Host specialization in habitat specialists and generalists

Generalists and specialists use different cues to find their habitat and essential resources. While generalists have the advantage of exploiting a wider range of resources, they are predicted to be less efficient in using one particular resource compared to specialists. The level of specialization of parasitoids can be either at the habitat or at the host level; strategies used by either type are expected to differ. We examined interactions between three aphid parasitoid species that are a habitat specialist Aphidius rhopalosiphi, a habitat generalist Aphidius ervi, and a host generalist Praon volucre on three cereal aphids, Sitobion avenae, Metopolophium dirhodum and Rhopalosiphum padi. We compared total parasitism rate across behavioral and physiological variation in a non-choice test. Next, we addressed total parasitism in two phases to examine: (1) the response of parasitoids to different hosts through the behavioral sequence from antennation through oviposition, and (2) the physiological suitability of different hosts for oviposition and larval development. Parasitization typically involved the following behavioral steps: (1) antennal contact, (2) abdominal bending, and (3) ovipositor insertion (acceptance). A. rhopalosiphi had the same number of antennal contacts with the three aphids but showed fewer instances of abdominal bending towards R. padi. Pre-contact host preference was found for A. ervi but it did not correspond to the level of acceptance. The number of antennal contacts by P. volucre corresponded to the parasitization level of the aphid species but more mummies were produced on M. dirhodum than on R. padi. These results suggest that parasitoid species that are habitat specialists react similarly to the different host species present in the same habitat, whereas generalist species exhibit clear preferences during host selection. Preferences were, however, not always related to host suitability.     

Soil seed-bank composition reveals the land-use history of calcareous grasslands

We compared soil seed banks and vegetation of recent (established on abandoned arable fields) and ancient (continuously managed as pastures at least since 1830) calcareous grasslands if there is any impact of former arable field use. The study was carried out in two regions of Southern Germany with well-preserved dry grassland vegetation: the western Jurassic mountains (Kaltes Feld) and the climatically drier eastern part of Southern Germany (Kallmünz).Total number of species in the seed bank was similar in both regions, but species composition partly differed, reflecting phytogeographical differences between the regions. The total number of emerged seedlings showed a large disparity (5457 compared to 2523 seedlings/m² in Kaltes Feld and Kallmünz, respectively).Though there were differences in seed bank composition and size, we found a uniform pattern of plant traits (affiliation to phytosociological groups, Raunkiaer plant life-forms and seed longevity), which depended on the age of the grassland.The main conclusion is that seed banks in contemporary calcareous grasslands still reflect the history of former land use – in this case arable cultivation, even though it occurred a long time ago (up to 150 years). Indicators of former arable fields are germinable seeds of weeds which have persisted in the soil to the present. By contrast, weedy species are completely absent from the seed banks of ancient grasslands. Soil seed banks of recent grasslands may be of substantial conservation importance because they may store seeds of rare and endangered weed species such as Kickxia spuria, Silene noctiflora and Stachys annua, the majority of which have already gone extinct from the current vegetation of the study sites.     

Spatial segregation of specialists and generalists in bird communities

Each species generally has a close relationship with one or more habitats and can therefore be classified as either specialist or generalist. We studied whether specialist and generalist species are spatially distributed independently of each other. Repeating the analysis for 100 of the most frequent terrestrial bird species recorded over the 10 000 sampled sites of the French Breeding Bird survey, we found that specialists were more abundant if the rest of the community was specialized, and that the inverse was also true. This pattern was far subtler than just a simple dichotomy: most species actually presented a maximum abundance at a value of community specialization similar to their own level of specialization. Bird communities appear very well defined along a specialist–generalist gradient. We believe this pattern becomes more apparent with habitat degradation. The consequences on both ecological services and community resilience may well be considerable.     

Flower handling by bumblebees: a comparison of specialists and generalists

The hypothesis that specialists have evolved behavioural adaptations to handle their preferred food types more efficiently than related generalist species was tested. Naive workers of the specialist bumblebee, Bombus consobrinus, were more efficient than those of generalist species in acquiring flower-handling skills on their specialty plant, Aconitum (monkshood). With no previous foraging experience, B. consobrinus workers began probing in the vicinity of the nectary and quickly located the nectar. Generalists, on the other hand, showed no predisposition toward correct foraging on Aconitum. They probed in the wrong places, and many gave up before finding the nectar.     

