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Aim Global warming is assumed to restructure mountain insect communities in space and time. Theory and observations along climate gradients predict that insect abundance and richness, especially of small‐bodied species, will increase with increasing temperature. However, the specific responses of single species to rising temperatures, such as spatial range shifts, also alter communities, calling for intensive monitoring of real‐world communities over time. Location German Alps and pre‐alpine forests in south‐east Germany. Methods We empirically examined the temporal and spatial change in wild bee communities and its drivers along two largely well‐protected elevational gradients (alpine grassland vs. pre‐alpine forest), each sampled twice within the last decade. Results We detected clear abundance‐based upward shifts in bee communities, particularly in cold‐adapted bumble bee species, demonstrating the speed with which mobile organisms can respond to climatic changes. Mean annual temperature was identified as the main driver of species richness in both regions. Accordingly, and in large overlap with expectations under climate warming, we detected an increase in bee richness and abundance, and an increase in small‐bodied species in low‐ and mid‐elevations along the grassland gradient. Community responses in the pre‐alpine forest gradient were only partly consistent with community responses in alpine grasslands. Main Conclusion In well‐protected temperate mountain regions, small‐bodied bees may initially profit from warming temperatures, by getting more abundant and diverse. Less severe warming, and differences in habitat openness along the forested gradient, however, might moderate species responses. Our study further highlights the utility of standardized abundance data for revealing rapid changes in bee communities over only one decade.

Environmental gradients generate and maintain biodiversity on Earth. Mountain slopes are among the most pronounced terrestrial environmental gradients, and the elevational structure of species and their interactions can provide unique insight into the processes that govern community assembly and function in mountain ecosystems. We recorded bumble bee–flower interactions over 3 years along a 1400‐m elevational gradient in the German Alps. Using nonlinear modeling techniques, we analyzed elevational patterns at the levels of abundance, species richness, species β‐diversity, and interaction β‐diversity. Though floral richness exhibited a midelevation peak, bumble bee richness increased with elevation before leveling off at the highest sites, demonstrating the exceptional adaptation of these bees to cold temperatures and short growing seasons. In terms of abundance, though, bumble bees exhibited divergent species‐level responses to elevation, with a clear separation between species preferring low versus high elevations. Overall interaction β‐diversity was mainly caused by strong turnover in the floral community, which exhibited a well‐defined threshold of β‐diversity rate at the tree line ecotone. Interaction β‐diversity increased sharply at the upper extreme of the elevation gradient (1800–2000 m), an interval over which we also saw steep decline in floral richness and abundance. Turnover of bumble bees along the elevation gradient was modest, with the highest rate of β‐diversity occurring over the interval from low‐ to mid‐elevation sites. The contrast between the relative robustness bumble bee communities and sensitivity of plant communities to the elevational gradient in our study suggests that the strongest effects of climate change on mountain bumble bees may be indirect effects mediated by the responses of their floral hosts, though bumble bee species that specialize in high‐elevation habitats may also experience significant direct effects of warming.

Animal pollinators are globally threatened by anthropogenic land use change and agricultural intensification. The yield of many food crops is therefore negatively impacted because they benefit from biotic pollination. This is especially the case in the tropics. For instance, fruit set of Coffea arabica has been shown to increase by 10–30% in plantations with a high richness of bee species, possibly influenced by the availability of surrounding forest habitat. Here, we performed a global literature review to (1) assess how much animal pollination enhances coffee fruit set, and to (2) examine the importance of the amount of forest cover, distance to nearby forest and forest canopy density for bee species richness and coffee fruit set. Using a systematic literature review, we identified eleven case studies with a total of 182 samples where fruit set of C. arabica was assessed. We subsequently gathered forest data for all study sites from satellite imagery. We modelled the effects of open (all forest with a canopy density of ≥25%), closed (≥50%) and dense (≥75%) forests on pollinator richness and fruit set of coffee. Overall, we found that animal pollination increases coffee fruit set by ~18% on average. In only one of the case studies, regression results indicate a positive effect of dense forest on coffee fruit set, which increased with higher forest cover and shorter distance to the forest. Against expectations, forest cover and distance to open forest were not related to bee species richness and fruit set. In summary, we provide strong empirical support for the notion that animal pollinators increase coffee fruit set. Forest proximity had little overall influence on bee richness and coffee fruit set, except when farms were surrounded by dense tropical forests, potentially because these may provide high-quality habitats for bees pollinating coffee. We, therefore, advocate that more research is done to understand the biodiversity value of dense forest for pollinators, notably assessing the mechanisms underlying the importance of forest for pollinators and their pollination services.

