Linking deforestation scenarios to pollination services and economic returns in coffee agroforestry systems.

The ecological and economic consequences of rain forest conversion and fragmentation for biodiversity, ecosystem functioning, and ecosystem services like protection of soils, water retention, pollination, or biocontrol are poorly understood. In human-dominated tropical landscapes, forest remnants may provide ecosystem services and act as a source for beneficial organisms immigrating into adjacent annual and perennial agro-ecosystems. In this study, we use empirical data on the negative effects of increasing forest distance on both pollinator diversity and fruit set of coffee to estimate future changes in pollination services for different land use scenarios in Sulawesi, Indonesia. Spatially explicit land use simulations demonstrate that depending on the magnitude and location of ongoing forest conversion, pollination services are expected to decline continuously and thus directly reduce coffee yields by up to 18%, and net revenues per hectare up to 14% within the next two decades (compared to average yields of the year 2001). Currently, forests in the study area annually provide pollination services worth 46 Euros per hectare. However, our simulations also revealed a potential win-win constellation, in which ecological and economic values can be preserved, if patches of forests (or other natural vegetation) are maintained in the agricultural landscape, which could be a viable near future option for local farmers and regional land use planners.     

Modeling scale up of anthropogenic impacts from individual pollinator behavior to pollination systems

Understanding how anthropogenic disturbances affect plant–pollinator systems has important implications for the conservation of biodiversity and ecosystem functioning. Previous laboratory studies show that pesticides and pathogens, which have been implicated in the rapid global decline of pollinators over recent years, can impair behavioral processes needed for pollinators to adaptively exploit floral resources and effectively transfer pollen among plants. However, the potential for these sublethal stressor effects on pollinator–plant interactions at the individual level to scale up into changes to the dynamics of wild plant and pollinator populations at the system level remains unclear. We developed an empirically parameterized agent-based model of a bumblebee pollination system called SimBee to test for effects of stressor-induced decreases in the memory capacity and information processing speed of individual foragers on bee abundance (scenario 1), plant diversity (scenario 2), and bee–plant system stability (scenario 3) over 20 virtual seasons. Modeling of a simple pollination network of a bumblebee and four co-flowering bee-pollinated plant species indicated that bee decline and plant species extinction events could occur when only 25% of the forager population showed cognitive impairment. Higher percentages of impairment caused 50% bee loss in just five virtual seasons and system-wide extinction events in less than 20 virtual seasons under some conditions. Plant species extinctions occurred regardless of bee population size, indicating that stressor-induced changes to pollinator behavior alone could drive species loss from plant communities. These findings indicate that sublethal stressor effects on pollinator behavioral mechanisms, although seemingly insignificant at the level of individuals, have the cumulative potential in principle to degrade plant–pollinator species interactions at the system level. Our work highlights the importance of an agent-based modeling approach for the identification and mitigation of anthropogenic impacts on plant–pollinator systems.     

Higher soybean production using honeybee and wild pollinators, a sustainable alternative to pesticides and autopollination

This is the first report showing that using honeybee (Apis mellifera) and wild pollinators complementary pollination can enhance soybean productivity (Glycine max). Current industrial production of soybean involves autopollination and high loads of pesticides. Therefore, growers have neglected possible biotic pollination despite suggestions that soybean benefit from insect pollinators. Reports advocating possible biotic pollination are based on experiments where bees are caged with flowering plants and the absence of pesticides, thus not in field conditions. Therefore, here we compared in field conditions soybean yield produced (1) independently of biotic pollinators, (2) with wild pollinators and (3) with honeybee colonies. Results showed an increase of +6.34 % of soybean yield in areas where wild pollinators had free access to flowers. The introduction of honeybee colonies further raised the yield of +18.09 %. Our findings therefore show that, though soybean is autogamous, allowing pollination by wild pollinators leads to higher yields. Moreover, adding honeybee mitigates pollination deficits and improves yield compared to current practices.     

