Elevation‐dependent effects of forest fragmentation on plant–bird interaction networks in the tropical Andes

Tropical forests harbor diverse ecological communities of plants and animals that are organized in complex interaction networks. The diversity and structure of plant–animal interaction networks may change along elevational gradients and in response to human‐induced habitat fragmentation. While previous studies have analyzed the effects of elevation and forest fragmentation on species interaction networks in isolation, to our knowledge no study has investigated whether the effects of forest fragmentation on species interactions may differ along elevational gradients. In this study, we analyzed main and interaction effects of elevation and forest fragmentation on plant–frugivore interaction networks at plant and bird species level. Over a period spanning two years, we recorded plant–frugivore interactions at three elevations (1000, 2000 and 3000 m a.s.l.) and in two habitat types (continuous and fragmented forest) in tropical montane forests in southern Ecuador. We found a consistent effect of elevation on the structure of plant–frugivore networks. We observed a decrease in the number of effective bird partners of plants and, thus, a decline in the redundancy of bird species with increasing elevation. Furthermore, bird specialization on specific plant partners increased towards high elevations. Fragmentation had a relatively weak effect on the interaction networks for both plant and bird species, but resulted in a significant increase in bird specialization in fragmented forests at high elevations. Our results indicate that forest fragmentation may have stronger effects on plant–frugivore interaction networks at high compared to low elevations because bird species richness declined more steeply towards high elevations than plant species richness. We conclude that conservation efforts should prioritize the maintenance of consumer diversity, for instance by maintaining stretches of continuous forest. This applies in particular to species‐poor communities, such as those at high elevations, as the ecological processes in these communities seem most sensitive towards forest fragmentation.     

Elevational trends of biodiversity and plant traits do not converge—a test in the Helan Range, NW China

When elevational gradients include combinations of different climatic gradients, such as a decline in temperature combined with an increase in moisture, vegetation and plant trait responses are difficult to explain. Here, we used plant species richness and morphometric traits data across steep elevational gradients in the Helan Range of Northwestern China in an attempt to separate general trends (temperature related) from regional peculiarity (moisture related). Based on the floristic data of the Helan Range as well as plot-based data, both drought (at low elevation) and low temperature (high elevation) are associated with low species richness presumably also explaining the peak in diversity at mid-elevation, where climatic conditions are moderate. However, this mid-elevation peak in diversity is not mirrored in trends of plant traits such as leaf size and inflorescence size, which show either unidirectional trends or no change with elevation (with impacts of drought and low temperature perhaps gradually replacing each other). Our analysis illustrated the taxonomic and plant functional type (PFT)-related biases in functional trait studies and showed that consistent patterns only emerge after careful data stratification, with taxonomy (family level) holding more promises than PFTs. Inflorescence size increased with elevation in major insect-pollinated families, a trend not seen in wind-pollinated (graminoid) families. However, the reproductive effort expressed as inflorescence/leaf length ratio increases with elevation in the majority of plant families, irrespective of their pollination system. The fact that these biometric responses to elevation do not correlate with responses in species richness (which peaks at mid-elevation) may reflect contrasting drivers of trait selection and biodiversity. Based on our plot-based data, this analysis also confirmed the usefulness of floristic archive data for testing ecological theory related to elevational gradients.     

