged forms. Ancient pairs of populations that subsequently fused are now observed as a single form, whereas those that did not fuse are observable as pairs and included in our study. We propose that fusion of two populations is more likely to occur when they have similar migratory routes and little other phenotypic differentiation that would cause reproductive isolation. By contrast, pairs with migratory divides are more likely to remain reproductively isolated, even when differing little in other phenotypic traits. These findings suggest that migratory differences may be one among several isolating barriers that prevent divergent populations from fusing and thereby increase the likelihood that they will continue differentiating as distinct species.     

Bio-energetic data show weak spatial but strong seasonal differences in wetland quality for waders in a Mediterranean migration bottleneck

1. Assessing the quality of wetlands as refuelling areas for migrating waterbirds based on resource distribution patterns is challenging. Resources in wetlands can vary both horizontally and vertically and may be differentially available to different bird species at different times of the year. Therefore, the extent to which wetland quality can be generalised across seasons and for a diversity of birds remains unresolved.
2. Spatiotemporal variation in abundance and quality of macrobenthos as food for migrating waders was studied in a set of wetland areas near a Mediterranean migration bottleneck in the Balkan peninsula, during both spring and autumn migration. Samples were subdivided into different depth layers to differentiate between parts of the sediment that are accessible to different groups of wader species. To quantify food availability and the resulting refuelling capacity in different wetland habitat types, abundance, wet weight, and lipid and protein content of invertebrate taxa were determined for each sample.
3. Invertebrate food availability and quality were markedly higher in spring than in autumn. Given the higher abundance and protein and lipid content of prey in spring, the total energy that could be harvested in spring (3.81 ± 0.79 kJ/m²) was about 7 times higher than in autumn (0.56 ± 0.12 kJ/m²). Most prey were found in the top layer of the sediment (0–22 mm depth), but about a third of the total energy was present in the deepest soil layer that only longer-billed species could reach (55–200 mm depth). Higher quality prey items such as larger-bodied and heavier polychaetes and bivalves were found in the deeper sediment layers.
4. For other taxa, there was no evident vertical pattern of increasing individual body size. Prey abundance differed between certain habitat types but, overall, food availability could not be linked to distinct habitats. In spring, redox potential tended to be higher, while pH, vegetation cover, conductivity, and temperature were lower than in autumn.
5. Different wetland habitat classes used in a conservation framework may provide similar food resources for waders. As a result, linking food availability to habitat classifications is not straightforward. Furthermore, seasonal variation in wetland quality requires a re-evaluation of the importance of wetland areas during spring and autumn migration. Finally, nutritional analyses are essential for determining the capacity of wetlands to support refuelling by migratory waterbirds.

     

Dynamics of a harvested moose population in a variable environment

1. Population size, calves per female, female mean age and adult sex ratio of a moose ( Alces alces ) population in Vefsn, northern Norway were reconstructed from 1967 to 1993 using cohort analysis and catch-at-age data from 96% (6752) of all individuals harvested.
2. The dynamics of the population were influenced mainly by density-dependent harvesting, stochastic variation in climate and intrinsic variation in the age-structure of the female segment of the population.
3. A time delay in the assignment of hunting permits in relation to population size increased fluctuations in population size.
4. Selective harvesting of calves and yearlings increased the mean age of adult females in the population, and, because fecundity in moose is strongly age-specific, the number of calves per female concordantly increased. However, after years with high recruitment, the adult mean age decreased as large cohorts entered the adult age-groups. This age-structure effect generated cycles in the rate of recruitment to the population and fluctuations introduced time-lags in the population dynamics.
5. An inverse relationship between recruitment rate and population density, mediated by a density-dependent decrease in female body condition, could potentially have constituted a regulatory mechanism in the dynamics of the population, but this effect was counteracted by a density-dependent increase in the mean age of adult females.
6. Stochastic variation in winter snow depth and summer temperature had delayed effects on recruitment rate and in turn population growth rate, apparently through effects on female body condition before conception.     