Patterns of resource use by animals can clarify how ecological communities have assembled in the past, how they currently function and how they are likely to respond to future perturbations. Bumble bees (Hymentoptera: Bombus spp.) and their floral hosts provide a diverse yet tractable system in which to explore resource selection in the context of plant–pollinator networks. Under conditions of resource limitation, the ability of bumble bees species to coexist should depend on dietary niche overlap. In this study, we report patterns and dynamics of floral morphotype preferences in a mountain bumble bee community based on ~13 000 observations of bumble bee floral visits recorded along a 1400 m elevation gradient. We found that bumble bees are highly selective generalists, rarely visiting floral morphotypes at the rates predicted by their relative abundances. Preferences also differed markedly across bumble bee species, and these differences were well-explained by variation in bumble bee tongue length, generating patterns of preference similarity that should be expected to predict competition under conditions of resource limitation. Within species, though, morphotype preferences varied by elevation and season, possibly representing adaptive flexibility in response to the high elevational and seasonal turnover of mountain floral communities. Patterns of resource partitioning among bumble bee communities may determine which species can coexist under the altered distributions of bumble bees and their floral hosts caused by climate and land use change.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown1-3. Conclusions that are derived from studies in non-mountainous areas are not suitable for predicting the effects of land-use changes on tropical mountains because the climatic environment rapidly changes with elevation, which may mitigate or amplify the effects of land use4,5. It is of key importance to understand how the interplay of climate and land use constrains biodiversity and ecosystem functions to determine the consequences of global change for mountain ecosystems. Here we show that the interacting effects of climate and land use reshape elevational trends in biodiversity and ecosystem functions on Africa's largest mountain, Mount Kilimanjaro (Tanzania). We find that increasing land-use intensity causes larger losses of plant and animal species richness in the arid lowlands than in humid submontane and montane zones. Increases in land-use intensity are associated with significant changes in the composition of plant, animal and microorganism communities; stronger modifications of plant and animal communities occur in arid and humid ecosystems, respectively. Temperature, precipitation and land use jointly modulate soil properties, nutrient turnover, greenhouse gas emissions, plant biomass and productivity, as well as animal interactions. Our data suggest that the response of ecosystem functions to land-use intensity depends strongly on climate; more-severe changes in ecosystem functioning occur in the arid lowlands and the cold montane zone. Interactions between climate and land use explained-on average-54% of the variation in species richness, species composition and ecosystem functions, whereas only 30% of variation was related to single drivers. Our study reveals that climate can modulate the effects of land use on biodiversity and ecosystem functioning, and points to a lowered resistance of ecosystems in climatically challenging environments to ongoing land-use changes in tropical mountainous regions.

Experimental exclosure of birds and bats constitutes a powerful tool to study the impacts of wildlife on pests and crop yields in agricultural systems. Though widely utilized, exclosure experiments are not standardized across studies. Indeed, key differences surrounding the design, materials, and protocols for implementing field-based exclosure experiments of flying vertebrates increase heterogeneity across studies, and limit our understanding of biodiversity-friendly land use management. We reviewed the available literature on studies in which bird and bat exclosures were applied to study pest control in agricultural settings, and isolated 30 studies from both tropical and temperate land use systems, involving 12 crop types across 14 countries. Focusing on exclosure effects on crop yield, we analyzed effect detectability for a subset of suitable data. We then analyzed the potential of exclosure methods and possible extensions to improve our understanding of complex food webs and ecosystem services affecting the productivity of agricultural systems. While preferences exist in materials (e.g., nylon nets and bamboo frames), experimental exclosure studies of birds and bats differed greatly in their respective design, related costs, and effort — limiting the generalization and transferability of results at larger spatial scales. Most studies were based on experiments conducted in the United States and the Neotropics, mainly in coffee and cacao farms. A lack of preliminary or long-term data with repeated measurements makes it impossible to apply power analysis in most studies. Common constraints include, among other things, the choice of material and experimental duration, as well as the consideration of local versus landscape factors. We discuss such limitations, related common pitfalls, and options for optimization to inform improved planning, design, and execution of exclosure studies. By doing so, we aim to promote more comparable and transferable approaches in future field research on biodiversity-mediated ecosystem services.

