State‐dependent dynamics of cycles in red grouse abundance

Fluctuating populations are frequently demonstrated to co‐vary in abundance over space, but the dynamics of coupling between populations that gives rise to this synchrony are poorly understood. Synchrony may arise through coupling that is weak and continuous, but in populations that cycle with a characteristic period, synchrony can be maintained through stronger coupling that acts only intermittently. Here, we apply a discrete Markov model that describes the state of a population trajectory to be in one of four possible states. The Markov model reveals the nature of the coupling that gives rise to the weakly synchronous cycles of red grouse abundance. Using time‐series data from 287 populations across the species range in the UK, we show that grouse populations appear mostly uncoupled through time, but that approximately one year in six, “collective forcing events” occur, where populations in a region are forced into synchrony to a significantly greater degree than would be expected if their dynamics proceeded independently. In the absence of these events, synchrony between populations dissipates within ~3 yr. Smaller, low abundance populations tend to make the less probable phase shifts required to synchronize with nearby high abundance populations, suggesting that these low abundance populations are more susceptible to the perturbations responsible for phase shifts than larger populations.     

Recent parallel divergence in body shape and diet source of alewife life history forms

Recent work suggests that juvenile alewives (Alosa pseudoharengus) share similar phenotypes among independently derived landlocked (freshwater resident) populations. Based on this observation, it is possible that the alewife life history forms represent a case of parallel adaptive divergence. To further evaluate this hypothesis, we describe patterns of body shape divergence between anadromous and landlocked alewife life history forms using geometric morphometrics. Our results suggest that body shape differs significantly between juveniles of the alewife forms: anadromous fish were more robust, with larger heads and deeper caudal peduncles, while landlocked fish from three independently isolated populations were more fusiform with thinner caudal peduncles and smaller heads. These differences matched population level dietary patterns, which suggest that anadromous fish consumed more littoral resources than landlocked fish. Finding consistent differences across populations of the same form supports the notion that landlocked alewives have diverged from their anadromous ancestors in a parallel manner, in response to pressures associated with being isolated in freshwater lakes. Comparing alewife phenotypes to expectations from the literature suggests that neither migration distance of the population, nor the relative availability of habitats in each lake, are likely drivers of the pattern we report. Instead, the pattern is consistent with the hypothesis that divergence between alewife forms results from the distinct effects of each form on its zooplankton prey.     

A Model of Functional Epistasis and Linkage Disequilibrium in Populations with Overlapping Generations

A model of functional epistasis is proposed in which it is assumed that coupling and repulsion genotypes differ in metabolic efficiency and thus in development time and net fecundity. The implications of this model are investigated for iteroparous populations with fluctuating rates of increase. It is found that the fluctuations in rate of increase can lead to large fluctuations in gamete frequency and D, the coefficient of linkage disequilibrium, but that D will almost always have a value of zero at some point during the populations' demographic cycle. Some of the model populations would be expected to be in a state of linkage disequilibrium only fleetingly: others would exhibit D-cycles interpretable as random fluctuation. Implications of the model for interpretations of existing data on linkage disequilibrium among enzyme loci in Drosophila are discussed.     

Frequency-dependent selection on aposematic prey: some experiments

Experiments to measure the strength of frequency-dependent selection on dimorphic populations of distasteful, brightly coloured prey were carried out, using pastry cylinders as prey and wild birds as predators. When the two forms were haphazardly intermingled, the birds took a relative excess of whichever form happened to be the rarer if all prey in the population were distasteful but selected the forms independently of frequency if they were all neutral in flavour. They did not take an excess of the rarer form from populations of distasteful prey when that form was distributed in a single clump rather than evenly dispersed among the commoner form. This confirms the expectation that rarer forms of aposematic animals are at a frequency-dependent disadvantage when they are intermingled with commoner forms but not when they occur in clusters.     

Trade‐offs in osmoregulation and parallel shifts in molecular function follow ecological transitions to freshwater in the Alewife

Adaptation to freshwater may be expected to reduce performance in seawater because these environments represent opposing selective regimes. We tested for such a trade‐off in populations of the Alewife (Alosa pseudoharengus). Alewives are ancestrally anadromous, and multiple populations have been independently restricted to freshwater (landlocked). We conducted salinity challenge experiments, whereby juvenile Alewives from one anadromous and multiple landlocked populations were exposed to freshwater and seawater on acute and acclimation timescales. In response to acute salinity challenge trials, independently derived landlocked populations varied in the degree to which seawater tolerance has been lost. In laboratory‐acclimation experiments, landlocked Alewives exhibited improved freshwater tolerance, which was correlated with reductions in seawater tolerance and hypo‐osmotic balance, suggesting that trade‐offs in osmoregulation may be associated with local adaptation to freshwater. We detected differentiation between life‐history forms in the expression of an ion‐uptake gene (NHE3), and in gill Na⁺/K⁺‐ATPase activity. Trade‐offs in osmoregulation, therefore, may be mediated by differentiation in ion‐uptake and salt‐secreting pathways.     

