Density-dependent selection closes an eco-evolutionary feedback loop in the stick insect Timema cristinae

Empirical demonstrations of feedbacks between ecology and evolution are rare. Here, we used a field experiment to test the hypothesis that avian predators impose density-dependent selection (DDS) on Timema cristinae stick insects. We transplanted wild-caught T. cristinae to wild bushes at 50 : 50 cryptic : conspicuous morph ratio and manipulated density by transplanting either 24 or 48 individuals. The frequency of the conspicuous morph was reduced by 73% in the low-density treatment, but only by 50% in the high-density treatment, supporting a hypothesis of negative DDS. Coupled with previous studies on T. cristinae, which demonstrate that maladaptive gene flow reduces population density, we support an eco-evolutionary feedback loop in this system. Furthermore, our results support the hypothesis that predator satiation is the mechanism driving DDS. We found no effects of T. cristinae density on the abundance or species richness of other arthropods. Eco-evolutionary feedbacks, driven by processes like DDS, can have implications for adaptive divergence and speciation.     

Correlated responses in death-feigning behavior,activity, and brain biogenic amine expression in red flour beetle <Emphasis Type="Italic">Tribolium castaneum</Emphasis> strains selected for walking distance

Dispersal ability may influence antipredator and mating strategies. A previous study showed a trade-off between predation avoidance and mating success in strains of the red flour beetle Tribolium castaneum selected for walking distance. Specifically, beetles derived from strains selected for longer walking distance suffered higher predation pressure and had higher male mating success than their counterparts derived from strains selected for shorter walking distance. In the study reported here, we compared the locomotor activity, biogenic amine expression in the brain, and death-feigning behavior of the red flour beetle strains selected for walking distance. The results indicated that individuals genetically predisposed to longer walking distance had higher locomotor activity and lower intensity of death-feigning behavior than those genetically predisposed to shorter walking distance. However, no significant differences were found in the expression of biogenic amines in the brain among strains selected for walking distance, although the level of dopamine in the brain differed from that of the strains divergently selected for duration of death-feigning behavior. The relationships between walking speed, activity, death-feigning behavior, and brain biogenic amines in T. castaneum are discussed.     

Is death-feigning adaptive? Heritable variation in fitness difference of death-feigning behaviour.

The adaptation of death-feigning (thanatosis), a subject that has been overlooked in evolutionary biology, was inferred in a model prey-and-predator system. We studied phenotypic variation among individuals, fitness differences, and the inheritance of death-feigning behaviour in the red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae). Two-way artificial selections for the duration of death-feigning, over 10 generations, showed a clear direct response in the trait and a correlated response in the frequency of death-feigning, thus indicating variation and inheritance of death-feigning behaviour. A comparison of the two selected strains with divergent frequencies of death-feigning showed a significant difference in the fitness for survival when a model predator, a female Adanson jumper spider, Hasarius adansoni Audouin (Araneomophae: Salticidae), was presented to the beetles. The frequency of predation was lower among beetles from strains selected for long-duration than among those for short-duration death-feigning. The results indicate the possibility of the evolution of death-feigning under natural selection.     

Predator-induced plasticity does not alter the pathway from evolution to ecology among locally adapted populations of <Emphasis Type="Italic">Daphnia</Emphasis>

In the growing field of eco-evolutionary dynamics, evidence for an influence of rapid shifts in phenotype on ecological processes is accumulating, yet, the contributions of phenotypic plasticity versus genetic change to these observed ecological changes are unclear. In one of the best studied ecosystems in terms of eco-evolutionary dynamics, landlocked versus anadromous alewife (Alosa pseudoharengus) have caused strong evolutionary divergence in their key zooplankton prey (Daphnia ambigua). We previously showed that such evolutionary differences have cascading ecological effects on consumer-resource dynamics and primary production. Yet, these locally adapted populations of Daphnia also differ in trait plasticity, which may, in turn, modify the pathway from evolution to ecology. Here we compared Daphnia from lakes with landlocked versus anadromous alewife for differences in rates of population growth in the presence and absence of predator cues over the course of a 39-day experiment. We predicted that predator-induced shifts in life history traits would facilitate faster rates of population growth. Contrary to our expectations, predator cue exposure did not alter rates of population growth. We instead found that Daphnia from lakes with landlocked alewife ultimately attained higher population densities (and exhibited faster population growth) when compared with Daphnia from lakes with anadromous alewife. Based on our previous work, these population level responses were unexpected, as Daphnia from lakes with landlocked alewife exhibit slower rates of somatic growth and delayed maturation. We discuss our results in lieu of the known differences in plasticity and how the population growth patterns may be influenced by resource limitation.     