2017-2018 Belmont Forum and BiodivERsA joint call for research proposals, under the BiodivScen ERA-Net COFUND programme, and with the funding organisations AEI, NWO, ECCyT and NSF Funding information, Allen-Perkins A, Magrach A, Dainese M, Garibaldi LA, Kleijn D, Rader R, Reilly JR, Winfree R, Lundin O, McGrady CM, Brittain C, Biddinger DJ, Artz DR, Elle E, Hoffman G, Ellis JD, Daniels J, Gibbs J, Campbell JW, Brokaw J, Wilson JK, Mason K, Ward KL, Gundersen KB, Bobiwash K, Gut L, Rowe LM, Boyle NK, Williams NM, Joshi NK, Rothwell N, Gillespie RL, Isaacs R, Fleischer SJ, Peterson SS, Rao S, Pitts-Singer TL, Fijen T, Boreux V, Rundlöf M, Viana BF, Klein AM, Smith HG, Bommarco R, Carvalheiro LG, Ricketts TH, Ghazoul J, Krishnan S, Benjamin FE, Loureiro J, Castro S, Raine NE, de Groot GA, Horgan FG, Hipólito J, Smagghe G, Meeus I, Eeraerts M, Potts SG, Kremen C, García D, Miñarro M, Crowder DW, Pisanty G, Mandelik Y, Vereecken NJ, Leclercq N, Weekers T, Lindstrom SAM, Stanley DA, Zaragoza-Trello C, Nicholson CC, Scheper J, Rad C, Marks EAN, Mota L, Danforth B, Park M, Bezerra ADM, Freitas BM, Mallinger RE, Oliveira da Silva F, Willcox B, Ramos DL, D da Silva E Silva F, Lázaro A, Alomar D, González-Estévez MA, Taki H, Cariveau DP, Garratt MPD, Nabaes Jodar DN, Stewart RIA, Ariza D, Pisman M, Lichtenberg EM, Schüepp C, Herzog F, Entling MH, Dupont YL, Michener CD, Daily GC, Ehrlich PR, Burns KLW, Vilà M, Robson A, Howlett B, Blechschmidt L, Jauker F, Schwarzbach F, Nesper M, Diekötter T, Wolters V, Castro H, Gaspar H, Nault BA, Badenhausser I, Petersen JD, Tscharntke T, Bretagnolle V, Willis Chan DS, Chacoff N, Andersson GKS, Jha S, Colville JF, Veldtman R, Coutinho J, Bianchi FJJA, Sutter L, Albrecht M, Jeanneret P, Zou Y, Averill AL, Saez A, Sciligo AR, Vergara CH, Bloom EH, Oeller E, Badano EI, Loeb GM, Grab H, Ekroos J, Gagic V, Cunningham SA, Åström J, Cavigliasso P, Trillo A, Classen A, Mauchline AL, Montero-Castaño A, Wilby A, Woodcock BA, Sidhu CS, Steffan-Dewenter I, Vogiatzakis IN, Herrera JM, Otieno M, Gikungu MW, Cusser SJ, Nauss T, Nilsson L, Knapp J, Ortega-Marcos JJ, González JA, Osborne JL, Blanche R, Shaw RF, Hevia V, Stout J, Arthur AD, Blochtein B, Szentgyorgyi H, Li J, Mayfield MM, Woyciechowski M, Nunes-Silva P, Halinski de Oliveira R, Henry S, Simmons BI, Dalsgaard B, Hansen K, Sritongchuay T, O'Reilly AD, Chamorro García FJ, Nates Parra G, Magalhães Pigozo C, Bartomeus I