Refuges from fire maintain pollinator–plant interaction networks

Fire is a major disturbance factor in many terrestrial ecosystems, leading to landscape transformation in fire‐prone areas. Species in mutualistic interactions are often highly sensitive to disturbances like fire events, but the degree and complexity of their responses are unclear. We use bipartite insect–flower interaction networks across a recently burned landscape to explore how plant–pollinator interaction networks respond to a recent major fire event at the landscape level, and where fire refuges were present. We also investigate the effectiveness of these refuges at different elevations (valley to hilltop) for the conservation of displaced flower‐visiting insects during fire events. Then, we explore how the degree of specialization of flower‐visiting insects changes across habitats with different levels of fire impact. We did this in natural areas in the Greater Cape Floristic Region (GCFR) biodiversity hotspot, which is species rich in plants and pollinators. Bees and beetles were the most frequent pollinators in interactions, followed by wasps and flies. Highest interaction activity was in the fire refuges and least in burned areas. Interactions also tracked flower abundance, which was highest in fire refuges in the valley and lowest in burned areas. Interactions consisted mostly of specialized flower visitors, especially in refuge areas. The interaction network and species specialization were lowest in burned areas. However, species common to at least two fire classes showed no significant difference in species specialization. We conclude that flower‐rich fire refuges sustain plant–pollinator interactions, especially those involving specialized species, in fire‐disturbed landscape. This may be an important shelter for specialized pollinator species at the time that the burned landscape goes through regrowth and succession as part of ecosystem recovery process after a major fire event.     

Pyrodiversity promotes interaction complementarity and population resistance

Theory predicts that network characteristics may help anticipate how populations and communities respond to extreme climatic events, but local environmental context may also influence responses to extreme events. For example, altered fire regimes in many ecosystems may significantly affect the context for how species and communities respond to changing climate. In this study, I tested whether the responses of a pollinator community to extreme drought were influenced by the surrounding diversity of fire histories (pyrodiversity) which can influence their interaction networks via changing partner availability. I found that at the community level, pyrodiverse landscapes promote functional complementarity and generalization, but did not consistently enhance functional redundancy or resistance to simulated co‐extinction cascades. Pyrodiversity instead supported flexible behaviors that enable populations to resist perturbations. Specifically, pollinators that can shift partners and network niches are better able to take advantage of the heterogeneity generated by pyrodiversity, thereby buffering pollinator populations against changes in plant abundances. These findings suggest that pyrodiversity is unlikely to improve community‐level resistance to droughts, but instead promotes population resistance and community functionality. This study provides unique evidence that resistance to extreme climatic events depends on both network properties and historical environmental context.     

Predicting the effect of competition on secondary plant extinctions in plant–pollinator networks

What are the limitations of models that predict the behavior of an ecological community based on a single type of species interaction? Using plant–pollinator network models as an example, we contrast the predicted vulnerability of a community to secondary extinctions under the assumption of purely mutualistic interactions versus mutualistic and competitive interactions. We find that competition among plant species increases the risk of secondary extinctions and extinction cascades. Simulations over a number of different network structures indicate that this effect is stronger in larger networks, more strongly connected networks and networks with higher plant:pollinator ratios. We conclude that efforts to model plant–pollinator communities will systematically over‐estimate community robustness to species loss if plant competition is ignored. However, because the effect of plant competition depends on network architecture, and because characterization of plant competition is work intensive, we suggest that efforts to account for plant competition in plant–pollinator network models should be focused on large, strongly connected networks with high plant:pollinator ratios.     

Bird and insect pollinators differ in specialization and potential pollination services along disturbance and resource gradients

Combined studies of the communities and interaction networks of bird and insect pollinators are rare, especially along environmental gradients. Here, we determined how disturbance by fire and variation in sugar resources shape pollinator communities and interactions between plants and their pollinating insects and birds. We recorded insect and bird visits to 21 Protea species across 21 study sites and for 2 years in Fynbos ecosystems in the Western Cape, South Africa. We recorded morphological traits of all pollinator species (41 insect and nine bird species). For each site, we obtained estimates of the time since the last fire (range: 2–25 calendar years) and the Protea nectar sugar amount per hectare (range: 74–62 000 g/ha). We tested how post-fire age and sugar amount influence the total interaction frequency, species richness and functional diversity of pollinator communities, as well as pollinator specialization (the effective number of plant partners) and potential pollination services (pollination service index) of insects and birds. We found little variation in the total interaction frequency, species richness and functional diversity of insect and bird pollinator communities, but insect species richness increased with post-fire age. Pollinator specialization and potential pollination services of insects and birds varied differently along the environmental gradients. Bird pollinators visited fewer Protea species at sites with high sugar amount, while there was no such trend for insects. Potential pollination services of insect pollinators to Protea species decreased with increasing post-fire age and resource amounts, whereas potential pollination services of birds remained constant along the environmental gradients. Despite little changes in pollinator communities, our analyses reveal that insect and bird pollinators differ in their specialization on Protea species and show distinct responses to disturbance and resource gradients. Our comparative study of bird and insect pollinators demonstrates that birds may be able to provide more stable pollination services than insects.     