Forcible eviction and prevention of recruitment in the clown anemonefish

How big an animal group will be depends on how the group's sizeis regulated and on the costs and benefits of living in thegroup. To determine which individuals regulate group size ofthe clown anemonefish, Amphiprion percula, I investigated thestrategies involved in the formation, maintenance, and dissolutionof its groups. Groups composed of a single breeding pair andof zero to four nonbreeding subordinates occupied individualsea anemones (Heteractis magnifica), which provided the fishwith oviposition sites and protection from predators. Groupsize increased linearly with anemone size. I used the residualsof this relationship as a measure of the degree of saturationof each anemone. Residents evicted low-rank subordinates andprevented the recruitment of additional subordinates at anemoneswith a high degree of saturation, but not at anemones with alow degree of saturation. These strategies indicate that residentscontrol group membership of their subordinates, and suggestthat residents might incur costs from the presence of subordinatesin more saturated anemones. In general, whenever residents cancontrol group membership, the prevention of recruitment andthe eviction of subordinates will set an upper limit on groupsize.     

Evidence of seasonal migration in a tropical subterranean vertebrate

Many animals undergo migrations that depend on a range of biotic or abiotic factors provoking daily, seasonal or annual movement of individuals and/or populations. Temperate amphibians frequently have their life cycles dominated by annual temperature fluctuations, but in the tropics it is often presumed that distinct rainy seasons will influence amphibians more than small changes in temperature. Here, we direct the seasonal changes in abundance hypothesis to a caecilian amphibian Boulengerula boulengeri found in monthly randomized quantitative surveys in the top 30 cm of soil. Meteorological data are used to interpret the significant changes found in relation to rainfall and temperature variables. Instead of the expected correlation of migration with rainfall, we find that frequency varies significantly and positively with temperature. In addition, data on the depth at which individual caecilians are collected suggest that animals undergo a vertical (rather than horizontal) migration within the soil. This is the first example of vertical migrations with temperature (as opposed to rainfall) of any tropical subterranean fauna. We discuss the possible influences that stimulate migration in this species (abiotic factors, feeding, reproduction and predation). The ecology of caecilians has lagged behind that of all other tetrapods, and this study has importance for future ecological studies, as well as biodiversity assessments and monitoring methodologies.     

The effects of the mating system on the evolution of migration in a spatially heterogeneous population

Summary Verbal explanations for the evolution of migration and dispersal often invoke inbreeding depression as an important force. Experimental work on plant populations indicates that while inbreeding depression may favor increased migration rates, adaptation to local environments may reduce the advantage to migrants. We formalize and test this hypothesis using a two-locus genetic model that incorporates lowered fitness in offspring produced by self-fertilization, and habitat differentiation. We also use the model to address questions about the general theory of genetic modifiers and the modifier reduction principle. We find that even under conditions when migration would increase the mean fitness of a population, migration may not be favored. This result is due to the associations that develop between genotypes at a locus subject to overdominant selection and at a neutral locus controlling the migration rate. Thus, it appears that, in this model, the forces of local adaptation, which favor a reduction in the migration rate, overwhelm those of inbreeding depression, which may favor dispersal.     

Reproductive migration of brown trout in a small Norwegian river studied by telemetry

The movement of 34 large (39–73 cm standard length) brown trout Salmo trutta was monitored using radio telemetry for up to 74 days in Brumunda, a small Norwegian river (mean annual discharge 3·3 m³ s⁻¹) flowing into the large Lake Mjøsa. The maximum range of movement in the river was 20 km. No clear relationships existed between individual movement and water discharge, temperature and barometric pressure. Brown trout migrated at all levels of water discharge. At low discharge (<2 m³ s⁻¹) movements were nocturnal. A weir 5·3 km from the outlet restricted ascending brown trout at low ( c . 6° C), but not at high ( c . 8° C) water temperatures. Spawning occurred in September to October and tagged individuals spent 2–51 days at the spawning sites. Mean migration speed from tagging to when the fish reached the spawning area, and from when they left the spawning areas and reached the lake was 1·0 and 2·3 km day⁻¹, respectively. All tagged brown trout that survived spawning returned to the lake after spawning.     