Negative relationship between ambient temperature and death-feigning intensity in adult Callosobruchus maculatus and Callosobruchus chinensis

Abstract.  Although the effects of temperature on insect behaviours are studied frequently, few studies report on the relationship between temperature and anti-predator behaviours. A negative relationship between ambient temperature and the intensity of death-feigning is found in adults of two seed beetle species, Callosobruchus maculatus (F.) and C. chinensis (L.) (Coleoptera: Bruchidae). Two traits representing the intensity of immobility, the frequency and the duration of death-feigning, are measured at different temperatures. Almost all adults feign death at 15 °C, but the frequency of death-feigning decreases at higher temperatures in C. maculatus , whereas all C. chinensis adults show this behaviour at 15 and 20 °C and almost all show it at 25 °C, but the frequency of death-feigning decreases at 30 and 35 °C. The difference between the two species might be due to the specific strain of each species used in the experiment. The duration of death-feigning is correlated negatively with the increase in ambient temperature in both species. The frequency at which adults feigned death is higher in females than in males in both species, but the duration of death-feigning is higher in females than in males only in C. maculatus . The relationships between temperature and death-feigning behaviours are discussed from physiological and ecological viewpoints.     

Research on meiotic chromosome pairing in <Emphasis Type="Italic">Roegneria sinica</Emphasis> var. <Emphasis Type="Italic">media</Emphasis> evaluated using genomic in situ hybridization

The analysis of chromosome pairing during meiosis is important for understanding the relationships between different genomes. To evaluate the diversity of chromosome pairing behavior in the wild species of Roegneria sinica var. media Keng with St and H genomes in Triticeae (Poaceae), differences and similarities in the meiotic chromosome pairing behaviors of the two genomes in two populations of R. sinica var. media, were analyzed using genomic in situ hybridization. Chromosome pairing at meiotic metaphase I in the two populations of R. sinica var. media mainly formed bivalents, although several univalents, trivalents and quadrivalents also occurred. Chromosome pairings occurred mainly between homologous chromosomes. However, some non-homologous pairings were observed under natural conditions. No significant differences in karyotype were found between the St and H genomes. Chromosome pairing behaviors differed between and within the two populations. Genetic variation occurred mainly within populations (94.04 %), and variation was more abundant in one population than the other. The genomes St and H differed, but there was some relationship between the two genomes. These findings suggest that homoeologous pairing of chromosomes or exchanges occurred between different genomes of the wild species in Triticeae during evolution. The findings also provide conclusive cytological evidence for genetic variation within the wild species, which forms the basis of their genetic diversity.     

Eco-evolutionary dynamics in herbivorous insect communities mediated by induced plant responses

It is increasingly recognized that the ecology of communities and evolution of species within communities are interdependent, and researchers have been paying attention to this rapidly emerging field of research, i.e., through studies on eco-evolutionary dynamics. Most of the studies on eco-evolutionary dynamics have been concerned with direct trophic interactions. However, community ecologists have shown that trait-mediated indirect effects play an important role in shaping the structure of natural communities. In particular, in terrestrial plant–insect systems, indirect effects mediated through herbivore-induced plant responses are common and have a great impact on the structure of herbivore communities. This review describes eco-evolutionary dynamics in herbivorous insect communities, and specifically focuses on the key role of herbivore-induced plant responses in eco-evolutionary dynamics. First, I review studies on the evolution of herbivore traits relevant to plant induction and discuss evolution in a community context mediated by induced plant responses. Second, I highlight how intraspecific genetic variation or evolution in herbivore traits can influence herbivore community structure. Finally, I propose the hypothetical model that induced plant responses supports eco-evolutionary feedback in herbivore communities. In this review, I argue that the application of the indirect interaction web approaches into studies on eco-evolutionary will provide profound insights into understanding of mechanisms of the generation and maintenance of biodiversity.     

