Comparing the conservatism of ecological interactions in plant–pollinator and plant–herbivore networks

Conservatism in species interaction, meaning that related species tend to interact with similar partners, is an important feature of ecological interactions. Studies at community scale highlight variations in conservatism strength depending on the characteristics of the ecological interaction studied. However, the heterogeneity of datasets and methods used prevent to compare results between mutualistic and antagonistic networks. Here we perform such a comparison by taking plant–insect communities as a study case, with data on plant–herbivore and plant–pollinator networks. Our analysis reveals that plants acting as resources for herbivores exhibit the strongest conservatism in species interaction among the four interacting groups. Conservatism levels are similar for insect pollinators, insect herbivores and plants as interacting partners of pollinators, although insect pollinators tend to have a slightly higher conservatism than the two others. Our results thus clearly support the current view that within antagonistic networks, conservatism is stronger for species as resources than for species as consumer. Although the pattern tends to be opposite for plant–pollinator networks, our results suggest that asymmetry in conservatism is much less pronounced between the pollinators and the plant they interact with. We discuss these differences in conservatism strength in relation with the processes structuring plant–insect communities.     

Restoration of plant–pollinator interaction networks via species translocation

The recent decline in pollinator biodiversity, notably in the case of wild bee populations, puts both wild and agricultural ecosystems at risk of ecological community collapse. This has triggered calls for further study of these mutualistic communities in order to more effectively inform restoration of disturbed plant–pollinator communities. Here, we use a dynamic network model to test a variety of translocation strategies for restoring a community after it experiences the loss of some of its species. We consider the reintroduction of extirpated species, both immediately after the original loss and after the community has reequilibrated, as well as the introduction of other native species that were originally absent from the community. We find that reintroducing multiple highly interacting generalist species best restores species richness for lightly disturbed communities. However, for communities that experience significant losses in biodiversity, introducing generalist species that are not originally present in the community may most effectively restore species richness, although in these cases the resultant community often shares few species with the original community. We also demonstrate that the translocation of a single species has a minimal impact on both species richness and the frequency of community collapse. These results have important implications for restoration practices in the face of varying degrees of community perturbations, the refinement of which is crucial for community management.     

Climate mediates roles of pollinator species in plant–pollinator networks

Aim

Understanding how climate conditions influence plant–pollinator interactions at the global scale is crucial to understand how pollinator communities and ecosystem function respond to environmental change. Here, we investigate whether climate drives differences in network roles of the main insect pollinator orders: Diptera, Coleoptera, Lepidoptera and Hymenoptera.

Location

Global.

Time period

1968–2020.

Major taxa studied

Diptera, Coleoptera, Lepidoptera and Hymenoptera.

Methods

We collated plant–pollinator networks from 26 countries and territories across the five main Köppen–Geiger climate zones. In total, we compiled data from 101 networks that included >1500 plant species from 167 families and >2800 pollinator species from 163 families. We assessed differences in the composition of plant–pollinator interactions among climate zones using a permutational ANOVA. We calculated standard network metrics for pollinator taxonomic groups and used Bayesian generalized mixed models to test whether climate zone influenced the proportion of pollinator network links and the level of pollinator generalism.

Results

We found that climate is a strong driver of compositional dissimilarities between plant–pollinator interactions. Relative to other taxa, bees and flies made up the greatest proportion of network links across climate zones. When network size was accounted for, bees were the most generalist pollinator group in the tropics, whereas non-bee Hymenoptera were the most generalist in arid zones, and syrphid flies were the most generalist in polar networks.

Main conclusions

We provide empirical evidence at the global scale that climate strongly influences the roles of different pollinator taxa within networks. Importantly, non-bee taxa, particularly flies, play central network roles across most climate zones, despite often being overlooked in pollination research and conservation. Our results identify the need for greater understanding of how global environmental change affects plant–pollinator interactions.     