Effects of pollinator density-dependent preferences on field margin visitations in the midst of agricultural monocultures: A modelling approach

Managed field margins offer a means of reducing the impact of agricultural monocultures within intensively managed environments. By providing refuge for wild plants and the pollinators associated with them, field margins can also contribute to enhancing the pollination services within the monoculture. However, the effects of the monoculture on pollinator behaviour need to be carefully considered. It is known that pollinators may show density-dependent preferences such as neophobia (an avoidance of unfamiliar items) when different types of flower are available within their environment, and the dominance of monoculture crops within the environment may consequently have adverse effects upon the preferences shown by pollinators living in the field margins within them. In order to examine how pollinator preferences for wild flowers are affected by monocultures, we modelled the effects of density-dependent preferences, flower densities, and the geometry of field margins within a monoculture landscape using numerical simulations. This was done by considering how the placement of pollinator nests within a simple, spatially explicit landscape consisting of fields of monoculture crops separated by margins containing wild flowers affected the ratio of wild and monoculture crops experienced by the pollinator, given that it could only forage within a limited distance from its nest. Increasing field margin width and decreasing monoculture field width both led to an increase in pollinators visiting wild flowers (which levelled off as width increased). The size of the monoculture fields had little additional effect once they had passed an intermediate width. Increasing wild flower density within the margins led to a shift away from preferring monocultures. When wild flowers were at low densities compared to the monoculture, even the addition of small amounts of extra wild flowers had a large effect in shifting foraging preferences away from the monoculture. The distance which pollinators normally forage over only has an effect upon preferences for wild flowers when the travel distance is small. This suggests that careful consideration of margin design might be extremely important for those species which do not travel far. Innate preferences for density-dependence and particular crop types may also have an effect on preference behaviour. We demonstrate that the way in which resources are presented to indigenous pollinators may be extremely important in influencing where they choose to forage within agricultural landscapes. Careful margin design, as well as increasing the density of wild flowers (such as by enhancing the wild seed bank within the margins), may lead to overall improvements in ecosystem function within intensively farmed monocultures.     

Advances in pollination ecology from tropical plantation crops

Although ecologists traditionally focus on natural ecosystems, there is growing awareness that mixed landscapes of managed and unmanaged systems provide a research environment for understanding basic ecological relationships on a large scale. Here, we show how tropical agroforestry systems can be used to develop ideas about the mechanisms that influence species diversity and subsequent biotic interactions at different spatial scales. Our focus is on tropical plantation crops, mainly coffee and cacao, and their pollinators, which are of basic ecological interest as partners in an important mutualistic interaction. We review how insect-mediated pollination services depend on local agroforest and natural habitats in surrounding landscapes. Further, we evaluate the functional significance of pollinator diversity and the explanatory value of species traits, and we provide an intercontinental comparison of pollinator assemblages. We found that optimal pollination success might be best understood as a consequence of niche complementarities among pollinators in landscapes harboring various species. We further show that small cavity-nesting bees and small generalist beetles were especially affected by isolation from forest and that larger-bodied insects in the same landscapes were not similarly affected. We suggest that mixed tropical landscapes with agroforestry systems have great potential for future research on the interactions between plants and pollinators.     

Rapid assessment of insect pollination services to inform decision-making

Pollinator declines have prompted efforts to assess how land-use change affects insect pollinators and pollination services in agricultural landscapes. Yet many tools to measure insect pollination services require substantial landscape-scale data and technical expertise. In expert workshops, 3 straightforward methods (desk-based method, field survey, and empirical manipulation with exclusion experiments) for rapid insect pollination assessment at site scale were developed to provide an adaptable framework that is accessible to nonspecialist with limited resources. These methods were designed for TESSA (Toolkit for Ecosystem Service Site-Based Assessment) and allow comparative assessment of pollination services at a site of conservation interest and in its most plausible alternative state (e.g., converted to agricultural land). We applied the methods at a nature reserve in the United Kingdom to estimate the value of insect pollination services provided by the reserve. The economic value of pollination services provided by the reserve ranged from US$6163 to US$11,546/year. The conversion of the reserve to arable land would provide no insect pollination services and a net annual benefit from insect-pollinated crop production of approximately $1542/year (US$24∙ha^–1∙year^–1). The methods had wide applicability and were readily adapted to different insect-pollinated crops: rape (Brassica napus) and beans (Vicia faba) crops. All methods were rapidly employed under a low budget. The relatively less robust methods that required fewer resources yielded higher estimates of annual insect pollination benefit.     