Evolving the division of labour: generalists,specialists and task allocation

The evolutionary dynamics of specialization, in the context of the division of labour, are investigated. Individuals associate in groups in which benefits are shared and costs borne individually; each individual is either a generalist who can perform all the necessary tasks, a specialist who performs a sub-set of the necessary tasks, or a parasite who contributes nothing to the group. The implications of the model are explored analytically and through both numerical and Monte Carlo methods. These methods demonstrate the evolution of populations towards stable arrangements of specialists and generalists. The fittest populations are those that divide tasks fairly and associate in large, highly specialized groups. Generalists have a distinct advantage in small groups, but the presence of generalists, ironically, lowers group fitness. Parasites are able to invade both specialized and non-specialized populations. A basic model for the continuous division of labour is also presented, demonstrating a tendency for populations to evolve increasingly unfair divisions of labour. This last result implies that an evolutionary ratchet favours disparity between the workload of specialist populations.     

Diversity and composition of larger moths in three different forest types of Southern Korea

Moth assemblages in three habitat types were investigated to find differences in species richness and abundances and to find any specific moth group(s) in each habitat. Study areas were in southern Korea: lowlands of Muan-county, Jeollanam-do Province and mid-elevations to highlands of Mt. Jirisan National Park. Moth monitoring, conducted at eight sites, was comprised of three habitat types—native forest, regenerative forest and rural landscape. A total of 4,803 individuals, consisting of 583 species in 362 genera, were identified. Species richness did not differ significantly by habitat type or elevation. ANOVA indicated that site, location and elevation significantly affected the species abundances of Drepanidae, Epiplemidae, Limacodidae, Noctuidae and Zygaenidae, while habitat type was not a significant factor. The moth abundances of Geometridae, Lasiocampidae, Lymantriidae, Notodontidae, Saturniidae, Sphingidae and Thyatiridae were not significantly associated with any independent variables. The diversity patterns of larger moths along environmental gradients and the relationship between moths and forest types were discussed.     

Distinct spatiotemporal succession of bacterial generalists and specialists in the lacustrine plastisphere

The assembly processes of generalists and specialists and their driving mechanisms during spatiotemporal succession is a central issue in microbial ecology but a poorly researched subject in the plastisphere. We investigated the composition variation, spatiotemporal succession, and assembly processes of bacterial generalists and specialists in the plastisphere, including non-biodegradable (NBMPs) and biodegradable microplastics (BMPs). Although the composition of generalists and specialists on NBMPs differed from that of BMPs, colonization time mainly mediated the composition variation. The relative abundance of generalists and the relative contribution of species replacement were initially increased and then decreased with colonization time, while the specialists initially decreased and then increased. Besides, the richness differences also affected the composition variation of generalists and specialists in the plastisphere, and the generalists were more susceptible to richness differences than corresponding specialists. Furthermore, the assembly of generalists in the plastisphere was dominated by deterministic processes, while stochastic processes dominated the assembly of specialists. The network stability test showed that the community stability of generalists on NBMPs and BMPs was lower than corresponding specialists. Our results suggested that different ecological assembly processes shaped the spatiotemporal succession of bacterial generalists and specialists in the plastisphere, but were less influenced by polymer types.     

Evolution restricts the coexistence of specialists and generalists: the role of trade-off structure

Environmental variability and adaptive foraging behavior have been shown to favor coexistence of specialists and generalists on an ecological timescale. This leaves unaddressed the question of whether such coexistence can also be expected on an evolutionary timescale. In this article, we study the attainability, through gradual evolution, of specialist-generalist coexistence, as well as the evolutionary stability of such communities when allowing for immigration. Our analysis shows that the potential for specialist-generalist coexistence is much more restricted than originally thought and strongly depends on the trade-off structure assumed. We establish that ecological coexistence is less likely for species facing a trade-off between per capita reproduction in different habitats than when the trade-off acts on carrying capacities alone. We also demonstrate that coexistence is evolutionarily stable whenever it is ecologically stable but that in most cases, such coexistence cannot be reached through gradual evolution. We conclude that an evolutionarily stable community of specialists and generalists may be created only through immigration from elsewhere or through mutations of large effect. Our results highlight that trade-offs in fitness-determining traits can have counterintuitive effects on the evolution of specialization.     

Polyphagy and diversification in tussock moths: Support for the oscillation hypothesis from extreme generalists

Theory on plasticity driving speciation, as applied to insect–plant interactions (the oscillation hypothesis), predicts more species in clades with higher diversity of host use, all else being equal. Previous support comes mainly from specialized herbivores such as butterflies, and plasticity theory suggests that there may be an upper host range limit where host diversity no longer promotes diversification. The tussock moths (Erebidae: Lymantriinae) are known for extreme levels of polyphagy. We demonstrate that this system is also very different from butterflies in terms of phylogenetic signal for polyphagy and for use of specific host orders. Yet we found support for the generality of the oscillation hypothesis, in that clades with higher diversity of host use were found to contain more species. These clades also consistently contained the most polyphagous single species. Comparing host use in Lymantriinae with related taxa shows that the taxon indeed stands out in terms of the frequency of polyphagous species. Comparative evidence suggests that this is most probably due to its nonfeeding adults, with polyphagy being part of a resulting life history syndrome. Our results indicate that even high levels of plasticity can drive diversification, at least when the levels oscillate over time.     