Contrasting effects of invasive plants in plant–pollinator networks

The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity. However, there is little information available about the effect of introduced species on the structure of such networks. We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant–pollinator networks. To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders. Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators. Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively. Invasive plants acted as pollination super generalists. The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant–pollinator networks. Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant–pollinator network. These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant–pollinator network.     

Size-specific interaction patterns and size matching in a plant�Cpollinator interaction web

Background and Aims

Many recent studies show that plant–pollinator interaction webs exhibit consistent structural features such as long-tailed distributions of the degree of generalization, nestedness of interactions and asymmetric interaction dependencies. Recognition of these shared features has led to a variety of mechanistic attempts at explanation. Here it is hypothesized that beside size thresholds and species abundances, the frequency distribution of sizes (nectar depths and proboscis lengths) will play a key role in determining observed interaction patterns.

Methods

To test the influence of size distributions, a new network parameter is introduced: the degree of size matching between nectar depth and proboscis length. The observed degree of size matching in a Spanish plant–pollinator web was compared with the expected degree based on joint probability distributions, integrating size thresholds and abundance, and taking the sampling method into account.

Key Results

Nectar depths and proboscis lengths both exhibited right-skewed frequency distributions across species and individuals. Species-based size matching was equally close for plants, independent of nectar depth, but differed significantly for pollinators of dissimilar proboscis length. The observed patterns were predicted well by a model considering size distributions across species. Observed size matching was closer when relative abundances of species were included, especially for flowers with openly accessible nectar and pollinators with long proboscises, but was predicted somewhat less successfully by the model that included abundances.

Conclusions

The results suggest that in addition to size thresholds and species abundances, size distributions are important for understanding interaction patterns in plant–pollinator webs. It is likely that the understanding will be improved further by characterizing for entire communities how nectar production of flowers and energetic requirements of pollinators covary with size, and how sampling methods influence the observed interaction patterns.Key words: Plant–pollinator community, flower morphology, generalization, nectar, pollination network, body size, size matching, specialization     

Urbanization alters the spatiotemporal dynamics of plant–pollinator networks in a tropical megacity

Urbanization is a major driver of biodiversity change but how it interacts with spatial and temporal gradients to influence the dynamics of plant–pollinator networks is poorly understood, especially in tropical urbanization hotspots. Here, we analysed the drivers of environmental, spatial and temporal turnover of plant–pollinator interactions (interaction β-diversity) along an urbanization gradient in Bengaluru, a South Indian megacity. The compositional turnover of plant–pollinator interactions differed more between seasons and with local urbanization intensity than with spatial distance, suggesting that seasonality and environmental filtering were more important than dispersal limitation for explaining plant–pollinator interaction β-diversity. Furthermore, urbanization amplified the seasonal dynamics of plant–pollinator interactions, with stronger temporal turnover in urban compared to rural sites, driven by greater turnover of native non-crop plant species (not managed by people). Our study demonstrates that environmental, spatial and temporal gradients interact to shape the dynamics of plant–pollinator networks and urbanization can strongly amplify these dynamics.     

Robustness of size–structure across ecological networks in pelagic systems

The study of biomass size distributions has become an important tool for addressing aquatic ecosystem complexity and the consequences of anthropogenic disturbances. However, it remains unclear how changes in pelagic food web topology affect the biomass size–structure. Employing a dynamic multispecies bioenergetic consumer-resource model, we simulated biomass trajectories over time in 10,000 virtual networks of varying topology to address which food web properties are important in determining size–structure in pelagic systems. The slopes of the normalized biomass size spectra (NBSS) and Pareto’s shape parameter (γ) of our modeled communities are consistent with theoretically expected values for steady-state systems and empirical values reported for several aquatic ecosystems. We found that the main drivers of the NBSS slope and Pareto’s γ were the slope of the relationship between body mass and trophic level, the maximum trophic level of the food web, and the stability of total community biomass. Our analyses showed a clear conservative trend in pelagic community size–structure as demonstrated by the robustness of the NBSS slope and Pareto’s γ against most of the topological changes in virtual networks. Nevertheless, these analyses also caution that major disturbances in large-bodied or top-trophic level individuals may disrupt this stable pattern.     