Combined Effects of Extreme Climatic Events and Elevation on Nutritional Quality and Herbivory of Alpine Plants

Climatic extreme events can cause the shift or disruption of plant-insect interactions due to altered plant quality, e.g. leaf carbon to nitrogen ratios, and phenology. However, the response of plant-herbivore interactions to extreme events and climatic gradients has been rarely studied, although climatic extremes will increase in frequency and intensity in the future and insect herbivores represent a highly diverse and functionally important group. We set up a replicated climate change experiment along elevational gradients in the German Alps to study the responses of three plant guilds and their herbivory by insects to extreme events (extreme drought, advanced and delayed snowmelt) versus control plots under different climatic conditions on 15 grassland sites. Our results indicate that elevational shifts in CN (carbon to nitrogen) ratios and herbivory depend on plant guild and season. CN ratios increased with altitude for grasses, but decreased for legumes and other forbs. In contrast to our hypotheses, extreme climatic events did not significantly affect CN ratios and herbivory. Thus, our study indicates that nutritional quality of plants and antagonistic interactions with insect herbivores are robust against seasonal climatic extremes. Across the three functional plant guilds, herbivory increased with nitrogen concentrations. Further, increased CN ratios indicate a reduction in nutritional plant quality with advancing season. Although our results revealed no direct effects of extreme climatic events, the opposing responses of plant guilds along elevation imply that competitive interactions within plant communities might change under future climates, with unknown consequences for plant-herbivore interactions and plant community composition.     

Effects of hummingbird morphology on specialization in pollination networks vary with resource availability

Specialization of species in interaction networks influences network stability and ecosystem functioning. Spatial and temporal variation in resource availability may provide insight into how ecological factors, such as resource abundance, and evolutionary factors, such as phylogenetically conserved morphological traits, influence specialization within mutualistic networks. We used independent measures of hummingbird abundance and resources (nectar), information on hummingbird traits and plant–hummingbird interactions to examine how resource availability and species' morphology influence the specialization of hummingbirds in three habitat types (forest, shrubs, cattle ranch) sampled over 10 sessions across two years in the southern Andes of Ecuador. Specialization of hummingbird species in the networks was measured by three indices: d' (related to niche partitioning), generality (related to niche width) and PSI (related to pollination services). Specialization indices d', generality and PSI of hummingbird species were influenced by resource availability. All indices indicated that specialization of hummingbirds increased when the availability of resources decreased. Variation in d' was also explained by an interaction between resource availability and bill length; hummingbirds with a long bill switched from being more specialized than other species when resource availability was low to being similarly specialized when availability was high. Overall, our results highlight the importance of ecological and evolutionary factors determining the specialization of species in interaction networks. We demonstrate in particular that ecological gradients in resource availability cause substantial changes in consumers' foraging behavior contingent on their morphology. Changes in pollinator specialization along resource gradients can have impacts on ecosystem functions, such as pollination by animals.     

What plant–pollinator network structure tells us about the mechanisms underlying the bidirectional biodiversity productivity relationship?