Effects of human demand on conservation planning for biodiversity and ecosystem services

Safeguarding ecosystem services and biodiversity is critical to achieving sustainable development. To date, ecosystem services quantification has focused on the biophysical supply of services with less emphasis on human beneficiaries (i.e., demand). Only when both occur do ecosystems benefit people, but demand may shift ecosystem service priorities toward human-dominated landscapes that support less biodiversity. We quantified how accounting for demand affects the efficiency of conservation in capturing both human benefits and biodiversity by comparing conservation priorities identified with and without accounting for demand. We mapped supply and benefit for 3 ecosystem services (flood mitigation, crop pollination, and nature-based recreation) by adapting existing ecosystem service models to include and exclude factors representing human demand. We then identified conservation priorities for each with the conservation planning program Marxan. Particularly for flood mitigation and crop pollination, supply served as a poor proxy for benefit because demand changed the spatial distribution of ecosystem service provision. Including demand when jointly targeting biodiversity and ecosystem service increased the efficiency of conservation efforts targeting ecosystem services without reducing biodiversity outcomes. Our results highlight the importance of incorporating demand when quantifying ecosystem services for conservation planning.     

Separating selection by diurnal and nocturnal pollinators on floral display and spur length in Gymnadenia conopsea

Most plants attract multiple flower visitors that may vary widely in their effectiveness as pollinators. Floral evolution is expected to reflect interactions with the most important pollinators, but few studies have quantified the contribution of different pollinators to current selection on floral traits. To compare selection mediated by diurnal and nocturnal pollinators on floral display and spur length in the rewarding orchid Gymnadenia conopsea, we manipulated the environment by conducting supplemental hand-pollinations and selective pollinator exclusions in two populations in central Norway. In both populations, the exclusion of diurnal pollinators significantly reduced seed production compared to open pollination, whereas the exclusion of nocturnal pollinators did not. There was significant selection on traits expected to influence pollinator attraction and pollination efficiency in both the diurnal and nocturnal pollination treatment. The relative strength of selection among plants exposed to diurnal and nocturnal visitors varied among traits and populations, but the direction of selection was consistent. The results suggest that diurnal pollinators are more important than nocturnal pollinators for seed production in the study populations, but that both categories contribute to selection on floral morphology. The study illustrates how experimental manipulations can link specific categories of pollinators to observed selection on floral traits, and thus improve our understanding of how species interactions shape patterns of selection.     

Evaluating the Potential for Conservation Development: Biophysical, Economic, and Institutional Perspectives

Abstract:  The widespread conversion of rural land to low-density residential development poses an immediate threat to biodiversity and to the provision of ecosystem services. Given that development will continue and environmental stakes are high, analyzing alternative growth strategies is critical. Conservation development is one such strategy that has the potential to benefit ecosystems and diverse stakeholders including developers, homebuyers, governments, and society as a whole. Conservation development clusters homes on one part of a property to manage the most ecologically important land for the conservation of biodiversity and ecosystem services. We draw on lessons learned from landscape ecology, open-space development, and regional planning to weigh the biophysical, economic, and institutional evidence for and against conservation development. Conservation development offers many potential environmental and economic advantages: relatively high home values and appreciation rates, lower development costs, and social and ecological benefits to society including landscape connectivity, protection and active stewardship of important ecological assets, and the maintenance of ecosystem services. But this approach also has shortcomings: it may require enlightened institutional regulations and regional planning (and/or ecologically aware developers), it is not always more profitable than conventional development and thus may require subsidies or incentives, and additional research is required to fully understand its benefits and drawbacks. With more information on the effects of clustering, the development of flexible zoning laws, and effective regional planning, conservation development could be a viable strategy for sustaining biodiversity and ecosystem services in changing landscapes.     