Many experiments have shown that biodiversity enhances ecosystem functioning. However, we have little understanding of how environmental heterogeneity shapes the effect of diversity on ecosystem functioning and to what extent this diversity effect is mediated by variation in species richness or species turnover. This knowledge is crucial to scaling up the results of experiments from local to regional scales. Here we quantify the diversity effect and its components-that is, the contributions of variation in species richness and species turnover-for 22 ecosystem functions of microorganisms, plants and animals across 13 major ecosystem types on Mt Kilimanjaro, Tanzania. Environmental heterogeneity across ecosystem types on average increased the diversity effect from explaining 49% to 72% of the variation in ecosystem functions. In contrast to our expectation, the diversity effect was more strongly mediated by variation in species richness than by species turnover. Our findings reveal that environmental heterogeneity strengthens the relationship between biodiversity and ecosystem functioning and that species richness is a stronger driver of ecosystem functioning than species turnover. Based on a broad range of taxa and ecosystem functions in a non-experimental system, these results are in line with predictions from biodiversity experiments and emphasize that conserving biodiversity is essential for maintaining ecosystem functioning.

1. Among the many concerns for biodiversity in the Anthropocene, recent reports of flying insect loss are particularly alarming, given their importance as pollinators, pest control agents, and as a food source. Few insect monitoring programmes cover the large spatial scales required to provide more generalizable estimates of insect responses to global change drivers. 2. We ask how climate and surrounding habitat affect flying insect biomass using data from the first year of a new monitoring network at 84 locations across Germany comprising a spatial gradient of land cover types from protected to urban and crop areas. 3. Flying insect biomass increased linearly with temperature across Germany. However, the effect of temperature on flying insect biomass flipped to negative in the hot months of June and July when local temperatures most exceeded long-term averages. 4. Land cover explained little variation in insect biomass, but biomass was lowest in forests. Grasslands, pastures, and orchards harboured the highest insect biomass. The date of peak biomass was primarily driven by surrounding land cover, with grasslands especially having earlier insect biomass phenologies. 5. Standardised, large-scale monitoring provides key insights into the underlying processes of insect decline and is pivotal for the development of climate-adapted strategies to promote insect diversity. In a temperate climate region, we find that the positive effects of temperature on flying insect biomass diminish in a German summer at locations where temperatures most exceeded long-term averages. Our results highlight the importance of local adaptation in climate change-driven impacts on insect communities.

Aim Species differ in their degree of specialization when interacting with other species, with significant consequences for the function and robustness of ecosystems. In order to better estimate such consequences, we need to improve our understanding of the spatial patterns and drivers of specialization in interaction networks. Methods Here, we used the extensive environmental gradient of Mt. Kilimanjaro (Tanzania, East Africa) to study patterns and drivers of specialization, and robustness of plant–pollinator interactions against simulated species extinction with standardized sampling methods. We studied specialization, network robustness and other network indices of 67 quantitative plant–pollinator networks consisting of 268 observational hours and 4,380 plant–pollinator interactions along a 3.4 km elevational gradient. Using path analysis, we tested whether resource availability, pollinator richness, visitation rates, temperature, and/or area explain average specialization in pollinator communities. We further linked pollinator specialization to different pollinator taxa, and species traits, that is, proboscis length, body size, and species elevational ranges. Results We found that specialization decreased with increasing elevation at different levels of biological organization. Among all variables, mean annual temperature was the best predictor of average specialization in pollinator communities. Specialization differed between pollinator taxa, but was not related to pollinator traits. Network robustness against simulated species extinctions of both plants and pollinators was lowest in the most specialized interaction networks, that is, in the lowlands. Conclusions Our study uncovers patterns in plant–pollinator specialization along elevational gradients. Mean annual temperature was closely linked to pollinator specialization. Energetic constraints, caused by short activity timeframes in cold highlands, may force ectothermic species to broaden their dietary spectrum. Alternatively or in addition, accelerated evolutionary rates might facilitate the establishment of specialization under warm climates. Despite the mechanisms behind the patterns have yet to be fully resolved, our data suggest that temperature shifts in the course of climate change may destabilize pollination networks by affecting network architecture., We studied specialization patterns in plant–pollinator interactions along a 3.4 km elevational gradient on Mt. Kilimanjaro (Tanzania) and demonstrated that specialization decreased with elevation across networks, communities, and pollinator species. Temperature predicted the degree of specialization better than pollinator richness, resource availability, area, or pollinator traits. Despite the mechanisms behind the patterns have yet to be fully resolved, we conclude that temperature shifts in the course of climate change may destabilize pollination networks by affecting network architecture.