Aphids Playing Possum – Defensive or Mutualistic Response?

Paper presents the phenomenon of thanatosis or death-feigning in selected aphids species. This specific reaction was predominantly analysed on the example of aphid subfamily Lachninae. Individuals of this group were used in experiments, during which a thanatotic response was induced with various results. The response differed from prolonged thanatosis, lasting for several minutes (Eulachnus rileyi), through shrinking behaviour (e. g. in Stomaphis graffii) to non-responsive species such as Cinara (Schizolachnus) pineti. The large interspecies variation of observed responses can be linked to other defensive mechanisms existing in the studied species, as well as to their mutualistic relationship with ants. The behaviour of shrinking is hypothesized to be the mutualistic response, developed from thanatosis, and being adapted to transportation by ant workers.     

Drop or fly? Negative genetic correlation between death-feigning intensity and flying ability as alternative anti-predator strategies

A prey animal may have the alternative of flying away or feigning death when it encounters predators. These alternatives have a genetic base as anti-predator strategies in the adzuki bean beetle, Callosobruchus chinensis. A negative genetic correlation between death-feigning intensity and flying ability was found in C. chinensis, i.e. lower flying ability is genetically connected to escaping by dropping from a perch and then feigning death, whereas higher flying ability does not correspond to death-feigning behaviour. Two bidirectional artificial selections for death-feigning duration and flying ability were conducted independently in C. chinensis. The strains selected for shorter (longer) duration of death-feigning had higher (lower) flying ability, while the strains selected for lower (higher) flying ability showed longer (shorter) duration of death-feigning. When the two traits were compared in 21 populations of C. chinensis derived from different geographical regions, a significant negative correlation was found between death-feigning intensity and flying ability. Based on these results, the choice between alternative escaping behaviours in animals is discussed from two points of view: phenotypic plasticity, an individual with two tactics; and pleiotropic genetic correlation, different individuals with opposite strategies.     

Responses to relaxed and reverse selection in strains artificially selected for duration of death-feigning behavior in the red flour beetle, <Emphasis Type="Italic">Tribolium castaneum</Emphasis>

Divergent lines selected artificially for many generations make it possible to answer two questions: (1) whether genetic variation still exists within the selected population; and (2) whether the selection itself is costly for the selected strain. In previous studies, the red flour beetle Tribolium castaneum was divergently selected artificially for duration of death-feigning, and strains selected for longer (L-strain) and shorter (S-strain) durations of death-feigning have been established (Miyatake et al. 2004, 2008). Because the selection experiments have been conducted for more than 27 generations, genetic variation may be eroded. Furthermore, because another previous study reported physiological costs to L-strains, the L-strains selected artificially for longer duration of death-feigning may have suffered more costs than the S-strains. In the present study, therefore, we relaxed the selection pressure after the 27th or 30th generation of S- and L-strains. We also carried out reverse selection during the most recent eight generations of S- and L-strains. The results showed that each strain clearly responded to relaxation of selection and reverse selection, suggesting that (1) additive genetic variation still existed in both strains after long-term selection, and (2) selection for shorter and longer duration of death-feigning was costly. These results suggest that anti-predator behavior is controlled by many loci, and longer or shorter duration of death-feigning is costly in a laboratory without predators.     