Cepaea nemoralis in a hostile environment: continuity, colonizations and morph-frequencies over time

In the 1930s Cyril Diver sampled and scored populations of the polymorphic snail Cepaea nemoralis on the South Haven Peninsula, Dorset. His recording methods make it possible to be confident about the absence of the species from parts of the peninsula, which has acid soils and changing habitats, and is generally unfavourable for the species. A resurvey in 1999 revealed a pattern of retreats and extensions of distribution related to local environmental change. Where populations have persisted over the period, the pattern of morph-frequency distributions has remained the same; there is equivocal evidence for a small increase in the frequency of midbanded shells. While most wetland populations have disappeared, new populations have become established along a roadside, and on newly-stabilized foredune ridges. Morph-frequencies in these new populations relate to those of the nearest established populations, and are sometimes monomorphic. The results are discussed in relation to the history of the peninsula, and to the role of migration in determining observed morph-frequency distributions.     

The first months at sea: marine migration and habitat use of sea trout Salmo trutta post‐smolts

The early migration and habitat use of brown trout Salmo trutta post‐smolts tagged with acoustic transmitters (n = 50) were investigated in a fjord system in central Norway from 30 April to 26 November 2014. The main aims were to investigate return rate, marine residence time and spatial use of the fjord system. Median seaward migration and return to fresh water dates were 22 May and 4 July, respectively. Of the 40 seaward migrating smolts, 26 returned to fresh water, giving a minimum return rate to fresh water of 65%. Entrance to the fjord from the river occurred mainly at night (80% of the S. trutta), however, no such diurnal pattern was observed during the return migration. Mean marine residence time was 38 days, but with large individual variation (22–99 days). The innermost parts of the study area were more utilized than the outer part of the fjord system during the sea residency, and with more use of the near shore habitat than the open, pelagic areas. Many post‐smolts also utilized the outer part of the fjord system, however, and 94% of the post‐smolts were recorded at least 14 km from the home river mouth. Marine survival and distribution in the fjord were size dependent with the largest individuals utilizing outer fjord areas and having higher return rates to fresh water. As far as is known, this is the first published study on temporal and spatial behaviour in the marine environment of first‐time S. trutta migrants during the full course of their first trip to sea.     

Postglacial population expansion drives the evolution of long-distance migration in a songbird

The evolution of long-distance migratory behavior from sedentary populations is a central problem in studies of animal migration. Three crucial issues that remain unresolved are: (1) the biotic and abiotic factors promoting evolution of migratory behavior, (2) the geographic origin of ancestral sedentary populations, and (3) the time scale over which migration evolves. We test the role of postglacial population expansions during the Quaternary in driving the evolution of songbird migration against prevailing views favoring the role of intraspecific competition. In contrast to previous attempts to investigate these questions using interspecific phylogenies, we adopt an intraspecific approach and examine the phylogeography of a North American songbird, the chipping sparrow (Spizella passerina), which exhibits both long-distance migratory behavior in temperate North America and sedentary behavior in Mexico and Central America. We show that migratory populations descend from sedentary populations in southern Mexico and that migration has evolved as a result of a northward population expansion into temperate North America since the last glacial maximum 18,000 years ago. Migration appears to have evolved rapidly in some species as populations colonized areas of high seasonality in the temperate zone. The phylogeography of the yellow-eyed junco (Junco phaeonotus), a strictly sedentary species, provides a null model supporting the view that northward range expansions were driven solely by environmental factors and not by a predisposition to evolve migratory behavior. These results provide the strongest evidence to date that historical climate patterns can drive the rapid evolution of avian migration in natural populations, and they suggest a general mechanism for the repeated evolution of migration within and across bird lineages.     