Topological change of Andean plant–pollinator networks along an altitudinal gradient

Pollination interaction networks exhibit structural regularities across a wide range of natural environments. Long-tailed degree distribution, nestedness, and modularity are the most prevalent topological patterns found in most bipartite networks analyzed up to day. In this work we evaluate the variation of these topological properties along an altitudinal gradient. To this end, we examined four plant–pollinator networks from the Chilean Andes at 33°S, in range from 1800 to 3600 m elevation. Our results indicate that network topology is strongly and systematically affected by elevation. At increasing altitude, the number of potential visitors per plant decreased, and species’ degree distributions are closer to random expectations. On the other hand, the nested structure of mutualistic interactions systematically decreased with elevation, and network modularity was significantly higher than random expectations over the entire altitudinal range. In addition, at increasing elevations the pollination networks were organized in fewer and more strongly connected modules. Our results suggest that the severe abiotic conditions found at increased elevations translate into less organized pollination networks.     

Variability and specialization of plant–pollinator systems in a northern maritime grassland

To detect plant specialization to pollinator insects and to further examine the pollination syndrome hypotheses, flowering angiosperms and insect visitors were recorded in a northern maritime grassland community. Associations between plant species and insect groups were analyzed using binomial tests based on census data obtained from two sites over 3years. Preference to an insect group by a plant species was expressed as a significant deviation of the actual proportion (i.e. the proportion of the number of flower visits by the insect group to the plant species in the total number of visits to the plant species) of the expected proportion (i.e. the proportion of the number of visits by the insect group in the total visits during its flowering period). Preference to a plant species by an insect group was similarly expressed using the number of flowers of the plant species visited by the insect group. Most significant preferences varied temporally or spatially. Variability in the preferences is suggested to have resulted from temporal and spatial variations in the abundance and species composition of both flowers and insects. However, in species showing variable preferences, significant specialization in the pollinator insect group (i.e. relatively constant, mutual preferences) were demonstrated for seven and five plant species at the two study sites, respectively. Most specializations were found in associations with bumblebees. Bumblebee specialists were significantly well represented in the flower shape types gullet and flag and the flower color types violet, which supports the pollination syndrome theory.     

Global versus local extinction in a network model of plant–pollinator communities

The loss of a species from an ecological community can trigger a cascade of additional extinctions; the complex interactions that comprise ecological communities make the dynamics and impacts of such a cascade challenging to predict. Previous studies have typically considered global extinctions, where a species cannot re-enter a community once it is lost. However, in some cases a species only becomes locally extinct, and may be able to reinvade from surrounding communities. Here, we use a dynamic, Boolean network model of plant–pollinator community assembly to analyze the differences between global and local extinction events in mutualistic communities. As expected, we find that compared to global extinctions, communities respond to local extinctions with lower biodiversity loss, and less variation in topological network properties. We demonstrate that in the face of global extinctions, larger communities suffer greater biodiversity loss than smaller communities when similar proportions of species are lost. Conversely, smaller communities suffer greater loss in the face of local extinctions. We show that targeting species with the most interacting partners causes more biodiversity loss than random extinctions in the case of global, but not local, extinctions. These results extend our understanding of how mutualistic communities respond to species loss, with implications for community management and conservation efforts.     

Delayed response in a plant–pollinator system to experimental grassland fragmentation

