Evolution of density‐dependent movement during experimental range expansions

Range expansions and biological invasions are prime examples of transient processes that are likely impacted by rapid evolutionary changes. As a spatial process, range expansions are driven by dispersal and movement behaviour. Although it is widely accepted that dispersal and movement may be context‐dependent, for instance density‐dependent, and best represented by reaction norms, the evolution of density‐dependent movement during range expansions has received little experimental attention. We therefore tested current theory predicting the evolution of increased movement at low densities at range margins using highly replicated and controlled range expansion experiments across multiple genotypes of the protist model system Tetrahymena thermophila. Although rare, we found evolutionary changes during range expansions even in the absence of initial standing genetic variation. Range expansions led to the evolution of negatively density‐dependent movement at range margins. In addition, we report the evolution of increased intrastrain competitive ability and concurrently decreased population growth rates in range cores. Our findings highlight the importance of understanding movement and dispersal as evolving reaction norms and plastic life‐history traits of central relevance for range expansions, biological invasions and the dynamics of spatially structured systems in general.     

Density‐dependent fitness,not dispersal movements,drives temporal variation in spatial genetic structure in dark‐eyed juncos (Junco hyemalis)

Some studies have found that dispersal rates and distances increase with density, indicating that density‐dependent dispersal likely affects spatial genetic structure. In an 11‐year mark–recapture study on a passerine, the dark‐eyed junco, we tested whether density affected dispersal distance and/or fine‐scale spatial genetic structure. Contrary to expectations, we found no effect of predispersal density on dispersal distance or the proportion of locally produced juveniles returning to the population from which they hatched. However, even though density did not affect dispersal distance or natal return rates, we found that density still did affect spatial genetic structure. We found significant positive spatial genetic structure at low densities of (postdispersal) adults but not at high densities. In years with high postdispersal (adult) densities that also had high predispersal (juvenile) densities in the previous year, we found negative spatial genetic structure, indicating high levels of dispersal. We found that density also affected fitness of recruits, and fitness of immigrants, potentially linking these population parameters with the spatial genetic structure detected. Immigrants and recruits rarely nested in low postdispersal density years. In contrast, in years with high postdispersal density, recruits were common and immigrants had equal success to local birds, so novel genotypes diluted the gene pool and effectively eliminated positive spatial genetic structure. In relation to fine‐scale spatial genetic structure, fitness of immigrants and new recruits is poorly understood compared to dispersal movements, but we conclude that it can have implications for the spatial distribution of genotypes in populations.     

Performance of individual vs. group sampling for inferring dispersal under isolation‐by‐distance

Models of isolation‐by‐distance formalize the effects of genetic drift and gene flow in a spatial context where gene dispersal is spatially limited. These models have been used to show that, at an appropriate spatial scale, dispersal parameters can be inferred from the regression of genetic differentiation against geographic distance between sampling locations. This approach is compelling because it is relatively simple and robust and has rather low sampling requirements. In continuous populations, dispersal can be inferred from isolation‐by‐distance patterns using either individuals or groups as sampling units. Intrigued by empirical findings where individual samples seemed to provide more power, we used simulations to compare the performances of the two methods in a range of situations with different dispersal distributions. We found that sampling individuals provide more power in a range of dispersal conditions that is narrow but fits many realistic situations. These situations were characterized not only by the general steepness of isolation‐by‐distance but also by the intrinsic shape of the dispersal kernel. The performances of the two approaches are otherwise similar, suggesting that the choice of a sampling unit is globally less important than other settings such as a study's spatial scale.     