Population consequences of mutational events: effects of antibiotic resistance on the <Emphasis Type="Italic">r</Emphasis>/<Emphasis Type="Italic">K</Emphasis> trade-off

What are the effects of a mutational event on population dynamics? This eco-evolutionary question has relevance not only to basic biological theories but also to conservation applications. We evaluated the relationship between maximum population growth rate (r _max) and carrying capacity (K) among strains of the bacterium Pseudomonas fluorescens. Each of 65 strains differed from their common ancestor by one naturally acquired phenotypic change conferring antibiotic resistance, brought about by a single mutational event, and each was grown in isolation in four environments. We found no evidence of a trade-off between r _max and K. Rather, strains with rapid growth rates also had high carrying capacity, with little interaction between strain and environment. We conclude that the extensive variation in overall fitness resulting from single mutational events likely masks whatever population trade-offs may exist.     

Song frequency correlates with latitude and individual body size in the cicada <Emphasis Type="Italic">Mogannia formosana</Emphasis> Matsumura (Hemiptera: Cicadidae)

Analyses of acoustic variation between and within populations can help to clarify the evolution and diversification of cicada calling songs. In this study, we analyzed the acoustic variation in the calling song of Mogannia formosana within Taiwan and between Taiwan and Green Island to assess the effects of geographic locations. Furthermore, chorusing males in the Green Island population were recorded and collected from the same habitat site during the same time period to investigate the relationship between individual body size and the acoustic features of calling songs. Among populations of M. formosana, we found that most of the acoustic variation in M. formosana calling songs was associated with frequency parameters, in which six frequency parameters changed significantly with latitude on the island of Taiwan. In contrast, temporal parameters, which were associated with principal components corresponding to less acoustic variation than that of the first principal component, were also found to be significant among populations but did not show consistent trends of difference with latitude. However, the geographically isolated Green Island population exhibited the lowest number of short echemes in segment B, which is the diagnostic structure employed to separate M. formosana from other M. species. This finding suggests that the temporal pattern of segment B in the calling songs of M. formosana might be important for both population differentiation and interspecific recognition. In a chorus of the Green Island population, the sound frequency of the last short echeme was found to be significantly correlated with individual body size. The possible role played by sexual selection in shaping sound frequency as a result of its reliable association with body size was discussed. We suggest that, in comparison with temporal elements, the frequency elements of segment B in calling songs of M. formosana in chorus can serve as a more reliable indicator of body size for female mate choice.     

Differential Rates of Male Genital Evolution in Sibling Species of <Emphasis Type="Italic">Drosophila</Emphasis>

Genital morphology in animals with internal fertilization is considered to be among the fastest evolving traits. Sexual selection is often proposed as the main driver of genital diversification but the exact selection mechanisms involved are usually unclear. In addition, the mechanisms operating may differ even between pairs of sibling species. We investigated patterns of male genital variation within and between natural populations of the cactophilic fly Drosophila koepferae ranging its entire geographic distribution and compared them with those previously observed in its sibling species, D. buzzatii. Using both mtDNA and nDNA markers we found that genital shape variation in D. koepferae is more restricted than expected for neutral evolution, suggesting the predominance of stabilizing selection. We also detected dissimilar patterns of divergence between populations of D. koepferae that were allopatric and sympatric with D. buzzatii. The constrained evolution inferred for D. koepferae’s genitalia clearly contrasts with the rapid divergence and higher morphological disparity observed in the populations of D. buzzatii. Finally, different possible scenarios of male genital evolution in each species and within the radiation of D. buzzatii cluster are discussed.     

Nuclear microsatellite markers reveal the low genetic structure of <Emphasis Type="Italic">Pinus mugo</Emphasis> Turra (dwarf mountain pine) populations in Europe

Twenty-one populations (555 individuals) covering the entire native range of Pinus mugo Turra (dwarf mountain pine) were investigated for genetic variation scored at 13 nuclear microsatellite markers (nSSRs). The main objective of the present study was to determine the genetic structure across the present distribution of the species and locate populations of different genetic compositions. Most of the genetic variation was observed within the populations (95%). The assignment of populations based on Bayesian clustering methods revealed that the Sudeten populations of P. mugo form a separate genetic cluster. These stands have likely been established through the founder effects of Alpine migrants. The distribution and level of SSR polymorphisms, along with no evidence of isolation by distance or phylogeographic structure, indicate that the present populations of P. mugo have diverged relatively recently and originate from a larger glacial distribution of the species. One peripheral stand from Italy had the lowest values of most calculated genetic variation indices. This stand could therefore be more susceptible to genetic drift and a negative impact of predicted environmental changes. We discuss our findings with respect to previously published results on the genetic and morphological variation of P. mugo and with consideration for the conservation genetics of the species.     

