Life on the edge: a hybrid zone in AZbinaria hippolyti (Gastropoda: Clausiliidae) from Crete

The hybrid zone between the Greek land snails Albinaria hippolyti aphrodite and Albinaria hippolyti harmonia runs exactly along the edge of a gorge. At two localities, we analysed patterns of variation in three enzyme loci, six conchological characters, and two anatomical characters across the zone. Clines in the various characters are largely coincident and concordant. They are narrow (approximately 30 m wide) where the ecological transition is abrupt, and wider (approximately 120 m wide) where a valley cuts into the gorge, making the environmental change more gradual. In spite of the obvious correlation with an ecological transition, we do not believe the zone is maintained by a selection gradient. Instead, direct and indirect evidence for hybrid disadvantage and evidence for directional dispersal at the edge of the gorge suggest that it is a tension zone, held at the ecotone by differential dispersal of at least one of the parental types. We speculate on the possibility that reinforcement may act in the hybrid zone. Our data on dispersal and reproductive isolating mechanisms lead us to believe that reinforcement may only be possible at localities where the hybrid zone is very narrow.     

Food plant and herbivore host species affect the outcome of intrinsic competition among parasitoid larvae

1. In nature, several parasitoid species often exploit the same stages of a common herbivore host species and are able to coexist despite competitive interactions amongst them. Less is known about the direct effects of resource quality on intrinsic interactions between immature parasitoid stages. The present study is based on the hypothesis that variation in the quality or type of plant resources on which the parasitoids indirectly develop may be complementary and thus facilitate niche segregation favouring different parasitoids in intrinsic competition under different dietary regimes. 2. The present study investigated whether two herbivore species, the cabbage butterflies Pieris brassicae and Pieris rapae (Pieridae), and the quality of two important food plants, Brassica oleracea and Brassica nigra (Brassicaceae), affect the outcome of intrinsic competition between their primary larval endoparasitoids, the gregarious Cotesia glomerata (Braconidae) and the solitary Hyposoter ebeninus (Ichneumonidae). 3. Hyposoter ebeninus is generally an intrinsically superior competitor over C. glomerata. However, C. glomerata survived more antagonistic encounters with H. ebeninus when both developed in P. brassicae rather than in P. rapae caterpillars, and while its host was feeding on B. nigra rather than B. oleracea. Moreover, H. ebeninus benefitted from competition by its higher survival in multiparasitised hosts. 4. These results show that both plant and herbivore species mediate the battleground on which competitive interactions between parasitoids are played out and may affect the outcomes of these interactions in ways that enable parasitoids to segregate their niches. This in turn may promote coexistence among parasitoid species that are associated with the same herbivore host.     

Arbuscular mycorrhizal fungi reduce growth and infect roots of the non‐host plant Arabidopsis thaliana

The arbuscular mycorrhizal (AM) symbiosis is widespread throughout the plant kingdom and important for plant nutrition and ecosystem functioning. Nonetheless, most terrestrial ecosystems also contain a considerable number of non‐mycorrhizal plants. The interaction of such non‐host plants with AM fungi (AMF) is still poorly understood. Here, in three complementary experiments, we investigated whether the non‐mycorrhizal plant Arabidopsis thaliana, the model organism for plant molecular biology and genetics, interacts with AMF. We grew A. thaliana alone or together with a mycorrhizal host species (either Trifolium pratense or Lolium multiflorum) in the presence or absence of the AMF Rhizophagus irregularis. Plants were grown in a dual‐compartment system with a hyphal mesh separating roots of A. thaliana from roots of the host species, avoiding direct root competition. The host plants in the system ensured the presence of an active AM fungal network. AM fungal networks caused growth depressions in A. thaliana of more than 50% which were not observed in the absence of host plants. Microscopy analyses revealed that R. irregularis supported by a host plant was capable of infecting A. thaliana root tissues (up to 43% of root length colonized), but no arbuscules were observed. The results reveal high susceptibility of A. thaliana to R. irregularis, suggesting that A. thaliana is a suitable model plant to study non‐host/AMF interactions and the biological basis of AM incompatibility.     