Density‐dependent sex‐biased development of macroptery in a water strider

In wing‐polymorphic insects, wing morphs differ not only in dispersal capability but also in life history traits because of trade‐offs between flight capability and reproduction. When the fitness benefits and costs of producing wings differ between males and females, sex‐specific trade‐offs can result in sex differences in the frequency of long‐winged individuals. Furthermore, the social environment during development affects sex differences in wing development, but few empirical tests of this phenomenon have been performed to date. Here, I used the wing‐dimorphic water strider Tenagogerris euphrosyne to test how rearing density and sex ratio affect the sex‐specific development of long‐winged dispersing morphs (i.e., sex‐specific macroptery). I also used a full‐sib, split‐family breeding design to assess genetic effects on density‐dependent, sex‐specific macroptery. I reared water strider nymphs at either high or low densities and measured their wing development. I found that long‐winged morphs developed more frequently in males than in females when individuals were reared in a high‐density environment. However, the frequency of long‐winged morphs was not biased according to sex when individuals were reared in a low‐density environment. In addition, full‐sib males and females showed similar macroptery incidence rates at low nymphal density, whereas the macroptery incidence rates differed between full‐sib males and females at high nymphal density. Thus complex gene‐by‐environment‐by‐sex interactions may explain the density‐specific levels of sex bias in macroptery, although this interpretation should be treated with some caution. Overall, my study provides empirical evidence for density‐specific, sex‐biased wing development. My findings suggest that social factors as well as abiotic factors can be important in determining sex‐biased wing development in insects.     

No evidence of sex‐biased dispersal in an island population of Common Blackbirds Turdus merula

Natal dispersal has major consequences for the dynamics and genetic structure of populations. Female‐biased natal dispersal, otherwise the norm in birds, is overridden when the place to move is limited, as on isolated islands. This effect was confirmed for the fist time in a European study system, the Common Blackbird breeding on Heligoland Island. Spatially restricted and sexually uniform natal dispersal may be a prerequisite for successful establishment of populations on remote islands or isolated habitat fragments, and this could play a major role in speciation processes.     

Symbiont physiology and population dynamics before and during symbiont shifts in a flexible algal‐cnidarian symbiosis

For cnidarians that can undergo shifts in algal symbiont relative abundance, the underlying algal physiological changes that accompany these shifts are not well known. The sea anemone Anthopleura elegantissima associates with the dinoflagellate Symbiodinium muscatinei and the chlorophyte Elliptochloris marina, symbionts with very different tolerances to light and temperature. We compared the performance of these symbionts in anemones maintained in an 8–11.5 month outdoor common garden experiment with simulated intertidal conditions and three levels of shading (2, 43, and 85% ambient irradiance). Symbiont densities, mitotic indices, photophysiology and pigments were assessed at three time points during the summer, a period of high irradiance and solar heating during aerial exposure. Whereas S. muscatinei was either neutrally or positively affected by higher irradiance treatments, E. marina responded mostly negatively to high irradiance. E. marina in the 85% irradiance treatment exhibited significantly reduced P_max and chlorophyll early in the summer, but it was not until nearly 3 months later that a shift in symbiont relative abundance toward S. muscatinei occurred, coincident with bleaching. Symbiont densities and proportions remained largely stable in all other treatments over time, and displacement of S. muscatinei by E. marina was not observed in the 2% irradiance treatment despite the potentially better performance of E. marina. While our results support the view that rapid changes in symbiont relative abundance are typically associated with symbiont physiological dysfunction and bleaching, they also show that significant temporal lags may occur between the onset of symbiont stress and shifts in symbiont relative abundances.     

Hybridization selects for prime‐numbered life cycles in Magicicada: An individual‐based simulation model of a structured periodical cicada population

We investigate competition between separate periodical cicada populations each possessing different life‐cycle lengths. We build an individual‐based model to simulate the cicada life cycle and allow random migrations to occur between patches inhabited by the different populations. We show that if hybridization between different cycle lengths produces offspring that have an intermediate life‐cycle length, then predation acts disproportionately to select against the hybrid offspring. This happens because they emerge in low densities without the safety‐in‐numbers provided by either parent population. Thus, prime‐numbered life cycles that can better avoid hybridization are favored. However, we find that this advantage of prime‐numbered cycles occurs only if there is some mechanism that can occasionally synchronize emergence between local populations in sufficiently many patches.     