Partial migration in a landlocked brown trout population

Population densities of landlocked lake‐migratory brown trout Salmo trutta were estimated in two distinct lotic sections, separated by a lentic segment, in the Greåna River, Sweden, and individual growth and habitat use were monitored for 835 tagged brown trout from September 1998 to June 2000. Residency dominated in the upstream section where density of 0+ and 1+ year brown trout was low and growth rate high. In contrast, >90% of the brown trout that migrated to the lake originated from the downstream section, where density was high and growth rate low. For ≥2+ year individuals, growth rate was similar between the two stream sections, but densities were higher in the upstream than in the downstream section. Lake‐migrants had higher growth rates than non‐migrants (residents) during the autumn of both years. From September to May, migrants increased their body mass by >35%, whereas non‐migrants increased by <5%. Approximately 70% of the brown trout moved <10 m and <2% moved between the two stream sections, indicating that the lentic habitat might function as a barrier for juveniles. Differences in migratory behaviour, density and growth between the upstream and the downstream section might indicate that environmental factors influence the decision to migrate. It cannot be excluded, however, that the observed differences are genetically programmed, selected by migration costs that favour migratory behaviour downstream and residency upstream.     

The role of microRNAs in prostate cancer migration,invasion, and metastasis

Prostate cancer (PCa) is considered the most prevalent malignancy and the second major cause of cancer-related death in males from Western countries. PCa exhibits variable clinical pictures, ranging from dormant to highly metastatic cancer. PCa suffers from poor prognosis and diagnosis markers, and novel biomarkers are required to define disease stages and to design appropriate therapeutic approach by considering the possible genomic and epigenomic differences. MicroRNAs (miRNAs) comprise a class of small noncoding RNAs, which have remarkable functions in cell formation, differentiation, and cancer development and contribute in these processes through controlling the expressions of protein-coding genes by repressing translation or breaking down the messenger RNA in a sequence-specific method. miRNAs in cancer are able to reflect informative data about the current status of disease and this might benefit PCa prognosis and diagnosis since that is concerned to PCa patients and we intend to highlight it in this paper.     

Habitat selection by feral cats and dingoes in a semi‐arid woodland environment in central Australia

Habitat use by feral cats and dingoes was examined within a heterogeneous semi‐arid woodland site in central Australia over 2 years. Density estimates of feral cats based on tracks were higher in mulga habitat than in open habitat. Isodar analysis implied that this pattern of habitat use by feral cats was consistent with the consumer‐resource model of density‐dependent habitat selection, which is an ideal free solution. The reason why mulga supported higher densities of feral cats was unclear. Foraging success of feral cats may be higher in the mulga because the stalk and ambush hunting tactics typically employed by felids are well suited to dense cover. Mulga may also have offered feral cats more protection from dingo predation. Dingo activity was distributed uniformly across habitats. The dingo isodar was statistically non‐significant, suggesting that habitat selection by dingoes was independent of density.     

The effect of local change in habitat quality on populations of migratory species

1. Habitat deterioration is a major problem world-wide as a result of processes such as change in land use, introduced species, human disturbance and exploitation of food supplies. Many studies have shown that habitat change can have considerable effect on the numbers of individuals using a site. For migratory species, however, the consequences for the total population cannot be deduced from local studies.
2. For a migratory species, the change in total population size Δ N , as a consequence of habitat change in the wintering area, can be calculated from Δ N  =  LM γ d '/( b ' +  d '), where γ is the expected proportional change in the number of birds using a site as a result of the habitat change, L is the area affected, M is the density of individuals using the site prior to habitat change, b ' is the strength of the per capita density-dependent breeding output, and d ' is the strength of the per capita density-dependent winter mortality. Similarly the consequences of habitat change in the breeding area can be calculated from Δ N  =  LM γ b '/( b ' +  d ').
3. The same approach can be used for predicting the consequences of improvements in habitat quality.
4. A worked example is given to illustrate how this approach could be used to predict the consequences for the total population of changes in the food supply of oystercatchers within one estuary.
5. There is a need for more measures of γ, the expected proportional change in the number of birds using a site as a result of various forms of habitat deterioration, and the strengths of density dependence.