An inherited virus influences the coexistence of parasitoid species through behaviour manipulation

The potential role of pathogens or parasites in maintaining species coexistence is well documented. However, the impact of vertically transmitted symbionts, that can markedly modify their host's biology, is largely unknown. Some females of the Drosophila parasitoid Leptopilina boulardi are infected with an inherited virus (LbFV). The virus forces females to lay supernumerary eggs in already parasitised hosts, thus allowing its horizontal transmission. Using two independent experimental procedures, we found that LbFV impacts inter-specific competition between L. boulardi and the related L. heterotoma. While L. boulardi rapidly outcompetes L. heterotoma in the absence of the virus, L. heterotoma was able to maintain or even to eliminate L. boulardi in the presence of LbFV. By forcing females to superparasitise, LbFV induced egg wastage in L. boulardi thus explaining its impact on the competition outcome. We conclude that this symbiont whose transmission is L. boulardi-density-dependant may affect the coexistence of Leptopilina species.     

Active suppression of D. melanogaster immune response by long gland products of the parasitic wasp Leptopilina boulardi

To develop inside their insect hosts, endoparasitoid wasps must either evade or overcome the host's immune system. Several ichneumonid and braconid wasps inject polydnaviruses that display well-studied immune suppressive effects. However, little is known about the strategies of immunoevasion used by other parasitoid families, such as figitid wasps. The present study provides experimental evidence, based on superparasitism and injection experiments, that the figitid species Leptopilina boulardi uses an active mechanism to suppress the Drosophila melanogaster host immune response, i.e. the encapsulation of the parasitoid eggs. The immune suppressive factors are localised in the long gland and reservoir of the female genital tractus, where virus-like particles (VLPs) have been observed. Parasitism experiments using a host tumorous strain indicate that these factors do not destroy host lamellocytes but that they impair the melanisation pathway. Interestingly, they are not susceptible to heating and are not depleted with prolonged oviposition experience, in contrast to observations reported for L. heterotoma, another figitid species. The mechanisms that prevent encapsulation of eggs from L. boulardi and L. heterotoma differ in several respects, suggesting that different physiological strategies of immunosuppression might be used by specialised and generalist parasitoids.     

Leptopilina heterotoma and L. boulardi: strategies to avoid cellular defense responses of Drosophila melanogaster

Eggs of three strains of the cynipid parasitoid Leptopilina heterotoma and a Tunisian strain (G317) of L. boulardi are not encapsulated by hemocytes of Drosophila melanogaster hosts, but the eggs of a Congolese strain (L104) of L. boulardi are encapsulated. To determine the reason for the difference in host response against the parasitoid eggs, lamellocytes (hemocytes that encapsulate foreign objects and form capsules around endogenous tissues in melanotic tumor mutants) were examined in host larvae parasitized by the five Leptopilina strains. Parasitization by the three L. heterotoma strains affected the morphology of host lamellocytes and suppressed endogenous melanotic capsule formation in melanotic tumor hosts. L104 did not alter the morphology of host lamellocytes nor block tumor formation in melanotic tumor mutant hosts. The morphology of some lamellocytes was affected by G317 parasitization but host lamellocytes were still capable of forming melanotic tumors and encapsulating dead supernumerary parasitoid larvae. Therefore, the eggs of strains affecting lamellocyte morphology are protected from encapsulation by the host's blood cells. L. heterotoma eggs float freely in the host hemocoel but L. boulardi eggs are attached to host tissue surfaces. Lamellocytes cannot infiltrate the attachment site so the capsule around the L104 egg remains incomplete. The wasp larva uses this gap in the capsule as an escape hatch for emergence.     

Niche-tracking migrants and niche-switching residents: evolution of climatic niches in New World warblers (Parulidae)

Differences in life-history traits between tropical and temperate lineages are often attributed to differences in their climatic niche dynamics. For example, the more frequent appearance of migratory behaviour in temperate-breeding species than in species originally breeding in the tropics is believed to have resulted partly from tropical climatic stability and niche conservatism constraining tropical species from shifting their ranges. However, little is known about the patterns and processes underlying climatic niche evolution in migrant and resident animals. We evaluated the evolution of overlap in climatic niches between seasons and its relationship to migratory behaviour in the Parulidae, a family of New World passerine birds. We used ordination methods to measure seasonal niche overlap and niche breadth of 54 resident and 49 migrant species and used phylogenetic comparative methods to assess patterns of climatic niche evolution. We found that despite travelling thousands of kilometres, migrants tracked climatic conditions across the year to a greater extent than tropical residents. Migrant species had wider niches than resident species, although residents as a group occupied a wider climatic space and niches of migrants and residents overlapped extensively. Neither breeding latitude nor migratory distance explained variation among species in climatic niche overlap between seasons. Our findings support the notion that tropical species have narrower niches than temperate-breeders, but does not necessarily constrain their ability to shift or expand their geographical ranges and become migratory. Overall, the tropics may have been historically less likely to experience the suite of components that generate strong selection pressures for the evolution of migratory behaviour.     