Advertising to the enemy: enhanced floral fragrance increases beetle attraction and reduces plant reproduction

Many organisms face challenges in avoiding predation while searching for mates. For plants, emitting floral fragrances to advertise reproductive structures could increase the attraction of detrimental insects along with pollinators. Very few studies have experimentally evaluated the costs and benefits of fragrance emission with explicit consideration of how plant fitness is affected by both pollinators and florivores. To determine the reproductive consequences of increasing the apparency of reproductive parts, we manipulated fragrance, pollination, and florivores in the wild Texas gourd, Cucurbita pepo var. texana. With enhanced fragrance we found an increase in the attraction of florivores, rather than pollinators, and a decrease in seed production. This study is the first to demonstrate that enhanced floral fragrance can increase the attraction of detrimental florivores and decrease plant reproduction, suggesting that florivory as well as pollination has shaped the evolution of floral scent.     

Floral scents: their roles in nursery pollination mutualisms

Mutualisms are interspecies interactions in which each participant gains net benefits from interacting with its partner. In nursery pollination mutualisms, pollinators reproduce within the inflorescence they pollinate. In these systems, each partner depends directly on the other for its reproduction. Therefore, the signal responsible for partner encounter is crucial in these horizontally transmitted mutualisms, in which the association between specific partners must be renewed at each generation. As in many other interspecies interactions, chemical signals are suspected to be important in the functioning of these mutualisms. We synthesized and compared the published data available on the role of floral scents in the functioning of the 16 known independently evolved nursery pollination mutualisms. So far, attraction of pollinators to their specific hosts has been investigated in only seven of these systems, and the majority of the studies have been conducted on one of them, fig/fig wasp interactions. While such unevenness of the information limits the potential for meta-analysis, some patterns emerge from this review concerning the role of flower volatiles in maintaining the specificity of pollinator attraction, in signaling the appropriate phenological stage for pollinator visit, in attracting the pollinator toward the rewardless sex in dioecious plant species and in aiding the location and exploitation of resources by parasites and predators associated with these mutualisms. Finally, we highlight new perspectives on the evolution of signals in these diversified systems depending on the age and the degree of specificity of the interaction, and on the effect of phylogenetic inertia on the evolutionary dynamics of plant signals.     

Planting gardens to support insect pollinators

Global insect pollinator declines have prompted habitat restoration efforts, including pollinator-friendly gardening. Gardens can provide nectar and pollen for adult insects and offer reproductive resources, such as nesting sites and caterpillar host plants. We conducted a review and meta-analysis to examine how decisions made by gardeners on plant selection and garden maintenance influence pollinator survival, abundance, and diversity. We also considered characteristics of surrounding landscapes and the impacts of pollinator natural enemies. Our results indicated that pollinators responded positively to high plant species diversity, woody vegetation, garden size, and sun exposure and negatively to the separation of garden habitats from natural sites. Within-garden features more strongly influenced pollinators than surrounding landscape factors. Growing interest in gardening for pollinators highlights the need to better understand how gardens contribute to pollinator conservation and how some garden characteristics can enhance the attractiveness and usefulness of gardens to pollinators. Further studies examining pollinator reproduction, resource acquisition, and natural enemies in gardens and comparing gardens with other restoration efforts and to natural habitats are needed to increase the value of human-made habitats for pollinators.     

Payments for Ecosystem Services as a Framework for Community-Based Conservation in Northern Tanzania

Abstract:  Payments for ecosystem services (PES) are an increasingly promoted approach to conservation. These approaches seek to develop financial mechanisms that create economic incentives for the maintenance of ecosystems and associated biodiversity by rewarding those who are responsible for provision of ecological services. There are, however, few cases in which such schemes have been used as a strategy for conserving wildlife in developing countries and very few operational examples of such schemes of any sort in sub-Saharan Africa. In savannah ecosystems, large mammal populations generally depend on seasonal use of extensive areas and are widely declining as a result of habitat loss, overexploitation, and policies that limit local benefits from wildlife. Community-based conservation strategies seek to create local incentives for conserving wildlife, but often have limited impact as a result of persistent institutional barriers that limit local rights and economic benefits. In northern Tanzania, a consortium of tourism operators is attempting to address these challenges through an agreement with a village that possesses part of a key wildlife dispersal area outside Tarangire National Park. The operators pay the community to enforce voluntary restrictions on agricultural cultivation and permanent settlement in a defined area of land. The initiative represents a potentially cost-effective framework for community-based conservation in an ecologically important area and is helping to reconcile historically conflicting local and national interests relative to land tenure, pastoralist livelihoods, and conservation. Wider adaptation of payments for ecosystem services approaches to settings where sustaining wildlife populations depends on local stewardship may help address current challenges facing conservation outside state-protected areas in savannah ecosystems in sub-Saharan Africa and other parts of the world.     