The Biodiversity – Ecosystem Functioning (B–EF) relationship remains a topic of ongoing debate with most studies focusing on primary productivity, and documenting that this relationship takes many forms. It remains unclear if biodiversity drives productivity or productivity shapes biodiversity or the relationship is bidirectional. B-EF studies explore almost exclusively the relationship between species richness and ecosystem functioning, while the role of biotic interactions, a key component of ecosystem functioning, has been neglected. Here, using data of 80 local plant–pollinator networks on 20 Aegean islands, and of gross primary productivity (GPP) from the MODIS satellite, we explored the bidirectional relationship between interaction network structure (nestedness and specialization), species richness (plants and pollinators) and mean and inter-annual variability of GPP. We found that nestedness and specialisation of plant–pollinator networks is driven by mean GPP. However, specialisation alone was a significant predictor of mean GPP, implying that networks tend to be more specialised in low-productivity areas. Pollinator species richness exerted a strong effect on mean GPP with the remaining factors playing a minor role, while the effect of mean GPP on pollinator species richness was weaker. Furthermore, the nestedness of plant–pollinator networks drives inter-annual variability of GPP with more nested networks displaying less variability, which is in accordance with the predictions of the insurance hypothesis. Plant and pollinator species richness were also associated with inter-annual variability of GPP.     

Additive effects of exotic plant abundance and land-use intensity on plant–pollinator interactions

The continuing spread of exotic plants and increasing human land-use are two major drivers of global change threatening ecosystems, species and their interactions. Separate effects of these two drivers on plant–pollinator interactions have been thoroughly studied, but we still lack an understanding of combined and potential interactive effects. In a subtropical South African landscape, we studied 17 plant–pollinator networks along two gradients of relative abundance of exotics and land-use intensity. In general, pollinator visitation rates were lower on exotic plants than on native ones. Surprisingly, while visitation rates on native plants increased with relative abundance of exotics and land-use intensity, pollinator visitation on exotic plants decreased along the same gradients. There was a decrease in the specialization of plants on pollinators and vice versa with both drivers, regardless of plant origin. Decreases in pollinator specialization thereby seemed to be mediated by a species turnover towards habitat generalists. However, contrary to expectations, we detected no interactive effects between the two drivers. Our results suggest that exotic plants and land-use promote generalist plants and pollinators, while negatively affecting specialized plant–pollinator interactions. Weak integration and high specialization of exotic plants may have prevented interactive effects between exotic plants and land-use. Still, the additive effects of exotic plants and land-use on specialized plant–pollinator interactions would have been overlooked in a single-factor study. We therefore highlight the need to consider multiple drivers of global change in ecological research and conservation management.     

Anthropogenic disturbance and rainfall variation threaten the stability of plant–ant interactions in the Brazilian Caatinga

Climate change is projected to exacerbate the effects of anthropogenic disturbance, with negative impacts on ecosystem stability and functioning. We evaluate the additive and combined effects of chronic anthropogenic disturbance (CAD) and rainfall variation on the temporal stability of mutualistic EFN‐bearing plant–ant networks in a Caatinga dry forest. We evaluated whether changes in the stability of these interactions are driven by changes in the stability of the communities of partners involved and/or in ant behavior. We sampled EFN‐bearing plant–ant networks in sixteen 20 × 20 m plots distributed across CAD and rainfall gradients. The stability of EFN‐bearing plant and attendant–ant communities were measured as the inverse of temporal differences in their community structure and composition. We also computed the stability of EFN‐bearing plant–ant networks by measuring the inverse of temporal differences in network specialization metrics. We found that, in general, the structure and composition of plant and ant interacting communities were similarly stable along both environmental gradients. Only CAD and its interaction with rainfall affected the temporal stability of EFN‐bearing plant diversity, which declined as CAD increased, with a more pronounced relationship in wetter areas. However, variation in levels of CAD and, to a lesser extent, rainfall greatly modulated the stability of EFN‐bearing plant–ant network specialization. CAD reduced the stability of network generality (specialization at the ant level), an effect that was much stronger in wetter areas. Meanwhile, the stability in network vulnerability (specialization at the plant level) decreased with the increase of CAD and the decrease of rainfall levels. Finally, there was a trend of decreasing stability in specialization of the overall network with increasing CAD. Our results suggest that changes in the structure of interaction networks are mainly driven by a switch in ant behavior rather than by changes in the structure and composition of plant and ant communities between years.     