Ecological distribution and niche segregation of sibling species: the case of bean beetles, Acanthoscelides obtectus Say and A. obvelatus Bridwell

Abstract.  1. Molecular techniques have greatly added to the number of known sympatric cryptic species in insects. Ecological differences between these newly distinguished species are little explored, but niches often appear to overlap strongly. These cases are good models for exploring new ideas about species coexistence and community structure.
2.  Acanthoscelides obtectus and A. obvelatus are two sister species of bean bruchids, which have been confused until the last decade. One important ecological difference between them has emerged, however: A. obtectus is multivoltine and now distributed worldwide, whereas A. obvelatus is univoltine and restricted to Mesoamerica. Where their ranges overlap, the two species share the same host plants and larvae can sometimes complete development in the same seed.
3. The analysis of 27 622 Mexican individuals of the two species in 2001-2002 and 2002-2003 indicates that their niches overlap, but are differentiated with respect to altitude and the kind of beans (wild vs. domesticated). The principal patterns in their relative abundance in different habitats, and at different seasons, were constant from one year to the next.
4. As sympatry of these species seems to be of recent origin, the observed niche differentiation may not have evolved in response to competition, but could instead be the consequence of physiological differences, evolved independently in each species in allopatry, that pre-adapted them for different altitudes and kinds of resources.
5. The combination of biological and historical factors thus appears to allow these two sibling species to coexist in sympatry, despite their broadly overlapping ecological niches.     

Pollination niche overlap between a parasitic plant and its host

Niche theory predicts that species which share resources should evolve strategies to minimise competition for those resources, or the less competitive species would be extirpated. Some plant species are constrained to co-occur, for example parasitic plants and their hosts, and may overlap in their pollination niche if they flower at the same time and attract the same pollinators. Using field observations and experiments between 1996 and 2006, we tested a series of hypotheses regarding pollination niche overlap between a specialist parasitic plant Orobanche elatior (Orobanchaceae) and its host Centaurea scabiosa (Asteraceae). These species flower more or less at the same time, with some year-to-year variation. The host is pollinated by a diverse range of insects, which vary in their effectiveness, whilst the parasite is pollinated by a single species of bumblebee, Bombus pascuorum, which is also an effective pollinator of the host plant. The two species therefore have partially overlapping pollination niches. These niches are not finely subdivided by differential pollen placement, or by diurnal segregation of the niches. We therefore found no evidence of character displacement within the pollination niches of these species, possibly because pollinators are not a limiting resource for these plants. Direct observation of pollinator movements, coupled with experimental manipulations of host plant inflorescence density, showed that Bombus pascuorum only rarely moves between inflorescences of the host and the parasite and therefore the presence of one plant is unlikely to be facilitating pollination in the other. This is the first detailed examination of pollination niche overlap in a plant parasite system and we suggest avenues for future research in relation to pollination and other shared interactions between parasitic plants and their hosts.     

Seasonal climatic niches diverge in migratory birds

Migratory birds occupy different geographical areas during breeding and non-breeding periods, and thus different factors may determine their range limits depending on each season. One such factor is the spatial climatic component of the niche, which is widely used to model species distributions, yet the temporal component is often neglected and is generally assumed to be constant. We tested the hypothesis that the climatic niche is conserved between breeding and non-breeding areas in 355 bird species migrating through Eurasian–African flyways. For this, we performed niche overlap analyses and compared niche differences between sister or phylogenetically closely related species, as well as linking the differences to migratory distances. For more than 80% of the species, there was no or very little overlap between their breeding and non-breeding climatic niches. For most closely related species, the degree of overlap of their breeding climatic niches was larger than the overlap observed within each species, but not for their wintering climatic niches, suggesting a phylogenetic conservation of breeding climatic niches. Finally, there was a clear negative relationship between migratory distances and climatic niche overlap within each species. Our results confirmed that the climatic niche of most Eurasian–African migratory species differs between both breeding and non-breeding ranges, suggesting distinctive seasonal climatic requirements. Given these results and the geographically uneven effects of climate change, the impact of global change is likely to have different effects in each seasonal range. Hence, both breeding and non-breeding climatic data need to be considered when using species distribution models.     