Collapse of a pollination web in small conservation areas

A suspected global decline in pollinators has heightened interest in their ecological significance. In a worst-case scenario, the decline of generalist pollinators is predicted to trigger cascades of linked declines among the multiple specialist plant species to which they are linked, but this has not been documented. I studied a portion of a pollination web involving a generalist pollinator, the oil-collecting bee Rediviva peringueyi, and a community of oil-secreting plants. Across 27 established conservation areas located in the Cape Floral Region, I found substantial variation in the bees' occurrence in relation to soil type and the successional stage of the vegetation. Anthropogenic declines were detectable against this background of naturally occurring variation: R. peringueyi was absent from small conservation areas (< 385 ha) in an urban matrix. In the absence of the bee, seed set failed in six specialist plant species that are pollinated only by R. peringueyi but remained high in a pollination generalist, which had replacement pollinators. The findings are consistent with theoretical predictions of the importance of generalist pollinators in maintaining the structure of pollination webs.     

Pollinator effectiveness varies with experimental shifts in flowering time

The earlier flowering times exhibited by many plant species are a conspicuous sign of climate change. Altered phenologies have caused concern that species could suffer population declines if they flower at times when effective pollinators are unavailable. For two perennial wildflowers, Tradescantia ohiensis and Asclepias incarnata, we used an experimental approach to explore how changing phenology affects the taxonomic composition of the pollinator assemblage and the effectiveness of individual pollinator taxa. After finding in the previous year that fruit set varied with flowering time, we manipulated flowering onset in greenhouses, placed plants in the field over the span of five weeks, and measured pollinator effectiveness as the number of seeds produced after a single visit to a flower. The average effectiveness of pollinators and the expected rates of pollination success were lower for plants of both species flowering earlier than for plants flowering at historical times, suggesting there could be reproductive costs to earlier flowering. Whereas for A. incarnata, differences in average seed set among weeks were due primarily to changes in the composition of the pollinator assemblage, the differences for T. ohiensis were driven by the combined effects of compositional changes and increases over time in the effectiveness of some pollinator taxa. Both species face the possibility of temporal mismatch between the availability of the most effective pollinators and the onset of flowering, and changes in the effectiveness of individual pollinator taxa through time may add an unexpected element to the reproductive consequences of such mismatches.     

Nectar secondary compounds affect self-pollen transfer: implications for female and male reproduction

Pollen movement within and among plants affects inbreeding, plant fitness, and the spatial scale of genetic differentiation. Although a number of studies have assessed how plant and floral traits influence pollen movement via changes in pollinator behavior, few have explored how nectar chemical composition affects pollen transfer. As many as 55% of plants produce secondary compounds in their nectar, which is surprising given that nectar is typically thought to attract pollinators. We tested the hypothesis that nectar with secondary compounds may benefit plants by encouraging pollinators to leave plants after visiting only a few flowers, thus reducing self-pollen transfer. We used Gelsemium sempervirens, a plant whose nectar contains the alkaloid gelsemine, which has been shown to be a deterrent to foraging bee pollinators. We found that high nectar alkaloids reduced the total and proportion of self-pollen received by one-half and one-third, respectively. However, nectar alkaloids did not affect female reproduction when we removed the potential for self-pollination (by emasculating all flowers on plants). We then tested the assumption that self-pollen in combination with outcrossed pollen depresses seed set. We found that plants were weakly self-compatible, but self-pollen with outcrossed pollen did not reduce seed set relative to solely outcrossed flowers. Finally, an exponential model of pollen carryover suggests that high nectar alkaloids could benefit plants via increased pollen export (an estimate of male function), but only when pollinators were efficient and abundant and plants had large floral displays. Results suggest that high nectar alkaloids may benefit plants via increased pollen export under a restricted set of ecological conditions, but in general, the costs of high nectar alkaloids in reducing pollination balanced or outweighed the benefits of reducing self-pollen transfer for estimates of female and male reproduction.     