Year‐to‐year variation in the topology of a plant–pollinator interaction network

Vigorous discussion of the degree of specialization in pollination interactions, combined with advances in the analysis of complex networks, has revitalized the study of entire plant–pollinator communities. Noticeably rare, however, are attempts to quantify temporal variation in the structure of plant–pollinator networks, and to determine whether the status of species as specialists or generalists is stable. Here we show that network structure varied through time in a montane meadow community from southern California, USA, in that pollinator species did not form the same links with plant species across years. Furthermore, composition of the generalized core group of species in the network varied among summers, as did the identity of those species involved in relationships that appeared to be reciprocally specialized within any one summer. These differences appear to be related to severe drought conditions experienced in the second summer of the 3 year study. In contrast to this variation, the pollinator community remained similarly highly nested in all three summers, even though species were packed into the nested matrix differently from year to year. These results suggest that plant–pollinator networks vary in detail through time, while retaining some basic topological properties. This dynamic aspect of community‐scale interactions has implications for both ecological and evolutionary inferences about pollination mutualisms.     

The importance of interannual variation and bottom–up nitrogen enrichment for plant–pollinator networks

Striking changes in food web structure occur with alterations in resource supply. Like predator–prey interactions, many mutualisms are also consumer–resource interactions. However, no studies have explored how the structure of plant–pollinator networks may be affected by nutrient enrichment. For three years, we enriched plots of subalpine plant communities with nitrogen and observed subsequent effects on plant–pollinator network structure. Although nitrogen enrichment affects floral abundance and rates of pollinator visitation, we found no effects of nitrogen enrichment on the core group of generalist plants and pollinators or on plant–pollinator network structure parameters, such as network topology (the identity and frequency of interactions) and the degree of nestedness. However, individual plant and pollinator taxa were packed into the nested networks differently among nitrogen treatments. In particular, pollinators visited different numbers and types of plants in the nested networks, suggesting weak, widespread effects of nitrogen addition on individual taxa. Independent of nitrogen enrichment, there were large interannual differences in network structure and interactions, due to species turnover among years and flexibility in interacting with new partners. These data suggest that the community structure of small‐scale mutualistic networks may be relatively robust to short‐term bottom–up changes in the resource supply, but sensitive to variation in the opportunistic behavior and turnover of plant and pollinator species among years.     

Plant–pollinator networks: adding the pollinator's perspective

Pollination network studies are based on pollinator surveys conducted on focal plants. This plant-centred approach provides insufficient information on flower visitation habits of rare pollinator species, which are the majority in pollinator communities. As a result, pollination networks contain very high proportions of pollinator species linked to a single plant species (extreme specialists), a pattern that contrasts with the widely accepted view that plant–pollinator interactions are mostly generalized. In this study of a Mediterranean scrubland community in NE Spain we supplement data from an intensive field survey with the analysis of pollen loads carried by pollinators. We observed 4265 contacts corresponding to 19 plant and 122 pollinator species. The addition of pollen data unveiled a very significant number of interactions, resulting in important network structural changes. Connectance increased 1.43-fold, mean plant connectivity went from 18.5 to 26.4, and mean pollinator connectivity from 2.9 to 4.1. Extreme specialist pollinator species decreased 0.6-fold, suggesting that ecological specialization is often overestimated in plant–pollinator networks. We expected a greater connectivity increase in rare species, and consequently a decrease in the level of asymmetric specialization. However, new links preferentially attached to already highly connected nodes and, as a result, both nestedness and centralization increased. The addition of pollen data revealed the existence of four clearly defined modules that were not apparent when only field survey data were used. Three of these modules had a strong phenological component. In comparison to other pollination webs, our network had a high proportion of connector links and species. That is, although significant, the four modules were far from isolated.     