Roles of endothermy in niche differentiation for ball-rolling dung beetles (Coleoptera: Scarabaeidae) along an altitudinal gradient

Abstract.  1. An analysis of whether niche differentiation in ball-rolling dung beetles can be explained by the way in which they regulate their body temperature was conducted.
2.  A priori assumptions were: (i) if thermoregulation affects niche partitioning, sympatric species must have different endothermic strategies that minimise encounters; or, alternatively (ii) if two co-occurring species show the same thermoregulation pattern and their flight periods overlap, they might be avoiding competition by exhibiting different resource preferences or different food relocation behaviour.
3. The ball-rolling dung beetles studied showed a hierarchical structure based on the species' endothermic capacity, measured as temperature excess [ T _ex= difference between body temperature ( T _b) and ambient temperature ( T _a)]. Those with a high T _ex (10–15 °C) were located exclusively at altitudes >1000 m a.s.l. On the coastal plains, species with a high T _ex were restricted to flying at night when the T _a was lower. Species with a lower T _ex (less than 10 °C higher than T _a) were found in the coastal plains zone.
4. Where there was sympatry with similar trophic habits, the species involved showed very different thermal niches, and where there was significant overlap of thermal niches between sympatric species, trophic habits of species were very different.
5. The results suggest that it is possible to use the concept of the thermal niche as a tool to explain interspecific interactions and the spatial distribution of species.     

Toxicity of venom of Asobara and Leptopilina species to Drosophila species

The Drosophila parasitoid Asobara japonica Belokobylskij (Hymenoptera: Braconidae) has highly toxic venom that kills host larvae if its injection is not followed by an injection of lateral oviduct components along with egg‐laying. In the present study, the venoms of seven other Drosophila parasitoids (Asobara rossica, Asobara rufescens, Asobara pleuralis, Leptopilina heterotoma, Leptopilina japonica, Leptopilina ryukyuensis, and Leptopilina victoriae) are tested against three kinds of Drosophila species (i.e. Drosophila species that are suitable as host for focal parasitoids, those that are resistant to the parasitoids, and a cosmopolitan species, Drosophila simulans). Venoms of the three Asobara species are not toxic to any of Drosophila species, whereas those of the four Leptopilina species are toxic to some Drosophila species. The toxicity of venom varies among Leptopilina species, and the susceptibility to venom also varies among host Drosophila species. Furthermore, toxicity and paralytic effects of venom are not correlated. Because the toxicity of venom is not adaptive for parasitoids, it may be an inevitable side effect of some components that play an essential role in parasitism.     

Interaction between host genotype and environmental conditions affects bacterial density in Wolbachia symbiosis

Regulation of microbial population density is a necessity in stable symbiotic interactions. In Wolbachia symbiosis, both bacterial and host genotypes are involved in density regulation, but environmental factors may also affect bacterial population density. Here, we studied the interaction between three strains of Wolbachia in two divergent homozygous lines of the wasp Leptopilina heterotoma at two different temperatures. Wolbachia density varied between the two host genotypes at only one temperature. Moreover, at this temperature, reciprocal-cross F1 insects displayed identical Wolbachia densities, which were intermediate between the densities in the two parental lines. While these findings confirm that the host genotype plays an important role in Wolbachia density, they also highlight its interaction with environmental conditions, making possible the evolution of local adaptations for the regulation of Wolbachia density.     

The joint effect of temperature and host species induce complex genotype-by-environment interactions in the larval parasitoid of Drosophila, Leptopilina heterotoma (Hymenoptera: Figitidae)

Phenotypic plasticity and genotype-by-environment interactions are usually studied by testing environmental factors separately. However, several environmental factors may vary simultaneously across the geographic range of a species, and thus interact to drive phenotypic and genotypic evolution. Here, we address the question of the interaction between two environmental factors in the case of a koinobiont parasitoid of Drosophila , Leptopilina heterotoma (Hymenoptera: Figitidae). Using two allopatric parasitoid lines, we analysed the joint effect of developmental temperature and host species on two major fitness components (pre-imaginal survival and fecundity) of emerged parasitoids. Our results indicate that both environmental factors strongly interact to shape the parasitoid phenotype. Moreover, complex genotype-by-environment interactions appeared that can even invert the relative fitness of the two parasitoid lines.     