Community structure and abundance of insects in response to early‐season aphid infestation in wild cabbage populations

1. Changes in the arthropod community structure can be attributed to differences in constitutively expressed plant traits or those that change depending on environmental conditions such as herbivory. Early‐season herbivory may have community‐wide effects on successive insect colonisation of host plants and the identity of the initially inducing insect may determine the direction and strength of the effects on the dynamics and composition of the associated insect community. 2. Previous studies have addressed the effect of early infestation with a chewing herbivore. In the present study, the effect of early infestation was investigated with a phloem‐feeding aphid [Brevicoryne brassicae L. (Hemiptera, Aphididae)] on the insect community associated with three wild cabbage (Brassica oleracea L.) populations, which are known to differ in defence chemistry, throughout the season in field experiments. 3. Aphid infestation had asymmetric effects on the associated insect community and only influenced the abundance of the natural enemies of aphids, but not that of chewing herbivores and their natural enemies. The effect size of aphid infestation further depended on the cabbage population. 4. Aphid feeding has been previously reported to promote host‐plant quality for chewing herbivores, which has been attributed to antagonism between the two major defence signalling pathways controlled by the hormones salicylic acid (SA) and jasmonic acid (JA), respectively. Our results show no effects of early infestation by aphids on chewing herbivores, suggesting the absence of long‐term JA–SA antagonism. 5. Investigating the effects of the identity of an early‐season coloniser and genotypic variation among plant populations on insect community dynamics are important in understanding insect–plant community ecology.     

Invertebrate grazing and riparian shade as controllers of nuisance algae in a eutrophic river

1. Algal growth in lotic systems is controlled either from the bottom‐up (e.g. nutrients and light, which determine growth rates) or from the top‐down (e.g. grazing pressure, which reduces accumulated biomass). Nutrient‐enriched streams that support large and diverse grazing macroinvertebrate populations and those with shaded riparian corridors rarely suffer from excessive algal growth. 2. In this study, the density of benthic algivorous macroinvertebrates was experimentally manipulated in shaded and open nutrient‐enriched stream habitats of the Owennagearagh River, south‐west Ireland. The ability of macroinvertebrate grazers and riparian shade to control benthic algal growth [particularly the nuisance alga, Cladophora glomerata (L. Kütz)] was investigated. Three sites with markedly different concentrations of plant nutrients (one site upstream and two sites downstream of the sewage outfall) were selected. The density of grazing invertebrates colonising ceramic tiles was reduced using high‐voltage localised electric pulses. Replicates of treatment (grazer‐excluded) and control (grazed) tiles were deployed in open and shaded (<25 and >80% canopy cover, respectively) patches of stream bed, in each site. 3. After 2‐week Cladophora cover, periphytic chlorophyll a and biofilm ash‐free dry mass (AFDM) were quantified for all experimental tiles. Values for all three parameters were highest on grazer‐excluded tiles from open patches. Grazed tiles from open patches accrued little Cladophora and had significantly lower levels of chlorophyll a and AFDM. Nutrient inputs were found to have an impact on the density of grazing invertebrates, with higher densities of Baetis nymphs at the most nutrient‐enriched site. 4. Our results demonstrate that in eutrophic, high‐light streams, filamentous algae can quickly accumulate to nuisance levels in the absence of invertebrate grazers. In future, greater attention should be paid to the role of grazing invertebrates in controlling nuisance algae in streams, in addition to algal–nutrient relationships.     

Contribution of biofilm‐dwelling consumers to pelagic–benthic coupling in a large river

1. Over the course of this 17‐month study, we assessed the potential loss of plankton (bacteria, algae, heterotrophic flagellates) to consumers (ciliates and rotifers) within mature biofilms established on natural substrata exposed to the main current of the River Rhine (Germany). Once a month, in flow cells in a bypass system to the River Rhine, we measured the clearance rates of the biofilm‐associated consumers on the different groups within the natural plankton. 2. Ciliates were the most dominant consumers, among which planktivorous groups, particularly peritrichs and (in spring and summer) heterotrichs dominated. Consumer biomass varied with season, with the highest density occurring directly after the appearance of the phytoplankton spring peak. 3. Clearance rates on plankton ranged from 96 to 565 L m^?2 d^?1 for bacteria and 66–749 L m^?2 d^?1 for algae, with a preference for algae in summer and for bacteria in winter. This pattern coincided with seasonal changes in the structures of the grazer communities. The consumers (both ciliates and rotifers with total standing stocks ranging between 19 and 572 mg C m^?2) imported a substantial amount of organic matter (between 15 and 137 mg C m^?2 d^?1) into the biofilm. 4. These results highlight the potential importance of consumers in the biofilm as a trophic link between the plankton and the benthos, a function that has hitherto mostly been attributed to filter‐feeding bivalves. In contrast to bivalves, the biofilm‐dwelling consumers show a more dynamic response towards the plankton density and composition. Such dynamic components need to be considered when estimating total plankton consumption by the benthos.     

