Regional neutrality evolves through local adaptive niche evolution.

Biodiversity in natural systems can be maintained either because niche differentiation among competitors facilitates stable coexistence or because equal fitness among neutral species allows for their long-term cooccurrence despite a slow drift toward extinction. Whereas the relative importance of these two ecological mechanisms has been well-studied in the absence of evolution, the role of local adaptive evolution in maintaining biological diversity through these processes is less clear. Here we study the contribution of local adaptive evolution to coexistence in a landscape of interconnected patches subject to disturbance. Under these conditions, early colonists to empty patches may adapt to local conditions sufficiently fast to prevent successful colonization by other preadapted species. Over the long term, the iteration of these local-scale priority effects results in niche convergence of species at the regional scale even though species tend to monopolize local patches. Thus, the dynamics evolve from stable coexistence through niche differentiation to neutral cooccurrence at the landscape level while still maintaining strong local niche segregation. Our results show that neutrality can emerge at the regional scale from local, niche-based adaptive evolution, potentially resolving why ecologists often observe neutral distribution patterns at the landscape level despite strong niche divergence among local communities.

The conflict between adaptation and dispersal for maintaining biodiversity in changing environments.

Dispersal and adaptation both allow species to persist in changing environments. Yet, we have limited understanding of how these processes interact to affect species persistence, especially in diverse communities where biotic interactions greatly complicate responses to environmental change. Here we use a stochastic metacommunity model to demonstrate how dispersal and adaptation to environmental change independently and interactively contribute to biodiversity maintenance. Dispersal provides spatial insurance, whereby species persist on the landscape by shifting their distributions to track favorable conditions. In contrast, adaptation allows species to persist by allowing for evolutionary rescue. But, when species both adapt and disperse, dispersal and adaptation do not combine positively to affect biodiversity maintenance, even if they do increase the persistence of individual species. This occurs because faster adapting species evolve to hold onto their initial ranges (i.e., monopolization effects), thus impeding slower adapting species from shifting their ranges and thereby causing extinctions. Importantly, these differences in adaptation speed emerge as the result of competition, which alters population sizes and colonization success. By demonstrating how dispersal and adaptation each independently and interactively contribute to the maintenance of biodiversity, we provide a framework that links the theories of spatial insurance, evolutionary rescue, and monopolization. This highlights the expectation that the maintenance of biodiversity in changing environments depends jointly on rates of dispersal and adaptation, and, critically, the interaction between these processes.

Transient Eco-Evolutionary Dynamics and the Window of Opportunity for Establishment of Immigrants.

AbstractTo what extent does landscape genetic structure bear the signature of arrival order of lineages during population assembly? Rapid genetic adaptation of resident populations founded by early colonists to local conditions might prevent establishment of later-arriving lineages, resulting in an evolution-mediated priority effect. This might result in a limited window of opportunity for establishment during which the resident population did not have sufficient time yet to monopolize the patch through local adaptation. The length of this window of opportunity is expected to depend on the degree to which early colonists and immigrants are preadapted to local habitat conditions. We present an intraspecific competition model of the initial transient population and evolutionary dynamics that quantifies the window of opportunity for establishment for asexual species. The model explicitly addresses the long-lasting effects of evolution-mediated priority effects by tracking lineages through time. Our results show that the difference in initial preadaptation between early colonists and late immigrants and the speed of evolution codetermine the window of opportunity for establishment. Our results also suggest that local populations should often be dominated by descendants of just a few early colonist lineages and that landscape genetic structure should often reflect the legacy of colonization history.

Adaptation reduces competitive dominance and alters community assembly.

A growing body of theory predicts that evolution of an early-arriving species in a new environment can produce a competitive advantage against later arriving species, therefore altering community assembly (i.e. the community monopolization hypothesis). Applications of the community monopolization hypothesis are increasing. However, experimental tests of the hypothesis are rare. Here, we provide a rare experimental demonstration of the community monopolization hypothesis using two archaeal species. We first expose one species to low- and high-temperature environments for 135 days. Populations in the high-temperature treatment evolved a 20% higher median performance when grown at high temperature. We then demonstrate that early arrival and adaptation reduce the abundance of a late-arriving species in the high-temperature environment by 63% relative to when both species arrive simultaneously and neither species is adapted to high temperature. These results are consistent with the community monopolization hypothesis and suggest that adaptation can reduce competitive dominance to alter community assembly. Hence, community monopolization might be much more common in nature than previously assumed. Our results strongly support the idea that patterns of biodiversity might often stem from a race between local adaptation and colonization of pre-adapted species.