Species traits and abundances predict metrics of plant–pollinator network structure,but not pairwise interactions

Plant–pollinator mutualistic networks represent the ecological context of foraging (for pollinators) and reproduction (for plants and some pollinators). Plant–pollinator visitation networks exhibit highly conserved structural properties across diverse habitats and species assemblages. The most successful hypotheses to explain these network properties are the neutrality and biological constraints hypotheses, which posit that species interaction frequencies can be explained by species relative abundances, and trait mismatches between potential mutualists respectively. However, previous network analyses emphasize the prediction of metrics of qualitative network structure, which may not represent stringent tests of these hypotheses. Using a newly documented temporally explicit alpine plant–pollinator visitation network, we show that metrics of both qualitative and quantitative network structure are easy to predict, even by models that predict the identity or frequency of species interactions poorly. A variety of phenological and morphological constraints as well as neutral interactions successfully predicted all network metrics tested, without accurately predicting species observed interactions. Species phenology alone was the best predictor of observed interaction frequencies. However, all models were poor predictors of species pairwise interaction frequencies, suggesting that other aspects of species biology not generally considered in network studies, such as reproduction for dipterans, play an important role in shaping plant–pollinator visitation network structure at this site. Future progress in explaining the structure and dynamics of mutualistic networks will require new approaches that emphasize accurate prediction of species pairwise interactions rather than network metrics, and better reflect the biology underlying species interactions.     

The geographical variation of network structure is scale dependent: understanding the biotic specialization of host–parasitoid networks

Research on the structure of ecological networks suggests that a number of universal patterns exist. Historically, biotic specialization has been thought to increase towards the Equator. Yet, recent studies have challenged this view showing non‐conclusive results. Most studies analysing the geographical variation in biotic specialization focus, however, only on the local scale. Little is known about how the geographical variation of network structure depends on the spatial scale of observation (i.e. from local to regional spatial scales). This should be remedied, as network structure changes as the spatial scale of observation changes, and the magnitude and shape of these changes can elucidate the mechanisms behind the geographical variation in biotic specialization. Here we analyse four facets of biotic specialization in host–parasitoid networks along gradients of climatic constancy, classifying the networks according to their spatial extension (local or regional). Namely, we analyse network connectance, consumer diet overlap, consumer diet breadth, and resource vulnerability at both local and regional scales along the gradients of both current climatic constancy and historical climatic change. While at the regional scale none of the climatic variables are associated to biotic specialization, at the local scale, network connectance, consumer diet overlap, and resource vulnerability decrease with current climatic constancy, whereas consumer generalism increases (i.e. broader diet breadths in tropical areas). Similar patterns are observed along the gradient of historical climatic change. We provide an explanation based on different beta‐diversity for consumers and resources across the geographical gradients. Our results show that the geographical gradient of biotic specialization is not universal. It depends on both the facet of biotic specialization and the spatial scale of observation.     

Top‐down network analysis characterizes hidden termite–termite interactions

The analysis of ecological networks is generally bottom‐up, where networks are established by observing interactions between individuals. Emergent network properties have been indicated to reflect the dominant mode of interactions in communities that might be mutualistic (e.g., pollination) or antagonistic (e.g., host–parasitoid communities). Many ecological communities, however, comprise species interactions that are difficult to observe directly. Here, we propose that a comparison of the emergent properties from detail‐rich reference communities with known modes of interaction can inform our understanding of detail‐sparse focal communities. With this top‐down approach, we consider patterns of coexistence between termite species that live as guests in mounds built by other host termite species as a case in point. Termite societies are extremely sensitive to perturbations, which precludes determining the nature of their interactions through direct observations. We perform a literature review to construct two networks representing termite mound cohabitation in a Brazilian savanna and in the tropical forest of Cameroon. We contrast the properties of these cohabitation networks with a total of 197 geographically diverse mutualistic plant–pollinator and antagonistic host–parasitoid networks. We analyze network properties for the networks, perform a principal components analysis (PCA), and compute the Mahalanobis distance of the termite networks to the cloud of mutualistic and antagonistic networks to assess the extent to which the termite networks overlap with the properties of the reference networks. Both termite networks overlap more closely with the mutualistic plant–pollinator communities than the antagonistic host–parasitoid communities, although the Brazilian community overlap with mutualistic communities is stronger. The analysis raises the hypothesis that termite–termite cohabitation networks may be overall mutualistic. More broadly, this work provides support for the argument that cryptic communities may be analyzed via comparison to well‐characterized communities.     