The fragmentation of natural habitat is considered to be a major threat to biodiversity. Decreasing habitat quality and quantity caused by fragmentation may lead to a disruption of plant–pollinator interactions and to a reduction in sexual reproduction in plant species. We conducted a 6-year field experiment to investigate the effects of small-scale fragmentation on plant–pollinator interactions and genetic diversity in the self-compatible Betonica officinalis. We examined the abundance and composition of pollinators, the foraging behaviour of bumblebees and the performance, outcrossing rate and genetic diversity of B. officinalis after 2 and 6 years in experimentally fragmented nutrient-poor, calcareous grassland in the northern Swiss Jura mountains. Fragments of different size (2.25 and 20.25 m²) were isolated by a 5-m-wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland. Experimental grassland fragmentation altered the composition of B. officinalis pollinators and reduced their flower visitation rate. Furthermore, the foraging behaviour of bumblebees was changed in the fragments. After 6 years of fragmentation seed weight was higher in fragments than in control plots. However, the densities of B. officinalis rosettes and inflorescences, plant height and inflorescence length were not affected by fragmentation. The outcrossing frequency of B. officinalis growing in fragments was reduced by 15% after 2 years and by 33% after 6 years of experimental fragmentation. This resulted in a significant reduction of the genetic diversity in seedlings emerging in fragments after 6 years. Our study shows that small-scale habitat fragmentation can disturb the interaction between B. officinalis and pollinators resulting in a reduced outcrossing frequency and genetic diversity in plants growing in fragments. However, the response to fragmentation was considerably delayed. This finding strengthens the claim for long-term field experiments with proper replications and controls to assess delayed effects of habitat fragmentation.     

The town Crepis and the country Crepis: How does fragmentation affect a plant–pollinator interaction?

In fragmented habitats, one cause of the decrease of plant diversity and abundance is the disruption of plant–animal interactions, and in particular plant–pollinator interactions. Since habitat fragmentation acts both on pollinator behaviour and plant reproduction, its consequences for the stability of such interactions are complex. An extreme case of habitat fragmentation occurs in urbanised areas where suitable habitat (in the present study small patches around ornamental trees) is embedded in a highly unsuitable environment (concrete matrix). Based on simple experiments, we ask whether pollinators can adapt their foraging behaviour in response to the amount of available resources (flowers) in the fragments and their isolation, as predicted by the optimal foraging theory. To do so we analysed the effect of fragmentation on the behaviour of pollinators visiting Crepis sancta (L.) Bornm. (Asteraceae), which forms large populations in the countryside and patchy populations in urban environments. More precisely we studied pollinator visitation rates, capitulum visit durations, capitulum search durations and capitulum size choice. Pollinators chose larger capitula in both types of populations and their foraging behaviour differed between the two population types in three ways: (1) pollinator visits were lower in urban fragmented populations, perhaps due to the lower accessibility of urban patches; (2) capitulum visit durations were longer in urban fragmented populations, a possible compensation of energy lost during flights among patches; and (3) capitulum search durations where longer in urban fragmented populations, which may represent an increase in capitulum prospecting effort. We discuss the possible impacts of such differences for plant population functioning in the two types of populations.     

Effects of elevated ultraviolet-B radiation on a plant–herbivore interaction

Enhanced ultraviolet-B (UV-B) radiation may have multiple effects on both plants and animals and affect plant–herbivore interactions directly and indirectly by inducing changes in host plant quality. In this study, we examined combined effects of UV-B and herbivory on the defence of the mountain birch (Betula pubescens ssp. czerepanovii) and also the effects of enhanced UV-B radiation on a geometrid with an outbreak cycle: the autumnal moth (Epirrita autumnata). We established an experiment mimicking ozone depletion of 30% (a relevant level when simulating ozone depletion above Northern Lapland). Both arctic species responded only slightly to the enhanced level of UV-B radiation, which may indicate that these species are already adapted to a broader range of UV-B radiation. UV-B exposure slightly induced the accumulation of myricetin glycosides but had no significant effect on the contents of quercetin or kaempferol derivatives. Mountain birch seedlings responded more efficiently to herbivory wounding than to enhanced UV-B exposure. Herbivory induced the activities of foliar oxidases that had earlier been shown to impair both feeding and growth of moth larvae. In contrast, the contents of foliar phenolics did not show the same response in different clones, except for a decrease in the contents of tannin precursors. The induction of foliar phenoloxidase activities is a specific defence response of mountain birches against insect herbivory. To conclude, our results do not support the hypothesis that the outbreak cycle of the autumnal moth can be explained by the cycles of solar activity and UV-B.     