LEGAL MEANS OF ENSURING COMPETITION IN PHARMACY.

OBJECTIVE: The aim: To research the consequences of pharmacy chains monopolization and establishment of legal means of neutralization of such consequences. PATIENTS AND METHODS: Materials and methods: The study is based on acts of the European Union, the United States, and Ukraine and international regulations and documents on health care. The study's materials were the results of a questionnaire survey of managers and specialists in a pharmacy on marketing contracts. The views of scientists on the above issue were also studied. The study analyzes generalized information from scientific journals using scientific, legal methods. Among the main research methods are systematic approach, analytical, statistical, comparative, dialectical, graphical, and a questionnaire survey of respondents. RESULTS: Results: Consolidation of massive pharmacy chains leads to an artificial increase in drug prices by almost 50 percent, which significantly reduces their availability to patients, and in many cases, makes treatment impossible due to lack of funds. CONCLUSION: Conclusions: As a result of further monopolization of the pharmacy market, the pharmaceutical industry, small pharmacy enterprises, and the complete distribution of medicines will be destroyed.

Dormancy in Metacommunities.

Although metacommunity ecology has improved our understanding of how dispersal affects community structure and dynamics across spatial scales, it has yet to adequately account for dormancy. Dormancy is a reversible state of reduced metabolic activity that enables temporal dispersal within the metacommunity. Dormancy is also a metacommunity-level process because it can covary with spatial dispersal and affect diversity across spatial scales. We develop a framework to integrate dispersal and dormancy, focusing on the covariation they exhibit, to predict how dormancy modifies the importance of species interactions, dispersal, and historical contingencies in metacommunities. We used empirical and modeling approaches to demonstrate the utility of this framework. We examined case studies of microcrustaceans in ephemeral ponds, where dormancy underlies metacommunity dynamics, and identified constraints on the dispersal and dormancy strategies of bromeliad-dwelling invertebrates. Using simulations, we showed that dormancy can alter classic metacommunity patterns of diversity in ways that depend on dispersal-dormancy covariation and spatiotemporal environmental variability. We propose that dormancy may also facilitate evolution-mediated priority effects if locally adapted seed banks prevent colonization by more dispersal-limited species. Last, we present testable predictions for the implications of dormancy in metacommunities, some of which may fundamentally alter our understanding of metacommunity ecology.

A rigorous comparison of sexual selection indexes via simulations of diverse mating systems.

Sexual selection is a cornerstone of evolutionary theory, but measuring it has proved surprisingly difficult and controversial. Various proxy measures--e.g., the Bateman gradient and the opportunity for sexual selection--are widely used in empirical studies. However, we do not know how reliably these measures predict the strength of sexual selection across natural systems, and most perform poorly in theoretical worst-case scenarios. Here we provide a rigorous comparison of eight commonly used indexes of sexual selection. We simulated 500 biologically plausible mating systems, based on the templates of five well-studied species that cover a diverse range of reproductive life histories. We compared putative indexes to the actual strength of premating sexual selection, measured as the strength of selection on a simulated "mating trait." This method sidesteps a key weakness of empirical studies, which lack an appropriate yardstick against which proxy measures can be assessed. Our model predicts that, far from being useless, the best proxy measures reliably track the strength of sexual selection across biologically realistic scenarios. The maximum intensity of precopulatory sexual selection s'max (the Jones index) outperformed all other indexes and was highly correlated with the strength of sexual selection. In contrast, the Bateman gradient and the opportunity for sexual selection were poor predictors of sexual selection, despite their continuing popularity.

Arrival order and release from competition does not explain why haplochromine cichlids radiated in Lake Victoria.