Despite decades of scientific effort, there is still no consensus on the determinants of broad-scale gradients of animal diversity. We argue that general drivers of diversity are unlikely to be found among the narrowly defined taxa which are typically analyzed in studies of broad-scale diversity gradients because ecological niches evolve largely conservatively. This causes constraints in the use of available niche space leading to systematic differences in diversity gradients among taxa. We instead advocate studies of phylogenetically diverse animal communities along broad environmental gradients. Such multi-taxa communities are less constrained in resource use and diversification and may be better targets for testing major classical hypotheses on diversity gradients. Besides increasing the spatial scale in analyses, expanding the phylogenetic coverage may be a second way to achieve higher levels of generality in studies of broad-scale diversity gradients.

1.Physiological and energetic mechanisms have been proposed to constrain body sizes of organisms along climatic gradients; however, these provide contrasting predictions. While Bergmann′s rule predicts increases of body sizes in cooler climates resulting from physiological constraints, energy-based community assembly rules suggest declines in the mean body size of species caused by increased extinction probabilities for large-bodied species in low-energy habitats. 2.We tested these contrasting hypotheses by quantifying trait distributions in bee communities along a 3.6 km elevational gradient at Mt. Kilimanjaro. Traditionally, intra- and interspecific trait shifts along environmental gradients have been investigated in isolation. However, a surge of theoretical approaches and studies on plants demonstrated that the explicit integration of trait variation among and within species can be essential for identifying the mechanisms that shape traits and related ecosystem functions along environmental gradients. We therefore studied variation of body size and related morphological traits at both the intra- and interspecific level. 3.We found support for both physiological constraints and energy-based community assembly rules as drivers of trait distribution in bee communities along elevational gradients, which, however, affected different levels of biotic organization, i.e. the population and community level. While the number of bee species with large body sizes declined with elevation, individuals within species became on average larger, resulting in contrasting trends in morphometric parameters at the community versus population level. 4.Furthermore, body size within bee communities became less variable at higher elevations, largely as a result of a non-random, directive loss of species, but paralleled by a decline in intraspecific variance, suggesting intensified filtering effects with increasing elevation. Similar patterns were found for other functional traits related to the foraging ecology of bees (tongue length, relative forewing length). 5.We conclude that along climatic gradients both physiological and energetic constraints shape trait distributions of pollinators, but at different levels of biological organization. This article is protected by copyright. All rights reserved.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown

Species’ functional traits set the blueprint for pair-wise interactions in ecological networks. Yet, it is unknown to what extent the functional diversity of plant and animal communities controls network assembly along environmental gradients in real-world ecosystems. Here we address this question with a unique dataset of mutualistic bird–fruit, bird–flower and insect–flower interaction networks and associated functional traits of 200 plant and 282 animal species sampled along broad climate and land-use gradients on Mt. Kilimanjaro. We show that plant functional diversity is mainly limited by precipitation, while animal functional diversity is primarily limited by temperature. Furthermore, shifts in plant and animal functional diversity along the elevational gradient control the niche breadth and partitioning of the respective other trophic level. These findings reveal that climatic constraints on the functional diversity of either plants or animals determine the relative importance of bottom-up and top-down control in plant–animal interaction networks., Differential responses of plant and animal functional diversity to climatic variation could affect trait matching in mutualistic interactions. Here, Albrecht et al. show that network structure varies across an elevational gradient owing to bottom-up and top-down effects of functional diversity.