Towards a mechanistic understanding of the impacts of nitrogen deposition on producer–consumer interactions

Nitrogen (N) deposition has increased substantially since the second half of the 20th century due to human activities. This increase of reactive N into the biosphere has major implications for ecosystem functioning, including primary production, soil and water chemistry and producer community structure and diversity. Increased N deposition is also linked to the decline of insects observed over recent decades. However, we currently lack a mechanistic understanding of the effects of high N deposition on individual fitness, species richness and community structure of both invertebrate and vertebrate consumers. Here, we review the effects of N deposition on producer–consumer interactions, focusing on five existing ecological frameworks: C:N:P ecological stoichiometry, trace element ecological stoichiometry, nutritional geometry, essential micronutrients and allelochemicals. We link reported N deposition-mediated changes in producer quality to life-history strategies and traits of consumers, to gain a mechanistic understanding of the direction of response in consumers. We conclude that high N deposition influences producer quality via eutrophication and acidification pathways. This makes oligotrophic poorly buffered ecosystems most vulnerable to significant changes in producer quality. Changes in producer quality between the reviewed frameworks are often interlinked, complicating predictions of the effects of high N deposition on producer quality. The degree and direction of fitness responses of consumers to changes in producer quality varies among species but can be explained by differences in life-history traits and strategies, particularly those affecting species nutrient intake regulation, mobility, relative growth rate, host-plant specialisation, ontogeny and physiology. To increase our understanding of the effects of N deposition on these complex mechanisms, the inclusion of life-history traits of consumer species in future study designs is pivotal. Based on the reviewed literature, we formulate five hypotheses on the mechanisms underlying the effects of high N deposition on consumers, by linking effects of nutritional ecological frameworks to life-history strategies. Importantly, we expect that N-deposition-mediated changes in producer quality will result in a net decrease in consumer community as well as functional diversity. Moreover, we anticipate an increased risk of outbreak events of a small subset of generalist species, with concomitant declines in a multitude of specialist species. Overall, linking ecological frameworks with consumer life-history strategies provides a mechanistic understanding of the impacts of high N deposition on producer–consumer interactions, which can inform management towards more effective mitigation strategies.     

Post hoc assessment of host plant use in a generalist invader: implications for understanding insect–plant interactions and weed biocontrol

Structured host-choice and no-choice tests were conducted to help clarify the host plant interactions of an insect herbivore that is simultaneously seen as broadly polyphagous and pestiferous (in Africa) and host restricted/beneficial (in Australia). The research reported here involves specification of the host range of the invasive population of Scirtothrips aurantii found on Bryophyllum in Australia and included tests involving three separate lists of plant species considered to have the potential for thrips attack (plants of horticultural concern, native species at risk of attack and species listed for screening in the search for specialist B. delagoense biocontrol agents). This procedure was developed specifically to deal with the S. aurantii situation in Australia. Because the test species is already present in the field, the conclusions from the tests could be evaluated independently against field sampling results. Host testing revealed that the fundamental host range of the Bryophyllum population of S. aurantii includes Macadamia integrifolia, Mangifera indica and Kalanchoe blossfeldianna. However, the choice tests (involving B. delagoense) and a field survey of Man. indica demonstrated conclusively that the realised host range of S. aurantii in the field is restricted to Crassulaceae. We recommend that host testing of generalist insects not be discounted out of hand (for biological control) because of their perceived polyphagy. Any evidence of populations being strongly associated with the weed species of interest (through quantified host association studies in the native range) suggests further scrutiny of that population is warranted, by means of the host testing methods developed here and in conjunction with appropriate tests of the population’s species status.     