The role of learning by a predator,Rivulus hartii,in the rare‐morph survival advantage in guppies

Negative frequency‐dependent selection (NFDS), where rare types are favoured by selection, can maintain diversity. However, the ecological processes that mediate NFDS are often not known. Male guppies (Poecilia reticulata) exhibit extreme diversity of colour patterning and, in a previous field experiment, rare morphs had a survival advantage. Here, we test the hypothesis that predators impose NFDS because they are efficient at capturing familiar prey morphs, but are less efficient at capturing unfamiliar morphs. Over a series of trials, we presented Rivulus hartii, a natural predator of guppies, with male guppies with the same colour patterning (A trials); then, for a second series of trials, we presented the rivulus with guppies with a new colour pattern (B trials). The success of rivulus at capturing guppies on the first attack increased over successive A trials. First attack success decreased significantly for the early B trials, and then increased during successive B trials, eventually reaching the same level as in the best A trials. This experiment demonstrates that learning, perhaps through long‐term search image formation, plays a role in predation success on familiar vs. unfamiliar prey morphs. These results support the hypothesis that predator learning contributes to the maintenance of the extreme male guppy polymorphism seen in nature.     

Estimating seed dispersal distance: A comparison of methods using animal movement and plant genetic data on two primate‐dispersed Neotropical plant species

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Parentage analyses identify local dispersal events and sibling aggregations in a natural population of Millepora hydrocorals,a free‐spawning marine invertebrate

Dispersal is a critical process for the persistence and productivity of marine populations. For many reef species, there is increasing evidence that local demography and self‐recruitment have major consequences on their genetic diversity and adaptation to environmental change. Yet empirical data of dispersal patterns in reef‐building species remain scarce. Here, we document the first genetic estimates of self‐recruitment and dispersal distances in a free‐spawning marine invertebrate, the hydrocoral Millepora cf. platyphylla. Using twelve microsatellite markers, we gathered genotypic information from 3,160 georeferenced colonies collected over 27,000 m² of a single reef in three adjacent habitats in Moorea, French Polynesia; the mid slope, upper slope, and back reef. Although the adult population was predominantly clonal (85% were clones), our parentage analysis revealed a moderate self‐recruitment rate with a minimum of 8% of sexual propagules produced locally. Assigned offspring often settled at <10 m from their parents and dispersal events decrease with increasing geographic distance. There were no discrepancies between the dispersal distances of offspring assigned to parents belonging to clonal versus nonclonal genotypes. Interhabitat dispersal events via cross‐reef transport were also detected for sexual and asexual propagules. Sibship analysis showed that full siblings recruit nearby on the reef (more than 40% settled at <30 m), resulting in sibling aggregations. Our findings highlight the importance of self‐recruitment together with clonality in stabilizing population dynamics, which may ultimately enhance local sustainability and resilience to disturbance.     

Resistance,tolerance and environmental transmission dynamics determine host extinction risk in a load‐dependent amphibian disease

While disease‐induced extinction is generally considered rare, a number of recently emerging infectious diseases with load‐dependent pathology have led to extinction in wildlife populations. Transmission is a critical factor affecting disease‐induced extinction, but the relative importance of transmission compared to load‐dependent host resistance and tolerance is currently unknown. Using a combination of models and experiments on an amphibian species suffering extirpations from the fungal pathogen Batrachochytrium dendrobatidis (Bd), we show that while transmission from an environmental Bd reservoir increased the ability of Bd to invade an amphibian population and the extinction risk of that population, Bd‐induced extinction dynamics were far more sensitive to host resistance and tolerance than to Bd transmission. We demonstrate that this is a general result for load‐dependent pathogens, where non‐linear resistance and tolerance functions can interact such that small changes in these functions lead to drastic changes in extinction dynamics.     