Aim Understanding the mechanisms controlling variation in species richness along environmental gradients is one of the most important objectives in ecology. Resource availability is often considered as the major driver of animal diversity. However, in ectotherms, temperature might play a predominant role as it modulates metabolic rates and the access of animals to resources. Here, we investigate the relative importance of resource availability and temperature in determining the diversity pattern of bees along a 3.6-km elevational gradient. Location Mount Kilimanjaro, Tanzania. Methods We assessed bee species richness and abundance with pan traps and floral resources with transect records on 60 study sites which were equally distributed over six near-natural and six disturbed habitat types along an elevational gradient from 870 to 4550 m a.s.l. We used path analysis to disentangle the effects of temperature, precipitation, floral resource abundance, bee abundance and land use on bee species richness. In addition, we monitored flower visitation rates during transect walks at different elevations to evaluate the temperature dependence of bee–flower interactions. Results Bee species richness continuously declined with elevation in natural and disturbed habitats. While the abundance of floral resources had a significant but only weak effect on species richness, the effect of temperature was strong. Temperature had a strong positive effect on species richness that was not mediated by bee abundance and an indirect effect via bee abundances. We observed higher levels of bee–flower interactions at higher temperatures, supporting the hypothesis that temperature limits diversity by constraining resource exploitation in ectotherms. Main conclusions Temperature and the availability of resources shape species richness patterns along environmental gradients. In ectothermic organisms like bees temperature seems to have the more important role, as it both limits the access to resources (abundance-mediated effect) and accelerates other (abundance-independent) ecological and evolutionary processes that drive the maintenance and origination of diversity.

The factors determining gradients of biodiversity are a fundamental yet unresolved topic in ecology. While diversity gradients have been analysed for numerous single taxa, progress towards general explanatory models has been hampered by limitations in the phylogenetic coverage of past studies. By parallel sampling of 25 major plant and animal taxa along a 3.7 km elevational gradient on Mt. Kilimanjaro, we quantify cross-taxon consensus in diversity gradients and evaluate predictors of diversity from single taxa to a multi-taxa community level. While single taxa show complex distribution patterns and respond to different environmental factors, scaling up diversity to the community level leads to an unambiguous support for temperature as the main predictor of species richness in both plants and animals. Our findings illuminate the influence of taxonomic coverage for models of diversity gradients and point to the importance of temperature for diversification and species coexistence in plant and animal communities., Explaining species richness patterns is a key question in ecology. Peters et al. sample diverse plant and animal groups across elevation on Mt. Kilimanjaro to show that, while disparate factors drive distributions of individual taxa, diversity overall decreases with elevation, mostly driven by effects of temperature.

The two moth species Heliothis virescens (Hv) and H. subflexa (Hs) are closely related, but have vastly different feeding habits. Hv is a generalist and an important pest in many crops in the USA, while Hs is a specialist feeding only on plants in the genus Physalis. In this study, we conducted a comparative population genetic analysis to assess whether and how generalist and specialist life styles are reflected in differences in population structures. In Hv 98% of the total variation occurred within populations. The overall differentiation (F(ST)) between regions was 0.006 and even lower between years (0.0039) and hosts (0.0028). Analyses of population structure suggest that all individuals form one genetically homogeneous population, except for at most 12 individuals (6%) that diverged from this cluster. Population homogeneity likely results from the high mobility of Hv and its generalist feeding behaviour. Hs exhibited substantially more population structure. Even though 96% of the total variation was attributable to within-population variability, F(ST)-values between Hs populations were 10 times higher than between Hv populations. Hs populations showed significant isolation by distance. Analyses of Hs population structure suggest at least two subpopulations and thus some degree of metapopulation structure. We speculate that the patchy distribution of Physalis - the exclusive food source of Hs - contributes to differences in population structure between these closely related species. The finding that the specialist shows more population differentiation than the generalist corroborates the notion that host specialization is not an evolutionary dead end but a dynamic trait.