The frequent occurrence of adaptive radiations on oceanic islands and in lakes is often attributed to ecological opportunity resulting from release from competition where arrival order among lineages predicts which lineage radiates. This priority effect occurs when the lineage that arrives first expands its niche breadth and diversifies into a set of ecological specialists with associated monopolization of the resources. Later-arriving species do not experience ecological opportunity and do not radiate. While theoretical support and evidence from microbial experiments for priority effects are strong, empirical evidence in nature is difficult to obtain. Lake Victoria (LV) is home to an exceptional adaptive radiation of haplochromine cichlid fishes, where 20 trophic guilds and several hundred species emerged in just 15 000 years, the age of the modern lake that was preceded by a complete desiccation lasting several thousand years. However, while about 50 other lineages of teleost fish also have established populations in the lake, none of them has produced more than two species and most of them did not speciate at all. Here, we test if the ancestors of the haplochromine radiation indeed arrived prior to the most competent potential competitors, 'tilapias' and cyprinids, both of which have made rapid radiations in other African lakes. We assess LV sediment core intervals from just before the desiccation and just after refilling for the presence of fossil fish teeth. We show that all three lineages were present when modern LV began to fill with water. We conclude that the haplochromines' extraordinary radiation unfolded in the presence of potentially competing lineages and cannot be attributed to a simple priority effect.

Local and regional founder effects in lake zooplankton persist after thousands of years despite high dispersal potential.

We reconstructed the genetic structure of a planktonic crustacean Daphnia longispina living in high mountain lakes and ponds in the Pyrenees to investigate whether it was shaped by persistent founder effects originating shortly after the last glacial maximum or by ongoing dispersal and effective migration (gene flow). We found that the genetic structure can largely be explained by a single colonization event following gradual deglaciation of the Pyrenees ~10 000-15 000 years ago. Nuclear genetic diversity declined steeply from southeast to northwest, suggestive of serial colonization of available habitats with advancing deglaciation. The spatial genetic structure suggests that founder effects were major determinants of the present-day diversity, both at the catchment level and at the level of individual water bodies, further supporting extremely low effective migration rates. This study reveals a prime example of a founder effect that is both long lasting and maintained at small spatial scales. Our data suggest a process of isolation by colonization as a result of strong priority effects and monopolization. We found evidence for the spread of haplotypes with Pyrenean ancestry across the Palaearctic over distances up to 5500 km, although the local genetic structure after colonization was hardly influenced by contemporary dispersal. Finally, our data also suggest that mitochondrial mutation rates in the studied populations were seven times higher than typically assumed. Overall, we show that founder effects can persist for centuries even at small spatial scales at which the potential for dispersal is high.

Drivers of population genetic differentiation in the wild: isolation by dispersal limitation, isolation by adaptation and isolation by colonization.

Empirical population genetic studies have been dominated by a neutralist view, according to which gene flow and drift are the main forces driving population genetic structure in nature. The neutralist view in essence describes a process of isolation by dispersal limitation (IBDL) that generally leads to a pattern of isolation by distance (IBD). Recently, however, conceptual frameworks have been put forward that view local genetic adaptation as an important driver of population genetic structure. Isolation by adaptation (IBA) and monopolization (M) posit that gene flow among natural populations is reduced as a consequence of local genetic adaptation. IBA stresses that effective gene flow is reduced among habitats that show dissimilar ecological characteristics, leading to a pattern of isolation by environment. In monopolization, local genetic adaptation of initial colonizing genotypes results in a reduction in gene flow that fosters the persistence of founder effects. Here, we relate these different processes driving landscape genetic structure to patterns of IBD and isolation by environment (IBE). We propose a method to detect whether IBDL, IBA and M shape genetic differentiation in natural landscapes by studying patterns of variation at neutral and non-neutral markers as well as at ecologically relevant traits. Finally, we reinterpret a representative number of studies from the recent literature by associating patterns to processes and identify patterns associated with local genetic adaptation to be as common as IBDL in structuring regional genetic variation of populations in the wild. Our results point to the importance of quantifying environmental gradients and incorporating ecology in the analysis of population genetics.