A taxonomic comparison of local habitat niches of tropical trees

The integration of ecology and evolutionary biology requires an understanding of the evolutionary lability in species’ ecological niches. For tropical trees, specialization for particular soil resource and topographic conditions is an important part of the habitat niche, influencing the distributions of individual species and overall tree community structure at the local scale. However, little is known about how these habitat niches are related to the evolutionary history of species. We assessed the relationship between taxonomic rank and tree species’ soil resource and topographic niches in eight large (24–50 ha) tropical forest dynamics plots. Niche overlap values, indicating the similarity of two species’ distributions along soil or topographic axes, were calculated for all pairwise combinations of co-occurring tree species at each study site. Congeneric species pairs often showed greater niche overlap (i.e., more similar niches) than non-congeneric pairs along both soil and topographic axes, though significant effects were found for only five sites based on Mantel tests. No evidence for taxonomic effects was found at the family level. Our results indicate that local habitat niches of trees exhibit varying degrees of phylogenetic signal at different sites, which may have important ramifications for the phylogenetic structure of these communities.     

Niche overlap of two sympatric-nesting hawks Accipiter spp. in the New Jersey-New York Highlands

The food and habitat niches of two nesting species of hawks Accipiter spp were studied in an extensively forested area of the Eastern Deciduous Forest Biome Nesting habitat was quantitated at 19 Cooper s hawk A cooperu nests and 16 northern goshawk A gentilis nests There was no significant trend for Cooper's hawks to nest in less mature forests than northern goshawks as reported previously for western North America Forest habitats did not differ markedly except that shrub cover was greater at Cooper's hawk nest sites, which were also on flatter terrain and closer to roads, forest openings, and human habitation However, these few differences resulted m reducing habitat-niche overlap considerably (0 538), as was calculated using principal components analysis Mean prey weight was significantly larger for the northern goshawk which follows its 2 2-fold body weight advantage over Cooper's hawk Although bird prey was of primary importance to both Accipiter, goshawks took twice the proportion of mammals compared to their smaller congener Food-niche overlap was lowest by prey species overlap (0 470), followed by prey size class overlap (0 529), and highest by vertical foraging zone overlap (0 816) The Cooper's hawk showed the greatest niche breadth for both food and habitat niches indicating it as more of a generalist Overall, niche complementarity of food and habitat dimensions resulted in niche overdispersion along food and habitat dimensions with a total niche overlap (0 504) that was below the competition threshold These results suggested that competition (past and current) was responsible for segregating niches     

Climatic niche evolution in turtles is characterized by phylogenetic conservatism for both aquatic and terrestrial species

Understanding how the climatic niche of species evolved has been a topic of high interest in current theoretical and applied macroecological studies. However, little is known regarding how species traits might influence climatic niche evolution. Here, we evaluated patterns of climatic niche evolution in turtles (tortoises and freshwater turtles) and whether species habitat (terrestrial or aquatic) influences these patterns. We used phylogenetic, climatic and distribution data for 261 species to estimate their climatic niches. Then, we compared whether niche overlap between sister species was higher than between random species pairs and evaluated whether niche optima and rates varied between aquatic and terrestrial species. Sister species had higher values of niche overlap than random species pairs, suggesting phylogenetic climatic niche conservatism in turtles. The climatic niche evolution of the group followed an Ornstein–Uhlenbeck model with different optimum values for aquatic and terrestrial species, but we did not find consistent evidence of differences in their rates of climatic niche evolution. We conclude that phylogenetic climatic niche conservatism occurs among turtle species. Furthermore, terrestrial and aquatic species occupy different climatic niches but these seem to have evolved at similar evolutionary rates, reinforcing the importance of habitat in understanding species climatic niches and their evolution.     

Differentiation of innate but not learnt responses to host‐habitat odours contributes to rapid host finding in a parasitoid genotype

Abstract In parasitoids, host‐habitat odour can influence host searching within the habitat. This is the case in Leptopilina sp. (Hymenoptera, Figitidae), a Drosophila parasitoid searching for larvae by ovipositor probes. This a behaviour can be conditioned to fruit odours. In a previous study, the latency of probing to a fruit odour is reported to have a genetic variability within a laboratory strain. This suggests a link between rapidity of host discovery and fitness. In the present study, this hypothesis is tested by comparing responses to host‐habitat odours between two genotypes of Leptopilina heterotoma Thomson, from the Mediterranean coast (Antibes) and from Burgundy (Tailly). The two genotypes present contrasting rhythms and levels of locomotor activity linked to contrasting interspecific competition in their area of origin. The high activity observed in the Mediterranean genotype is interpreted as an adaptive response to a limited time‐window to win against a competitor species absent in Burgundy. The present study finds differentiation in innate but not learnt responses to host‐habitat odours. The more active genotype (Antibes) has a higher probability and a shorter latency of innate probing to the odours than the less active genotype (Tailly); Antibes females also find larvae and complete infestations more rapidly. Learning equalizes the probability and the latency of probing to the odours in both strains, and increases the probing duration. Innate responses to host‐habitat odours would allow time‐limited insects to increase their reproductive rate, when host predictability is high in the habitat. Selection of faster innate responses to host and habitat cues without evolution of learnt responses indicates that the initial host discovery is more crucial to fitness than subsequent ones.     