Host dispersal shapes the population structure of a tick‐borne bacterial pathogen

Birds are hosts for several zoonotic pathogens. Because of their high mobility, especially of longdistance migrants, birds can disperse these pathogens, affecting their distribution and phylogeography. We focused on Borrelia burgdorferi sensu lato, which includes the causative agents of Lyme borreliosis, as an example for tick‐borne pathogens, to address the role of birds as propagation hosts of zoonotic agents at a large geographical scale. We collected ticks from passerine birds in 11 European countries. B. burgdorferi s.l. prevalence in Ixodes spp. was 37% and increased with latitude. The fieldfare Turdus pilaris and the blackbird T. merula carried ticks with the highest Borrelia prevalence (92 and 58%, respectively), whereas robin Erithacus rubecula ticks were the least infected (3.8%). Borrelia garinii was the most prevalent genospecies (61%), followed by B. valaisiana (24%), B. afzelii (9%), B. turdi (5%) and B. lusitaniae (0.5%). A novel Borrelia genospecies “Candidatus Borrelia aligera” was also detected. Multilocus sequence typing (MLST) analysis of B. garinii isolates together with the global collection of B. garinii genotypes obtained from the Borrelia MLST public database revealed that: (a) there was little overlap among genotypes from different continents, (b) there was no geographical structuring within Europe, and (c) there was no evident association pattern detectable among B. garinii genotypes from ticks feeding on birds, questing ticks or human isolates. These findings strengthen the hypothesis that the population structure and evolutionary biology of tick‐borne pathogens are shaped by their host associations and the movement patterns of these hosts.     

Mark‐recapture modeling accounting for state uncertainty provides concurrent estimates of survival and fecundity in a protected harbor seal population

Harbor seal breeding behavior and habitats constrain opportunities for individual‐based studies, and no current estimates of both survival and fecundity exist for any of the populations studied worldwide. As a result, the drivers underlying the variable trends in abundance exhibited by harbor seal populations around the world remain uncertain. We developed an individual‐based study of harbor seals in northeast Scotland, whereby data were collected during daily photo‐identification surveys throughout the pupping seasons between 2006 and 2011. However, a consequence of observing seals remotely meant that information on sex, maturity‐stage, or breeding status was not always available. To provide unbiased estimates of survival rates we conditioned initial release of individuals on the first time sex was known to estimate sex‐specific survival rates, while a robust design multistate model accounting for uncertainty in breeding status was used to estimate reproductive rate of multiparous and ≥3‐yr‐old females. Survival rates were estimated at 0.95 (95% CI = 0.91–0.97) for females and 0.92 (0.83–0.96) for males, while reproductive rate was estimated at 0.89 (0.75–0.95) for multiparous and 0.69 (0.64–0.74) for ≥3‐yr‐old females. Stage‐based population modeling indicated that this population should be recovering, even under the current shooting quotas implemented by the recent management plan.     

The role of immigration and local adaptation on fine‐scale genotypic and phenotypic population divergence in a less mobile passerine

Dispersal and local patterns of adaptation play a major role on the ecological and evolutionary trajectory of natural populations. In this study, we employ a combination of genetic (25 microsatellite markers) and field‐based information (seven study years) to analyse the impact of immigration and local patterns of adaptation in two nearby (< 7 km) blue tit (Cyanistes caeruleus) populations. We used genetic assignment analyses to identify immigrant individuals and found that dispersal rate is female‐biased (72%). Data on lifetime reproductive success indicated that immigrant females produced fewer local recruits than their philopatric counterparts whereas immigrant males recruited more offspring than those that remained in their natal location. In spite of the considerably higher immigration rates of females, our results indicate that, in absolute terms, their demographic and genetic impact in the receiving populations is lower than that in immigrant males. Immigrants often brought novel alleles into the studied populations and a high proportion of them were transmitted to their recruits, indicating that the genetic impact of immigrants is not ephemeral. Although only a few kilometres apart, the two study populations were genetically differentiated and showed strong divergence in different phenotypic and life‐history traits. An almost absent inter‐population dispersal, together with the fact that both populations receive immigrants from different source populations, is probably the main cause of the observed pattern of genetic differentiation. However, phenotypic differentiation (P_ST) for all the studied traits greatly exceeded neutral genetic differentiation (F_ST), indicating that divergent natural selection is the prevailing factor determining the evolutionary trajectory of these populations. Our study highlights the importance of integrating individual‐ and population‐based approaches to obtain a comprehensive view about the role of dispersal and natural selection on structuring the genotypic and phenotypic characteristics of natural populations.     