Phylogenetic affinities of an enigmatic nannoplankton,Braarudosphaera bigelowii based on the SSU rDNA sequences

The phylogenetic position of an enigmatic calcareous nannoplankton Braarudosphaera bigelowii, whose taxonomic position has long been questioned, was investigated using molecular genetic methods. The SSU rDNA sequences of B. bigelowii were obtained using a single cell PCR technique independently from two single cells isolated from field samples. A BLAST search revealed that the SSU rDNA of B. bigelowii was most similar to those of the division Haptophyta. Subsequent phylogenetic analyses showed that B. bigelowii was included in a clade comprising members of the orders Isochrysidales and Coccolithales, the genus Chrysoculter and two unidentified haptophytes. Morphometric measurements of extant B. bigelowii populations were made to obtain precise information on their intraspecific size variation. The results showed that these populations could be subdivided into a small form (< 2.4 μm), an intermediate form A (4.0–5.3 μm), and an intermediate form B (5.3–7.2 μm), although the borders of the latter two forms overlapped. The SSU rDNA sequences showed the presence of 10-bp substitutions plus six insertions/deletions between specimens of intermediate forms A and B. The levels of DNA variation between them suggested that these two forms are genetically distinct from each other.     

Multiple differences in calling songs and other traits between solitary and gregarious Mormon crickets from allopatric mtDNA clades

Background  

In acoustic species, traits such as male calling song are likely to diverge quickly between allopatric populations due to sexual selection, and divergence in parameters such as carrier frequency, chirp structure, and other important song characters can influence sexual isolation. Here we make use of two forms of Mormon crickets to examine differences in a broad suite of traits that have the potential to influence speciation via sexual isolation. Mormon crickets in "gregarious" populations aggregate into dense migratory bands, and females are the sexually competitive sex (sex-role reversal). There is also a non-outbreak "solitary" form. These two forms are largely but not perfectly correlated with a significant mtDNA subdivision within the species that is thought to have arisen in allopatry. Combined information about multiple, independently evolving traits, such as morphology and structural and behavioural differences in calling song, provides greater resolution of the overall differences between these allopatric populations, and allows us to assess their stage of divergence. We test two predictions, first that the forms differ in song and second that gregarious males are more reluctant to sing than solitary males due to sex role reversal. We also tested for a difference in the relationship between the size of the forewing resonator, the mirror, and carrier frequency, as most models of sound production in crickets indicate that mirror size should predict carrier frequency.     

Plastid DNA haplotype variation in Dactylorhiza incarnata (Orchidaceae): evidence for multiple independent colonization events into Scandinavia

The early marsh orchid, Dactylorhiza incarnata (L.) Soó s. l., grows in medium-rich to rich fens and marshes over much of Europe and parts of Asia. The species is highly polymorphic and different forms may grow together at the same site. In the present study, I tested the hypothesis that these forms represent different migrant populations that have colonized Scandinavia independently of each other, possibly from different source areas. Accessions from Scandinavia and elsewhere were screened for variation at three size-variable plastid marker loci, one polyA repeat, one polyA-polyTA-polyT repeat and one 9 bp indel. Ten haplotypes were defined on basis on the combined variation pattern. The common occurrence of several haplotypes in southern Scandinavia and adjacent areas to the south and the east of the Baltic Sea suggests that D. incarnata has been dispersed on repeated occasions across the Baltic. Also, there was some correlation between haplotype composition and morphological form on the island of Gotland, in agreement with the independent colonization hypothesis. Material from northernmost Sweden, Finland and northwest Russia was fixed for a single widespread haplotype, indicating that populations in this area are located farther away from the Pleistocene refugia. Dactylorhiza incarnata ssp. lobelii from southwest Norway was characterized by a haplotype that was not encountered elsewhere in Scandinavia. Given its proximity to British populations dominated by the same haplotype, it is suggested that D. incarnata ssp. lobelii was established independently of the other Scandinavian populations, from coastal refugia located in western Europe.     