Temporal variation in spatial organization of a rodent community in the southwestern Kyzylkum desert (Middle Asia)

The distribution of niches in resource space and the niche patterns of a 14-species community of Middle Asian desert rodents were studied during two years - at low and high rodent density - using discriminant function analysis Nineteen quantitative environmental parameters (soil structure and vegetation), measured in 550 plots within 22 1 -ha grids, were considered The first three canonical axes of resource space account for 72% of the variance The first two axes represent complex environmental gradients the first axis represents a general landscape gradient from sand to clay soils, the second axis reflects a gradient of in creasing productivity The third axis reflects with in-habitat environmental variation All community parameters, as well as parameters of individual species niches, were unstable between years At the same time, different parameters vary in different extent Position of niche centroids along macro-habitat axes, as. well as macrohabitat niche breadth, were relatively stable between years, but these parameters for microhabitat axis and values of niche overlap were much more variable A strong correlation between changes m relative between-habitat niche breadth and differences in average niche overlap with relative changes in species abundances indicate density dependence of these parameters Changes in niche overlap is a consequence of between-year differences in guild patterns Guild structure was pronounced at high density when the level of niche overlap was intermediate At low density, when the level of niche overlap decreased, guild structure was incon-spicous Different levels of diversity differed in their sensitivity to density changes α-diversity was relatively constant as a result of between-year stability of niche centroid positions However level of ß-diversity varied significantly between years reflecting changes in the level of niche overlap, because a decrease in niche overlap leads to an increase in the rate of species turnover     

Niches in the Anthropocene: passerine assemblages show niche expansion from natural to urban habitats

Human‐mediated habitat transformation is increasingly evident around the world. Yet, how this transformation influences species’ niche width and overlap remains unclear. On the one hand, human‐mediated habitat transformation promotes increased species similarity through trait‐based filtering, and an increased prevalence of generalist species with broad niches, resulting in functional homogenization. On the other hand, species that colonize transformed habitats could use empty niches, resulting in decreased species similarity and an expansion of assemblage‐level niche space. Here we explore these two alternatives in eight highly diverse passerine assembles in natural, rural and urban habitats in south and southwest China, a rapidly developing region of the world. Based on stable isotopes, we found that species’ niche width increased from natural to human‐made habitats, but there were no differences in niche overlap among habitats. Therefore, we found evidence for niche expansion, with generalists appearing to use empty niches created by human habitat modification, and with assemblages being comprised of complementary species. Further research is needed to determine whether increased between‐ or within‐individual niche variation is the main driver of niche expansion in transformed habitats.     

A comprehensive framework for the study of species co‐occurrences,nestedness and turnover

Binary presence–absence matrices (rows = species, columns = sites) are often used to quantify patterns of species co‐occurrence, and to infer possible biotic interactions from these patterns. Previous classifications of co‐occurrence patterns as nested, segregated, or modular have led to contradictory results and conclusions. These analyses usually do not incorporate the functional traits of the species or the environmental characteristics of the sites, even though the outcomes of species interactions often depend on trait expression and site quality. Here we address this shortcoming by developing a method that incorporates realized functional and environmental niches, and relates them to species co‐occurrence patterns. These niches are defined from n‐dimensional ellipsoids, and calculated from the n eigenvectors and eigenvalues of the variance–covariance matrix of measured environmental or trait variables. Average niche overlap among species and the spatial distribution of niches define a triangle plot with vertices of species segregation (low niche overlap), nestedness (high niche overlap), and modular co‐occurrence (clusters of overlapping niches). Applying this framework to temperate understorey plant communities in southwest Poland, we found a consistent modular structure of species occurrences, a pattern not detected by conventional presence–absence analysis. These results suggest that, in our case study, habitat filtering is the most important process structuring understorey plant communities. Furthermore, they demonstrate how incorporating trait and environmental data into co‐occurrence analysis improves pattern detection and provides a stronger theoretical framework for understanding community structure.