Ecological and molecular genetic features of <Emphasis Type="Italic">Ixodes persulcatus</Emphasis> and <Emphasis Type="Italic">I. pavlovskyi</Emphasis> ticks in Southern West Siberia

Ixodiae ticks have been counted and sampled in six distant locations within the Altai-Sayan Mountain Area and northern forest steppe in 2012. In total, 1829 individuals of I. persulcatus and 719 of I. pavlovskyi ticks were collected by flagging. I. persulcatus is found in all six examined locations and I. pavlovskyi is found in Western Altai, Northeastern Altai, and Northern Altai, as well as in forest parks near Akademgorodok of Novosibirsk. In total, 16S rRNA and COI sequences have been determined in 144 and 164 I. persulcatus individuals, as well as in 60 and 55 I. pavlovskyi. Selectively equivalent mutant forms are present in I. pavlovskyi populations of Northeastern Altai and Northern Altai. The hydrothermal regime in the forest parks near Novosibirsk is optimal for the I. pavlovskyi ticks. These ticks have recently colonized the forest parks near Novosibirsk and therefore are genetically close to I. pavloskyi from the Altai–Sayan Mountain Area populations.     

Morphological variation of <Emphasis Type="Italic">Melanargia russiae</Emphasis> (Esper, 1783) (Lepidoptera,Satyridae) from the main part of the range and in case of its expansion to the north under climate change conditions

A climate-related shift in the range boundaries of the western–central Eurasian subboreal species Melanargia russiae in the Ural region from the northern forest-steppe zone to pine–birch forests was found. Morphological variation of M. russiae wings from the boundary northern populations and populations from the main part of the range in the Urals were studied. The results of a complex analysis of the morphological traits (size, wing shape, and wing pattern eyespots) confirm the hypothesis that the local population formed in the south of Sverdlovsk oblast and contradict the hypothesis of the migratory origin of M. russiae imago.     

Microsatellite DNA analysis of population structure in <Emphasis Type="Italic">Cornops aquaticum</Emphasis> (Orthoptera: Acrididae), over a cline for three Robertsonian translocations

The grasshopper Cornops aquaticum occurs between Mexico (23°N) and Uruguay and Central Argentina (35°S). It was recently introduced as a pest control agent of the neotropical water-hyacinth Eichhornia crassipes in South Africa. The information about the amount and distribution of genetic variability of the native populations may optimise the results of biological control programmes. Here we analyse microsatellite variability at the south of C. aquaticum’s distribution, coinciding with a cline for three polymorphic Robertsonian translocations along the Paraná River in order to: (1) estimate the amount of intrapopulation variation and its correlation with geographic/climatic variables, (2) infer interpopulation genetic variation and assess connectivity between local populations and (3) compare chromosome, morphometric and molecular variation patterns to analyse the probable causes involved in the maintenance of intraspecific variation. Our sample of 170 individuals of C. aquaticum from seven Argentine populations between latitudes 27°S to 34°S showed 211 alleles across seven microsatellite loci. Genetic diversity was estimated through average number of alleles, allelic richness, expected heterozygosity and observed heterozygosity. The analysis of molecular variance showed significant genetic differentiation among populations. Pairwise comparisons of F_ST/R_ST and Bayesian population assignment method and the discriminant analysis of principal components revealed that the two southernmost populations are more differentiated. Genetic diversity is negatively correlated with Southern latitude and with Robertsonian translocation frequencies. Our results showed that the Paraná River’s middle course populations are genetically undifferentiated and more genetically diverse than the highly chromosomally polymorphic downstream ones. The chromosomal polymorphisms are associated with increased body size in the direction in which larger size is adaptive. This may be relevant for C. aquaticum’s role as a pest control agent, since chromosome variability would enhance the ability of the species for a successful settlement in its new habitats, especially in temperate regions of the world.     