Structure of plant–Hymenoptera networks in two coastal shrub sites in Mexico

The study of interaction networks between plants and pollinators allows us to explore interaction patterns at the community level, detect changes in visit frequency and evaluate the nestedness of the networks. The latter allows rare plant species to be visited by more abundant species of pollinators, potentially allowing community diversity to be maintained, and this approach makes it possible to discern the rewiring (changes in connections) of species when their preferred resource is not available. In this study, the topology, species identity and rewiring were compared between two contrasting sites, one within a conservation area and the other subjected to continuous disturbance. The networks of both sites were significantly nested and shared a high number of common species of both plants and pollinators. However, the sites differed notably in the number of exclusive interactions, suggesting a high percentage of interaction rewiring. The introduced bee species, Apis mellifera, was the most frequent species at both sites and also the most connected in terms of the number of its interactions. This is explained by its generalist foraging characteristics that allow it to form part of the networks’ core group. In general, our results underscore the importance of knowing the identity of the participating species when studying networks, and how connections change between them, as well as the potential effect of habitat destruction and the role of invasive species in the rearrangement of the interactions; all factors that can exert an influence on the functioning of plant–pollinator networks.     

Antagonistic effects of floral scent in an insect–plant interaction

In southwestern USA, the jimsonweed Datura wrightii and the nocturnal moth Manduca sexta form a pollinator–plant and herbivore–plant association. Because the floral scent is probably important in mediating this interaction, we investigated the floral volatiles that might attract M. sexta for feeding and oviposition. We found that flower volatiles increase oviposition and include small amounts of both enantiomers of linalool, a common component of the scent of hawkmoth-pollinated flowers. Because (+)-linalool is processed in a female-specific glomerulus in the primary olfactory centre of M. sexta, we hypothesized that the enantiomers of linalool differentially modulate feeding and oviposition. Using a synthetic mixture that mimics the D. wrightii floral scent, we found that the presence of linalool was not necessary to evoke feeding and that mixtures containing (+)- and/or (−)-linalool were equally effective in mediating this behaviour. By contrast, females oviposited more on plants emitting (+)-linalool (alone or in mixtures) over control plants, while plants emitting (−)-linalool (alone or in mixtures) were less preferred than control plants. Together with our previous investigations, these results show that linalool has differential effects in feeding and oviposition through two neural pathways: one that is sexually isomorphic and non-enantioselective, and another that is female-specific and enantioselective.     

Floral divergence, pollinator partitioning and the spatiotemporal pattern of plant–pollinator interactions in three sympatric Adenophora species

Floral divergence among congeners may relate to differential utilization of pollinators and contribute to reducing overlap in pollination niches. To investigate whether and how floral differences are associated with differential utilization of pollinators in three sympatric Adenophora species, we analyzed floral traits and evaluated the contribution of different visitors to pollination. We compared visitation rates of different pollinator categories in different years and sites. A suite of floral traits differed among the three Adenophora species, suggesting adaptation to diurnal versus nocturnal pollination and an intermediate condition. However, many visitor species were shared among the three plant species, suggesting that floral traits did not rigorously filter visitors. Effective pollinators were large bees and moths. The importance of large bees as pollinators decreased whereas that of moths increased along the gradient from typically bee-pollinated to moth-pollinated flowers. The intermediate species (A. khasiana) differed substantially from the other two species in pollinator species but not in pollinator categories. The principal pollinator category of each species was constant across years and sites except in the intermediate species where it differed between two sites. Overall, the three sympatric species of Adenophora partition pollinators by floral divergence and the principal pollinators coincide with the predictions based on floral syndromes.     

Role of plant myosins in motile organelles: Is a direct interaction required?

Plant organelles are highly motile, with speed values of 3–7 m m/s in cells of land plants and about20–60 m m/s in characean algal cells. This movement is believed to be important for rapid distribution of materials around the cell, for the plant's ability to respond to environmental biotic and abiotic signals and for proper growth. The main machinery that propels motility of organelles within plant cells is based on the actin cytoskeleton and its motor proteins the myosins.Most plants express multiple members of two main classes:myosin VIII and myosin XI. While myosin VIII has been characterized as a slow motor protein, myosins from class XI were found to be the fastest motor proteins known in al kingdoms. Paradoxically, while it was found that myosins from class XI regulate most organelle movement, it is not quite clear how or even if these motor proteins attach to the organelles whose movement they regulate.