Variability within sex pheromone signalling systems is generally believed to be low because of strong stabilizing selection; yet the noctuid moth Heliothis subflexa (Hs) shows significant intraspecific variation. One possible explanation is that females may alter their sex pheromone blend depending on prevailing olfactory cues in the habitat, which we termed the ‘experience hypothesis’. This could be adaptive if Hs females experiencing the pheromone of another species, Heliothis virescens (Hv), responded to reduce the frequency of heterospecific matings. We exposed Hs females to no pheromone, Hs pheromone or Hv pheromone in the first 3 days of their adult lives. Hs females in the latter treatment produced significantly more of the acetate Z11-16:OAc, which inhibits the attraction of Hv males. To our knowledge, this is the first study showing adaptive phenotypic plasticity in a moth sex pheromone and suggests that behavioural differentiation may precede genetic divergence in the sexual signals of moths.

Wild animals substantially support crop production by providing ecosystem services, such as pollination and natural pest control. However, the strengths of synergies between ecosystem services and their dependencies on land-use management are largely unknown. Here, we took an experimental approach to test the impact of land-use intensification on both individual and combined pollination and pest control services in coffee production systems at Mount Kilimanjaro. We established a full-factorial pollinator and vertebrate exclosure experiment along a land-use gradient from traditional homegardens (agroforestry systems), shaded coffee plantations to sun coffee plantations (total sample size = 180 coffee bushes). The exclusion of vertebrates led to a reduction in fruit set of ca 9%. Pollinators did not affect fruit set, but significantly increased fruit weight of coffee by an average of 7.4%. We found no significant decline of these ecosystem services along the land-use gradient. Pest control and pollination service were thus complementary, contributing to coffee production by affecting the quantity and quality of a major tropical cash crop across different coffee production systems at Mount Kilimanjaro.

Females of the tobacco budworm, Heliothis virescens (F.), exhibit distinct geographical and temporal variation in sex pheromone composition, but the causes and significance of this variation are largely unexplored. Here we assessed whether 1) female pheromone variation was related to the host plants of origin, and 2) pheromone lures with varying amounts of Z9–14:Ald or 16:Ald were differentially attractive to males. Variation in female pheromone did not seem to be related to the host plants from which the eggs or larvae were collected, which may be because field-collected larvae were reared for three to five larval stages on artificial diet. By varying the concentration of Z9–14:Ald within the range in the female pheromone gland, we found males were more attracted as the amount increased from 1 to 10% relative to Z11–16:Ald, but significantly less with the highest concentration of 25%. In contrast, with 16:Ald, similar numbers of tobacco budworm males were caught in all traps where 16:Ald ranged from 0 to 200%. These results show that variation in Z9–14:Ald but not 16:Ald is evolutionary significant and likely subject to stabilizing selection in the field.

Evolutionary diversification of sexual communication systems in moths is perplexing because signal and response are under stabilizing selection in many species, and this is expected to constrain evolutionary change. In the mothHeliothis virescens, we consistently found high phenotypic variability in the female sex pheromone blend within each of four geographically distant populations. Here, we assess the heritability, genetic basis and behavioural consequences of this variation. Artificial selection with field-collected moths dramatically increased the relative amount of the saturated compound 16:Ald and decreased its unsaturated counterpart Z11–16:Ald, the major sex pheromone component (high line). In a cross between the high- and low-selected lines, one quantitative trait locus (QTL) explained 11–21% of the phenotypic variance in the 16:Ald/Z11–16:Ald ratio. Because changes in activity of desaturase enzymes could affect this ratio, we measured their expression levels in pheromone glands and mapped desaturase genes onto our linkage map. A delta-11-desaturase had lower expression in females producing less Z11–16:Ald; however, this gene mapped to a different chromosome than the QTL. A model in which the QTL is a trans-acting repressor of delta-11 desaturase expression explains many features of the data. Selection favouring heterozygotes which produce more unsaturated components could maintain a polymorphism at this locus.