Know your enemy: The dragonfly Erythrodiplax fusca (Libellulidae) uses eavesdropping to obtain information about potential rivals.

Animals can eavesdrop on other competitors during territorial defense disputes to better choose rivals weaker than themselves and increase the chances of monopolizing resources. In dragonflies, males often compete for access to water bodies, which attract sexually receptive females to mate and lay eggs. During the breeding season, satellite males can observe fights between territory owners and intruders and, therefore, obtain information about potential rivals through visual cues. Consequently, weaker competitors may face more intense competition than stronger ones when defending a territory. In this study, we conducted field experiments with the dragonfly Erythrodiplax fusca to investigate whether eavesdropping on territorial disputes, using visual cues, affects the intensity of competition that territory owners face. We recorded the number of intruders that engage in disputes against males that recently occupied territories in two groups: the "eavesdropping" group (i.e., individuals with access to rivals' prior information) and the control group (i.e., competitors with no access to prior information). The number of intruders was greater in the eavesdropping group compared to the control group. This effect depended on the interaction between the size of the territory owners and the presence of eavesdropping. The number of intruders decreased with increase in the size of the owners in the presence of eavesdropping, but this relationship did not occur in the control group. We discuss the implications of our findings for the male decision-making process to initiate agonistic disputes and how investigating eavesdropping behavior can improve current models of conflict resolution in animals.

Can outcomes of dyadic interactions be consistent across contexts among wild zebrafish?

Winner-loser relations among group-living individuals are often measured by the levels of aggressive interactions between them. These interactions are typically driven by competition for resources such as food and mates. It has been observed in recent studies on zebrafish that dominant males generally have higher total reproductive success than their less aggressive subordinate counterparts. This study aimed to test whether males who monopolized a food resource (winners) also displayed higher levels of aggression than the males who were unsuccessful (losers). Further, the study also tested whether the same 'winner' males were also able to monopolize interactions with females during courtship. The results from these experiments showed that while males monopolizing food resources (winners) demonstrated higher levels of agonistic interactions than the losers, the average number of courtship interactions initiated by either of the males (i.e. winners/losers) with a female was not significantly different. A significant relationship was obtained between the number of aggressive interactions and feeding latencies of males in the context of food monopolization. This indicated that there could be a linkage between boldness defined by feeding latency in a novel environment and agonistic responses. The probable role of nature of resources, resource availability and distribution in determining the outcomes of dyadic contests is discussed.

Resource defense and monopolization in a marked population of ruby-throated hummingbirds (Archilochus colubris).

Resource defense behavior is often explained by the spatial and temporal distribution of resources. However, factors such as competition, habitat complexity, and individual space use may also affect the capacity of individuals to defend and monopolize resources. Yet, studies frequently focus on one or two factors, overlooking the complexity found in natural settings. Here, we addressed defense and monopolization of nectar feeders in a population of free-ranging ruby-throated hummingbirds marked with passive integrated transponder (PIT tags). Our study system consisted of a 44 ha systematic grid of 45 feeders equipped with PIT tag detectors recording every visit made at feeders. We modeled the number of visits by competitors (NVC) at feeders in response to space use by a focal individual potentially defending a feeder, number of competitors, nectar sucrose concentration, and habitat visibility. Individuals who were more concentrated at certain feeders on a given day and who were more stable in their use of the grid throughout the season gained higher exclusivity in the use of those feeders on that day, especially for males competing against males. The level of spatial concentration at feeders and its negative effect on NVC was, however, highly variable among individuals, suggesting a continuum in resource defense strategies. Although the apparent capacity to defend feeders was not affected by competition or nectar sucrose concentration, the level of monopolization decreased with increasing number of competitors and higher nectar quality. Defense was enhanced by visibility near feeders, but only in forested habitats. The reverse effect of visibility in open habitats was more difficult to interpret as it was probably confounded by perch availability, from which a bird can defend its feeder. Our study is among the first to quantify the joint use of food resource by overlapping individuals unconstrained in their use of space. Our results show the importance of accounting for variation in space use among individuals as it translated into varying levels of defense and monopolization of feeders regardless of food resource distribution.