Density‐dependent effects on reproductive performance in a recovering population of White‐tailed Eagles Haliaeetus albicilla

Understanding the mechanisms that shape density‐dependent processes and population dynamics is often essential for species conservation. Two key mechanisms of density‐dependent reductions in reproductive performance are a limited access to foraging habitats (the habitat heterogeneity hypothesis) and territorial aggression towards conspecifics (the interference competition hypothesis) at high population densities. Disentangling the relative importance of these mechanisms within populations below their carrying capacity is important for the evaluation of the success of conservation measures. However, relatively few studies have attempted to quantify the relative importance of both mechanisms for the reproductive performance of a population. Many raptor populations are ideal model systems to investigate density‐dependent effects because they are currently recovering from human‐induced reductions during the last decades. Using a 14‐year dataset, we combined analyses of individual reproductive performance with a mechanistic population model to investigate early signs of density‐dependent regulation in a population of White‐tailed Eagles Haliaeetus albicilla in north‐east Germany. We found a negative effect of the number of neighbouring breeding pairs and a positive effect of water surface area (as a proxy for the availability of favourable foraging habitat) on breeding success and on the average number of nestlings. The mean nearest neighbour distance between breeding pairs has decreased, and the mean distance of nests to the nearest water body has increased over the last 14 years. Moreover, the population model indicates that even though the population is still growing, carrying capacity could be reached at about 500–950 territorial pairs. These results suggest that the selection of nesting sites is determined by a trade‐off between the distance to favourable foraging habitat and the distance to neighbouring breeding pairs. To avoid increasing competition with conspecifics, due to continued population growth, breeding pairs seem to select increasingly suboptimal habitats. Therefore, our results suggest that the habitat heterogeneity and interference competition hypotheses are not necessarily mutually exclusive as mechanisms of density‐dependent population regulation, but can determine the reproductive performance of a raptor population simultaneously. Thus, a future decline in breeding success does not necessarily reflect a decrease in habitat quality but may rather be a consequence of density‐dependent mechanisms. This information may be useful for the interpretation of population trends and for the development of appropriate management strategies for recovering raptor populations.     

Parentage analysis of Ansell's mole‐rat family groups indicates a high reproductive skew despite relatively relaxed ecological constraints on dispersal

To better understand evolutionary pathways leading to eusociality, interspecific comparisons are needed, which would use a common axis, such as that of reproductive skew, to array species. African mole‐rats (Bathyergidae, Rodentia) provide an outstanding model of social evolution because of a wide range of social organizations within a single family; however, their reproductive skew is difficult to estimate, due to their cryptic lifestyle. A maximum skew could theoretically be reached in groups where reproduction is monopolized by a stable breeding pair, but the value could be decreased by breeding‐male and breeding‐female turnover, shared reproduction and extra‐group mating. The frequency of such events should be higher in species or populations inhabiting mesic environments with relaxed ecological constraints on dispersal. To test this prediction, we studied patterns of parentage and relatedness within 16 groups of Ansell's mole‐rat (Fukomys anselli) in mesic miombo woodland. Contrary to expectation, there was no shared reproduction (more than one breeder of a particular sex) within the studied groups, and proportion of immigrants and offspring not assigned to current breeding males was low. The within‐group parentage and relatedness patterns observed resemble arid populations of ‘eusocial’ Fukomys damarensis, rather than a mesic population of ‘social’ Cryptomys hottentotus. As a possible explanation, we propose that the extent ecological conditions affect reproductive skew may be markedly affected by life history and natural history traits of the particular species and genera.