Diversification, Sympatric Speciation, and Trophic Polymorphism of Arctic Charr, Salvelinus Alpinus Complex, in Transbaikalia

In northern Transbaikalia, independently evolving landlocked populations of Arctic charr are found in mountain lakes. To assess the diversity of charr in this region, speciation modes involved in the evolution of charr forms, and the role of trophic polymorphism in their divergence, we studied the morphology and feeding of dwarf, small, and large forms of Arctic charr from a number of Transbaikalian lakes. Meristic data on charr from five lakes support the earlier conclusion that the three forms do not represent separate lineages but have independently diverged in sympatry in each of the lakes. In 10 lakes, the dwarf form showed varying degrees of differentiation from normal (small and large) charr in meristic characters (up to morphologically distinct and presumably reproductively isolated groupings), which is viewed as various levels of sympatric divergence. Indexes of gill raker length in fish from 20 lakes vary among populations of both dwarf and normal charr, with forms having short and long rakers being sympatric in some of these lakes. However, the index can be used only for comparing charr of different forms up to about 32cm fork length (FL) because it is strongly negatively correlated with size in larger fish. The study of charr diets in 21 lakes indicates that large charr are piscivorous whereas dwarf and small charr feed on a wide range of invertebrates, partitioning these resources in different ways. Planktivores, including very specialized ones, and non-planktivores (benthic feeders, insectivores) can be identified within the small and dwarf forms. The proportion of plankton in the diets of dwarf and small charr is positively correlated with the number and length of gill rakers while the proportion of benthos is negatively correlated. Allopatric planktivorous and non-planktivorous small charr differ in body proportions; parallel emergence of such morphotypes in different parts of the range is a characteristic feature of the Salvelinus alpinus complex.     

Coupled oscillations in food webs: balancing competition and mutualism in simple ecological models

As in other oscillating systems, oscillations of consumer resource pairs in ecological systems may be coupled such that complex behavior results. The form of that coupling may determine the nature and extent of this behavior. Two biologically significant forms of coupling are here investigated: first, where consumers consume each other's resources (CR coupling, representing competition between the two consumers), and second, where the resources are in competition with one another (RR coupling, potentially representing indirect mutualism between the two consumers). Interestingly, CR coupling leads to in-phase synchrony of the oscillations, whereas RR coupling leads to antiphase synchrony. With either form of coupling, if the coupling remains weak, synchronous behavior is generated in the two systems. At strong levels of coupling, when the two forms act simultaneously, a balance between competition and mutualism is generated, which is manifest differently at different levels of resource coupling.     

Spatio-temporal dynamics of the grey-sided vole in Hokkaido: identifying coupling using state-based Markov-chain modelling

Explaining synchronization of cyclical or fluctuating populations over geographical regions presents ecologists with novel analytical challenges. We have developed a method to measure synchrony within spatial-temporal datasets of population densities applicable to both periodic and irregularly fluctuating populations. The dynamics of each constituent population is represented by a discrete Markov model. The state of a population trajectory at each time-point is classified as one of 'increase', 'decrease', 'peak' or 'trough'. The set of populations at any time-point is characterized by the frequency distribution of these different states, and the time-evolution of this frequency distribution used to test the hypothesis that the dynamics of each population proceeds independently of the others. The analysis identifies years in which population coupling results in synchronous states and onto which states the system converges, and identifies those years in which synchrony remains high but is accounted for by coupling observed in previous years. It also enables identification of which pairs of sites show the highest levels of coupling. Applying these methods to populations of the grey-sided vole on Hokkaido reveals them to be fluctuating in greater synchrony than would be expected from independent dynamics, and that this level of synchrony is maintained through intermittent coupling acting in ca. 1 year in four or five. High synchrony occurs between sites with similar vegetation and of similar altitude indicating that coupling may be mediated through shared environmental stimuli. When coupling is indicated, convergence is equally likely to occur on a peak state as a trough, indicating that synchronization may be brought about by the response of populations to a combination of different stimuli rather than by the action of any single process.     

Taxonomic significance of seed sculpture and pollen ecto-mycoflora at the infraspecific level: Brassica tournefortii case study

Several populations of Brassica tournefortii (Brassicaceae) occurring in Egypt are investigated from the micromorphological point of view (seed ornamentations). The species is known to show a notable phenotypic plasticity and five morphotypes was identified in the past. Furthermore, a soil analysis as well as a study of the fungal species from anthers were carried out. The aim of the study is to verify the taxonomic value of the morpholotypes of B. tournefortii and their ecologic relationship with soil variables. The results obtained demonstrated that the five morphological forms can be distinguished based on the seed sculpture. The Canonical Correspondence Analysis (CCA) exhibited a clear correlation between the soil variables and the identified forms. Six species of fungi were detected from the ecto-anthers in the Forms (F2-F4), while F1 was lacking the fungal species. The study revealed that the morphological plasticity of studied B. tournefortii depends on ecological factors.     