Habitat structure and predators: choices and consequences for rodent habitat specialists and generalists

Summary Both habitat structure and risk of predation are thought to influence rodent community composition in different habitats, but experiments on the degree to which these factors determine the use of habitat by rodents are lacking. I sought to discover (1) if cover density altered habitat choice and (2) if cover density affected the vulnerability to predators of two rodents, a habitat specialist and a habitat generalist. In laboratory experiments, the habitat specialist, the red-backed vole (Clethrionomys gapperi), preferred greater densities of both vertical (wall) and horizontal (ceiling) cover. The habitat generalist, the deer mouse (Peromyscus maniculatus), also preferred greater densities of vertical and horizontal cover, but its preferences were weaker and more inconsistent than those of C. gapperi In tests of vulnerability to domestic ferrets, C. gapperi were more vulnerable in arenas with less vertical cover, while P. maniculatus did not differ in vulnerability between the two vertical cover densities used. Vulnerability to predators in differing densities of horizontal cover was not tested because of the reduced differences in preference for this cover type between the rodent species. Risk of predation is one explanation for C. gapper's inherent preference for denser cover.     

Diversity of tiger moths in a Neotropical hotspot: determinants of species composition and identification of biogeographic units

The Brazilian Atlantic Forest is a Biodiversity Hotspot, yet many biological groups in this biome are poorly known. We compiled information on the diversity of Atlantic Forest tiger moths (Arctiidae) and assessed the resemblance among localities, whether arctiid assemblages are concordant with major vegetation types, and the importance of environmental factors in structuring the variation among assemblages. Additionally, we developed a procedure composed of subsampling and procrustean analysis to assess the robustness of the results from community composition ordinations when localities differ in species richness. To do this, we built a database from specimens deposited in the ten most important Brazilian entomological collections, and mapped species richness in one-degree latitude/longitude grid cells. We employed Principal Coordinates Analysis to assess similarities among the best-sampled localities. We obtained 8,667 records including 1,193 species, representing 60 and 20% of the estimated Brazilian and Neotropical faunas, respectively. Our subsampling procedure indicated that the ordination was not greatly affected by differences in species richness, and was congruent with major vegetation types. Lowland localities on the seacoast were quite distinct in species composition. A second group included localities in montane areas in the southeast part of the biome. The last group included localities in the southern region and those with Araucaria forests, and was associated with long distances from the ocean, even distribution of precipitation throughout the year, and large annual temperature ranges.     

Importance of grazing and soil acidity for plant community composition and trait characterisation in coastal dune grasslands

Question: Does grazing by large herbivores affect species composition or community‐wide variation in plant functional traits? Location: Dune grasslands at the Belgian coast. Methods: Plant cover and soil data were collected in 146 plots that were randomly selected at 26 grazed and ungrazed grassland sites. Plant community composition was assessed by Detrended Correspondence Analysis and mean values of plant trait categories were calculated across the plots. Results: Differentiation of plant composition and community‐wide plant trait characteristics was largely determined by grazing, soil acidity and their interaction. In ungrazed situations, a clear floristic distinction appears between acidic (non‐calcareous) and alkaline (calcareous) grasslands. In grazed situations, these floristic differences largely disappeared, indicating that grazing results in a decrease of natural variation in species composition. At higher soil pH, a larger difference in plant community composition and community‐wide plant traits was observed between grazed and ungrazed plots. In ungrazed situations, shifts in plant functional traits along the acidity gradient were observed. Conclusions: Grazing is responsible for shifts in plant community composition, and hence a decrease in plant diversity among grasslands at opposing acidity conditions in coastal dune grasslands. Therefore, care should be taken when introducing grazing as a system approach for nature conservation in dune grasslands as it may eliminate part of the natural variation in plant diversity along existing abiotic gradients.     

Nestedness in assemblages of gyrodactylids (Monogenea: Gyrodactylidea) parasitising two species of cyprinid--with reference to generalists and specialists

The structure of gyrodactylid assemblages in individual fishes of two species of cyprinid was determined. A total of 100 specimens of minnow, Phoxinus phoxinus, and 137 specimens of roach, Rutilus rutilus, were investigated for presence of gyrodactylids. Host specificity, specialists vs. generalists, was noted in each host fish. A nested pattern was recorded in parasite assemblages of minnow, the host with a dominant number of specialist gyrodactylids. A non-nested pattern was observed in parasite assemblages of roach, the host with a dominant number of generalist gyrodactylids. The host specificity appears to be a meaningful factor that determines the pattern of gyrodactylid assemblages of both fish hosts.     