Herbivore‐induced plant responses in Brassica oleracea prevail over effects of constitutive resistance and result in enhanced herbivore attack

1. Plant responses to herbivore attack may have community‐wide effects on the composition of the plant‐associated insect community. Thereby, plant responses to an early‐season herbivore may have profound consequences for the amount and type of future attack. 2. Here we studied the effect of early‐season herbivory by caterpillars of Pieris rapae on the composition of the insect herbivore community on domesticated Brassica oleracea plants. We compared the effect of herbivory on two cultivars that differ in the degree of susceptibility to herbivores to analyse whether induced plant responses supersede differences caused by constitutive resistance. 3. Early‐season herbivory affected the herbivore community, having contrasting effects on different herbivore species, while these effects were similar on the two cultivars. Generalist insect herbivores avoided plants that had been induced, whereas these plants were colonised preferentially by specialist herbivores belonging to both leaf‐chewing and sap‐sucking guilds. 4. Our results show that community‐wide effects of early‐season herbivory may prevail over effects of constitutive plant resistance. Induced responses triggered by prior herbivory may lead to an increase in susceptibility to the dominant specialists in the herbivorous insect community. The outcome of the balance between contrasting responses of herbivorous community members to induced plants therefore determines whether induced plant responses result in enhanced plant resistance.     

What drives long‐distance movements in the nomadic Grey Teal Anas gracilis in Australia?

In contrast to northern temperate environments, where day length and temperature changes are obvious proximate cues for movement to resource-rich breeding habitats, the cues for movement used by birds in an often resource-poor, stochastic environment are less obvious. We recorded long-distance movements of 23 Grey Teal Anas gracilis using satellite telemetry for up to 879 days and examined the relationship between those movements and environmental factors, such as heavy rainfall and flooding, at the destination site. We identified 32 long-distance (> 150 km) movements that met our criterion for minimally interrupted flight between origin and destination. Thirteen of these flights coincided with rainfall and/or flooding events up to 1050 km from the origin. However, some ducks moved without any clear beneficial conditions at the destination onto small wetlands in regions with little surface water. The data suggest that there are two types of long-distance movement – ranging and directed. These flights occurred over distances up to 1200 km across the arid inland. The rates and distances of movement suggest that long-distance movements of Grey Teal entail high energy costs as in waterfowl elsewhere. We conclude that the proximate controls of directed movements need not be very different from those of their temperate counterparts.     

Predicting adaptation of phenology in response to climate change, an insect herbivore example

Climate change has led to an advance in phenology in many species. Synchrony in phenology between different species within a food chain may be disrupted if an increase in temperature affects the phenology of the different species differently, as is the case in the winter moth egg hatch–oak bud burst system. Operophtera brumata (winter moth) egg hatch date has advanced more than Quercus robur (pedunculate oak) bud burst date over the past two decades. Disrupted synchrony will lead to selection, and a response in phenology to this selection may lead to species genetically adapting to their changing environment. However, a prerequisite for such genetic change is that there is sufficient genetic variation and severe enough fitness consequences. So far, examples of observed genetic change have been few. Using a half-sib design, we demonstrate here that O. brumata egg-hatching reaction norm is heritable, and that genetic variation exists. Fitness consequences of even a few days difference between egg hatch and tree bud opening are severe, as we experimentally determined. Estimates of genetic variation and of fitness were then combined with a climate scenario to predict the rate and the amount of change in the eggs' response to temperature. We predict a rapid response to selection, leading to a restoration of synchrony of egg hatch with Q. robur bud opening. This study shows that in this case there is a clear potential to adapt – rapidly – to environmental change. The current observed asynchrony is therefore not due to a lack of genetic variation and at present it is unclear what is constraining O. brumata to adapt. This kind of model may be particularly useful in gaining insight in the predicted amount and rate of change due to environmental changes, given a certain genetic variation and selection pressure.     

Climate change leads to decreasing bird migration distances

Global climate change has led to warmer winters in NW Europe, shortening the distance between suitable overwintering areas and the breeding areas of many bird species. Here we show that winter recovery distances have decreased over the past seven decades, for birds ringed during the breeding season in the Netherlands between 1932 and 2004. Of the 24 species included in the analysis, we found in 12 a significant decrease of the distance to the wintering site. Species from dry, open areas shortened their distance the most, species from wet, open areas the least, while woodland species fall in between the other two habitats. The decline in migration distance is likely due to climate change, as migration distances are negatively correlated with the Dutch temperatures in the winter of recovery. With a shorter migration distance, species should be better able to predict the onset of spring at their breeding sites and this could explain the stronger advancement of arrival date found in several short distance species relative to long-distance migrants.