Relationship and genetic structure among autoecious and heteroecious populations of Cronartium pini in northern Fennoscandia

Epidemics of Scots pine blister rust, caused by Cronartium pini, have become an increasing problem in northern Finland and Sweden. The biology of the rust fungus is complex, with two different life cycle forms that cannot be morphologically distinguished, and it is unclear to what extent the two forms contribute to the epidemics. Genetic structure of fourteen populations of C. pini were investigated in Fennoscandia. Distinction between the two life cycle forms, a heteroecious and an autoecious one, was made by determining zygosity using microsatellite markers, and AFLP markers were developed to analyse population genetic relationships. The results showed that the two life cycle forms are clearly differentiated and occur in separate populations. Within the life cycle forms, geographic differentiation was evident, probably due to restricted gene flow as well as connection with different alternating hosts. The host-alternating form dominated in the epidemic regions in northern Fennoscandia. Implications for silvicultural practices are discussed.     

How noise and coupling influence leading indicators of population extinction in a spatially extended ecological system

Anticipating critical transitions in spatially extended systems is a key topic of interest to ecologists. Gradually declining metapopulations are an important example of a spatially extended biological system that may exhibit a critical transition. Theory for spatially extended systems approaching extinction that accounts for environmental stochasticity and coupling is currently lacking. Here, we develop spatially implicit two-patch models with additive and multiplicative forms of environmental stochasticity that are slowly forced through population collapse, through changing environmental conditions. We derive patch-specific expressions for candidate indicators of extinction and test their performance via a simulation study. Coupling and spatial heterogeneities decrease the magnitude of the proposed indicators in coupled populations relative to isolated populations, and the noise regime and the degree of coupling together determine trends in summary statistics. This theory may be readily applied to other spatially extended ecological systems, such as coupled infectious disease systems on the verge of elimination.     

Parallel evolution of the melanic form of the California legless lizard, Anniella pulchra, inferred from mitochondrial DNA sequence variation

Abstract.— The phylogenetic relationships among populations of the fossorial California legless lizard, Anniella pulchra , were examined by sequencing a 990-bp region of the mitochondrial cytochrome b gene. The mitochondrial DNA gene tree was then compared with the geographic distributions of two currently recognized subspecies: A. p. nigra , a melanic form restricted to two disjunct coastal populations, and A. p. pulchra , a more widely distributed, silvery form. We tested the null hypothesis that all A. p. nigra form a clade that is monophyletic with respect to A. p. pulchra . Our results strongly reject the monophyletic origin of the melanic forms and suggest that the two populations of the nominal subspecies A. p. nigra may have arisen independently from different ancestral populations in a parallel evolutionary response to selection in cool, coastal habitats.     

Stability and pattern formation for competing populations with asymmetric nonlocal coupling

We consider a model of two competing species with asymmetric nonlocal coupling in a competition for resources. The nonlocal coupling is via convolution integrals and the asymmetry is via convolution kernel functions which are not even functions of their arguments. The nonlocality is due to species mobility, so that at any fixed point in space the competition for resources depends not just on the populations at that point but on a suitably weighted average of the populations. We introduce two parameters, δ  , describing the extent of the coupling, with δ=0$\delta =0$ corresponding to local coupling, and α  , describing the extent of the asymmetry, with α=0$\alpha =0$ corresponding to symmetric nonlocal interactions. We consider the case where the model admits a stable coexistence equilibrium solution. We perform a linear stability analysis and show that this solution can be destabilized by sufficient nonlocality, i.e., when δ increases beyond a critical value. We consider two specific kernel functions, (i) an asymmetric Gaussian and (ii) an asymmetric stepfunction. We compute the stability boundary as a function of α, and for δ beyond the stability boundary we determine unstable wavenumber bands. We compute nonlinear patterns for δ significantly beyond the stability boundary. Patterns consist of arrays of islands, regions of nonzero population, separated by either near-deadzones where the populations are small, but nonzero, or by deadzones where populations are exponentially small and essentially extinct. We find solutions consisting of propagating traveling waves of islands, solutions exhibiting colony formation, where a colony is formed just ahead of an island and eventually grows as the parent island decays, and modulated traveling waves, where competition between the two species allows propagation and inhibits colony formation. We explain colony formation and the modulated traveling waves as due to a positive feedback mechanism associated with small variations in the amplitude of the parent island.     