The experimental evolution of specialists,generalists, and the maintenance of diversity

Environmental heterogeneity may be a general explanation for both the quantity of genetic variation in populations and the ecological niche width of individuals. To evaluate this hypothesis, I review the literature on selection experiments in heterogeneous environments. The niche width usually – but not invariably – evolves to match the amount of environmental variation, specialists evolving in homogeneous environments and generalists evolving in heterogeneous environments. The genetics of niche width are more complex than has previously been recognized, particularly with respect to the magnitude of costs of adaptation and the putative constraints on the evolution of generalists. Genetic variation in fitness is more readily maintained in heterogeneous environments than in homogeneous environments and this diversity is often stably maintained through negative frequency‐dependent selection. Moreover environmental heterogeneity appears to be a plausible mechanism for at least two well‐known patterns of species diversity at the landscape scale. I conclude that environmental heterogeneity is a plausible and possibly very general explanation for diversity across the range of scales from individuals to landscapes.     

Soil pH and phosphorus drive species composition and richness in semi-natural heathlands and grasslands unaffected by twentieth-century agricultural intensification

Background: Increased soil phosphorus (P) caused by agricultural intensification has been associated with decreased plant species richness (SR) in central Europe. How plant communities and soil P gradients are related in unimproved open habitats remains unclear.

Aims: The aim of this article was to characterise the relationship between soil chemical parameters and plant species composition and richness in unimproved open habitats.

Methods: The influence of soil chemical parameters (pH, P, K, Mg) on species composition was assessed, using data from 40 heathland and 54 grassland plots, by non-metric multidimensional scaling and permutational multivariate analysis of variance. The relationship between soil chemical parameters and SR was tested by linear mixed effects models.

Results: A direct relationship between heathland community composition and pH was observed, explaining 10% of variation in species composition, while P, Mg and pH together explained 17% of variation in grassland composition. In heathlands, SR increased with increasing pH, whereas in grasslands, SR decreased with increasing soil P.

Conclusions: Soil chemical parameters were substantially related to plant community composition and richness. In an area spared from a century of agricultural intensification, reduced pH appeared to constrain SR in heathlands, while even slight P increases (<10 mg kg^?1) depressed plant SR in semi-natural grasslands.     

Short-time effects of land-use changes on O-horizon in subalpine grasslands

Due to economic needs, a process of abandonment of unfavourably situated areas and intensified cultivation on favoured areas has taken place in the alpine region. This abandonment not only means the loss of a cultural landscape based on centuries of care but also interference with an established ecological balance. We wanted to investigate the impact of land-use, such as mowing, fertilisation and irrigation and the effects of abandonment on the O-horizon type, quantity and quality. Therefore, we selected areas with different types of land use but with equivalent site conditions in the Passeier Valley (Bolzano-South Tyrol province) for investigation. The areas studied were as follows: an organically fertilised and irrigated, intensively managed hay meadow mown once a year; an organically fertilised, intensively managed hay meadow mown once a year; a lightly managed hay meadow mown every second year; a lightly managed hay meadow mown every 3–4 years; an area densely covered with dwarf shrubs abandoned 10 years ago; and a reforested area abandoned 30 years ago. These selected areas represent different intensities of land use or stages of succession. On these areas, vegetation, phytomass, litter quantity, humus type, O-horizon quantity, litter decomposition and C and N storage, were analysed in detail. Abandonment causes the percentage of coverage with grasses and herbs to decrease from 100 to 10%, while lignified species increase from 0 to 90%. This leads to growth of the phytomass pools, which in turn causes an increase in the annual litter quantity (from 117 to 525 g m⁻²). Significant changes in litter quality in the case of reduced use or abandonment increases the litter’s resistance to decomposition, decreasing overall decomposition. Litter quantity and quality then have a significant impact on the O-horizon quantity. This increases strongly with decreasing intensity of land use or longer times since abandonment (115–1,180 g m⁻²). This leads to a significant increase of C and N storage and a change in O-horizon type ranging from no definite O-horizon or mull O-horizons on regularly used areas to moder on abandoned areas.     

Working together to respond to the challenges of EU policy to replace animal testing

This paper presents a personal perspective on efforts during the past 15 years to replace animal testing for assessing the safety of chemicals and products. It is based on an invited lecture--the FRAME Annual Lecture--given in October 2005, with the theme of "making progress by working together" (government-industry-academia-NGOs). Where we have achieved some successes, these have clearly been due to effective cooperation and collaboration between the relevant stakeholders. In recent times, there has not been this same level of active commitment and coordination. This needs to change, since, if we are to make good progress in the years to come in responding to the new challenges of the EU policy to replace animal testing, this will undoubtedly require us to work together, hopefully facilitated by effective leadership and coordination from the EU policy-makers themselves.