Mitochondrial sequence polymorphism in Chinese populations of <Emphasis Type="Italic">Hylobitelus xiaoi</Emphasis> (Coleoptera: Curculionidae)

The Chinese weevil, Hylobitelus xiaoi Zhang, is a major pest affecting the pine industry throughout southern China, but its dispersal is still poorly understood. We aimed to investigate its dispersal by assessing the genetic structure of seven geographical populations, using the distribution of two mitochondrial genes (cytochrome c oxidase subunit I and cytochrome b). A 1212-bp fragment of the two mitochondrial genes was sequenced for 70 individuals from seven geographical populations. The gene sequences included 861 conserved sites, 351 variable sites, 254 parsim-info sites, and 97 singleton sites. These polymorphic sites defined 24 haplotypes with a haplotype diversity of 0.825. Nucleotide diversity was low (0.04568). The differentiation parameter (0.619) was much greater than the coefficient of genetic differentiation (0.285). An analysis of molecular variance suggested that most of the variation was due to within-population differences (61.68%). This molecular data also demonstrated that there is significant divergence among the seven sampled populations of H. xiaoi. This is probably due to habitat fragmentation, preventing effective gene flow between sites. Our molecular evidence supports the view that the Chinese weevil is a native pest, and that outbreaks of H. xiaoi are caused by large plantations of the exotic slash pine.     

Latitudinal Influence on the Sexual Dimorphism of the Marine Fish <Emphasis Type="Italic">Bathygobius soporator</Emphasis> (Gobiidae: Teleostei)

Environmental gradients in a marine setting may have significant effects on morphological variations and evolutionary patterns, including sexual dimorphism variations within and between fish populations. We analyzed sexual shape and size dimorphism in accordance with Rensch and Bergmann’s rules in five coastal populations of the gobiid Bathygobius soporator along 4000 km of the Brazilian coastline. The populations differ significantly in sexual body shape dimorphism, with a tendency toward reduced intrapopulation dimorphism, increasing with latitude. Body size variation was significant between populations and population vs. sex, and inverse to Bergmann’s rule. Moreover, size dimorphism among populations of B. soporator does not follow Rensch’s rule. These data represent a rare example of inter and intrapopulation spatial variation in sexual dimorphism associated with latitude in marine fish. This suggests a complex and particularized scenario of biotic and abiotic interactions acting on local populations of B. soporator in extensive coastal areas of the Western Atlantic, with profound implications for species evolution.     

Bluffing and Waiting: Handling Effects and Post-Release Immobility in a Death-Feigning Snake (Natrix natrix)

Tonic immobility is a common response of animals to capture by a predator; in some cases, the behaviour is elaborated into death-feigning. Death-feigning is usually interpreted as a last-resort anti-predator tactic that depends on the predator ceasing or slowing its attack when the prey is apparently dead, thus buying time for the prey to escape if the predator's attention is directed elsewhere, even momentarily. I tested the effects of different handling regimes on the expression of death-feigning in the grass snake ( Natrix natrix ). In one test, degree of handling had a significant effect on frequency of death-feigning in small snakes, with snakes that received more handling feigning death more often. A second test showed that different initial handling regimes also affected the probability of death-feigning in large snakes, with snakes initially held by the tail feigning death least frequently and those initially held by the head most frequently. Thus, increased apparent threat to a vulnerable part of the body is more likely to result in death-feigning. It remains to be seen whether death-feigning can reduce the threat posed by real predators, but immobility is frequently carried over into the post-release period, during which the animal presumably weighs its chances and awaits an opportunity to escape. Regardless of cause, death-feigning snakes exhibited significantly more frequent and longer post-release immobility than did non-death-feigning snakes, which typically fled immediately upon release. Overall, small snakes feigned death less frequently than adults and were more likely to flee upon release, suggesting that immobility is a riskier anti-predator defence strategy for them.