Population Variation in the Life History of a Land Fish,Alticus arnoldorum,and the Effects of Predation and Density

Life history variation can often reflect differences in age-specific mortality within populations, with the general expectation that reproduction should be shifted away from ages experiencing increased mortality. Investigators of life history in vertebrates frequently focus on the impact of predation, but there is increasing evidence that predation may have unexpected impacts on population density that in turn prompt unexpected changes in life history. There are also other reasons why density might impact life history independently of predation or mortality more generally. We investigated the consequences of predation and density on life history variation among populations of the Pacific leaping blenny, Alticus arnoldorum. This fish from the island of Guam spends its adult life out of the water on rocks in the splash zone, where it is vulnerable to predation and can be expected to be sensitive to changes in population density that impact resource availability. We found populations invested more in reproduction as predation decreased, while growth rate varied primarily in response to population density. These differences in life history among populations are likely plastic given the extensive gene flow among populations revealed by a previous study. The influence of predation and density on life history was unlikely to have operated independently of each other, with predation rate tending to be associated with reduced population densities. Taken together, our results suggest predation and density can have complex influences on life history, and that plastic life history traits could allow populations to persist in new or rapidly changing environments.     

The origin of a Robertsonian chromosomal translocation in house mice inferred from linked microsatellite markers

The western European house mouse, Mus domesticus, includes many distinct Robertsonian (Rb) chromosomal races. Two competing hypotheses may explain the distribution of Rb translocations found in different populations: they may have arisen independently multiple times, or they may have arisen once and been spread through long-distance dispersal. We investigated the origin of the Rb 5.15 translocation using six microsatellite loci linked to the centromeres of chromosomes 5 and 15 in 84 individuals from three Rb populations and four neighboring standard-karyotype populations. Microsatellite variation on the 5.15 metacentric chromosomes was significantly reduced relative to the amount of variation found on acrocentric chromosomes 5 and 15, suggesting that linked microsatellite loci can track specific mutational events. Phylogenetic analyses resulted in trees which are consistent with multiple origins of the 5.15 metacentric chromosomes found in the three Rb populations. These results suggest that cytologically indistinguishable mutations have arisen independently in natural populations of house mice.     

EVIDENCE FOR MULTIPLE ORIGINS AND SYMPATRIC DIVERGENCE OF TROPHIC ECOTYPES OF SMELT (OSMERUS) IN NORTHEASTERN NORTH AMERICA

The rainbow smelt, Osmerus mordax (Mitchill), is an osmerid fish that exhibits extensive life-history diversity throughout watersheds of northeastern North America. There are both ???sea-run (anadromous) and lake-resident (lacustrine) populations and the latter have diversified further into “dwarf-” and “normal-sized” life-history types. Anadromous and lacustrine smelt may inhabit the same watershed and there are several instances where dwarf and normal populations reside within the same lake. We assayed variation among smelt for morphological traits linked to feeding performance in fishes to see if trophic ecology might promote life-history diversity in Osmerus. We also examined mitochondrial DNA (mtDNA) restriction site variation among forms to assess their evolutionary interrelationships. Dwarf smelt had significantly more gill rakers, larger eyes, but shorter upper jaws than normal lake and anadromous smelt. The populations clustered into two trophic “morphotypes”; an anadromous/normal lake group of populations and a group consisting only of dwarf smelt. The mtDNAs of 444 smelt from 16 populations were digested with 12 restriction enzymes revealing 93 composite mtDNA genotypes that clustered (UPGMA) into two major phylogenetic groups differing by 0.78% in sequence. Both genetic groups were present in dwarf and normal smelt as well as in anadromous fish. Further, geographic proximity, rather than trophic morphotype, appeared to be the major determinant of genetic affinities among populations. In two lakes, however, dwarf and normal smelt populations had significantly different mtDNA genotype frequency distributions indicating that the forms are reproductively isolated within both lakes. A clustering analysis of population affinities suggested that the divergence of sympatric dwarf and normal populations had occurred independently in the two lakes. We concluded (1) that trophic ecology is an important factor promoting differentiation in smelt life histories; (2) that smelt ecotypes are polyphyletic and there have been multiple, independent divergences of Osmerus life-history types throughout northeastern North America; and (3) that the biological and mtDNA differences between coexisting dwarf and normal lake smelt argue strongly that their genetic isolation may have developed sympatrically.