Plant invasion and speciation along elevational gradients on the oceanic island La Palma,Canary Islands

Ecosystems that provide environmental opportunities but are poor in species and functional richness generally support speciation as well as invasion processes. These processes are expected not to be equally effective along elevational gradients due to specific ecological, spatial, and anthropogenic filters, thus controlling the dispersal and establishment of species. Here, we investigate speciation and invasion processes along elevational gradients. We assess the vascular plant species richness as well as the number and percentage of endemic species and non‐native species systematically along three elevational gradients covering large parts of the climatic range of La Palma, Canary Islands. Species richness was negatively correlated with elevation, while the percentage of Canary endemic species showed a positive relationship. However, the percentage of Canary–Madeira endemics did not show a relationship with elevation. Non‐native species richness (indicating invasion) peaked at 500 m elevation and showed a consistent decline until about 1,200 m elevation. Above that limit, no non‐native species were present in the studied elevational gradients. Ecological, anthropogenic, and spatial filters control richness, diversification, and invasion with elevation. With increase in elevation, richness decreases due to species–area relationships. Ecological limitations of native ruderal species related to anthropogenic pressure are in line with the absence of non‐native species from high elevations indicating directional ecological filtering. Increase in ecological isolation with elevation drives diversification and thus increased percentages of Canary endemics. The best preserved eastern transect, including mature laurel forests, is an exception. The high percentage of Canary–Madeira endemics indicates the cloud forest's environmental uniqueness—and thus ecological isolation—beyond the Macaronesian islands.     

Drought negatively affects communities on a foundation tree: growth rings predict diversity

Understanding how communities respond to extreme climatic events is important for predicting the impact of climate change on biodiversity. The plant vigor and stress hypotheses provide a theoretical framework for understanding how arthropods respond to stress, but are rarely tested at the community level. Following a record drought, we compared the communities of arthropods on pinyon pine (Pinus edulis) that exhibited a gradient in physical traits related to environmental stress (e.g., growth rate, branch dieback, and needle retention). Six patterns emerged that show how one of the predicted outcomes of climate change in the southwestern USA (i.e., increased drought severity) alters the communities of a foundation tree species. In accordance with the plant vigor hypothesis, increasing tree stress was correlated with an eight to tenfold decline in arthropod species richness and abundance. Trees that were more similar in their level of stress had more similar arthropod communities. Both foliage quantity and quality contributed to arthropod community structure. Individual species and feeding groups differed in their responses to plant stress, but most were negatively affected. Arthropod richness (r ² = 0.48) and abundance (r ² = 0.48) on individual trees were positively correlated with the tree’s radial growth during drought. This relationship suggests that tree ring analysis may be used as a predictor of arthropod diversity, which is similar to findings with ectomycorrhizal fungi. A contrast of our findings on arthropod abundance with published data on colonization by mutualistic fungi on the same trees demonstrates that at low stress these two communities respond differently, but at high stress both are negatively affected. These results suggest that the effect of extreme climatic events such as drought on foundation tree species are likely to decrease multi-trophic diversity and shift arthropod community composition, which in turn could cascade to affect other associated taxa.     

Contrasting effects of land‐use changes on herbivory and pollination networks

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