Do rewardless orchids show a positive relationship between phenotypic diversity and reproductive success?

Among rewardless orchids, pollinator sampling behavior has been suggested to drive a positive relationship between population phenotypic variability and absolute reproductive success, and hence population fitness. We tested this hypothesis by constructing experimental arrays using the rewardless orchid Dactylorhiza sambucina, which is dimorphic for corolla color. We found no evidence that polymorphic arrays had higher mean reproductive success than monomorphic arrays for pollinia removal, pollen deposition, or fruit set. For pollinia removal, monomorphic yellow arrays had significantly greater reproductive success, and monomorphic red the least. A tendency for yellow arrays to have higher pollen deposition was also found. We argue that differential population fitness was most likely to reflect differential numbers of pollinators attracted to arrays, through preferential long-distance attraction to arrays with yellow inflorescences. Correlative studies of absolute reproductive success in 52 populations of D. sambucina supported our experimental results. To our knowledge this is the first study to suggest that attraction of a greater number of pollinators to rewardless orchids may be of greater functional importance to population fitness, and thus ecology and conservation, than are the behavioral sequences of individual pollinators.     

Detecting Insect Pollinator Declines on Regional and Global Scales

Recently there has been considerable concern about declines in bee communities in agricultural and natural habitats. The value of pollination to agriculture, provided primarily by bees, is >$200 billion/year worldwide, and in natural ecosystems it is thought to be even greater. However, no monitoring program exists to accurately detect declines in abundance of insect pollinators; thus, it is difficult to quantify the status of bee communities or estimate the extent of declines. We used data from 11 multiyear studies of bee communities to devise a program to monitor pollinators at regional, national, or international scales. In these studies, 7 different methods for sampling bees were used and bees were sampled on 3 different continents. We estimated that a monitoring program with 200–250 sampling locations each sampled twice over 5 years would provide sufficient power to detect small (2–5%) annual declines in the number of species and in total abundance and would cost U.S.$2,000,000. To detect declines as small as 1% annually over the same period would require >300 sampling locations. Given the role of pollinators in food security and ecosystem function, we recommend establishment of integrated regional and international monitoring programs to detect changes in pollinator communities. Detección de Declinaciones de Insectos Polinizadores a Escalas Regional y Global     

Producer-level benefits of sustainability certification

Initiatives certifying that producers of goods and services adhere to defined environmental and social-welfare production standards are increasingly popular. According to proponents, these initiatives create financial incentives for producers to improve their environmental, social, and economic performance. We reviewed the evidence on whether these initiatives have such benefits. We identified peer-reviewed, ex post, producer-level studies in economic sectors in which certification is particularly prevalent (bananas, coffee, fish products, forest products, and tourism operations), classified these studies on the basis of whether their design and methods likely generated credible results, summarized findings from the studies with credible results, and considered how these findings might guide future research. We found 46 relevant studies, most of which focused on coffee and forest products and examined fair-trade and Forest Stewardship Council certification. The methods used in 11 studies likely generated credible results. Of these 11 studies, nine examined the economic effects and two the environmental effects of certification. The results of four of the 11 studies, all of which examined economic effects, showed that certification has producer-level benefits. Hence, the evidence to support the hypothesis that certification benefits the environment or producers is limited. More evidence could be generated by incorporating rigorous, independent evaluation into the design and implementation of projects promoting certification.     

Predicting the economic impact of an invasive species on an ecosystem service.

Quantifying the impact of alien invasive species on ecosystem services is an essential step in developing effective practices and policy for invasive species management. Here we develop a stochastic bioeconomic model that enables the economic impact of an invasive pest to be estimated before its arrival, based on relatively poorly specified ecological and economic parameters. We developed the model by using a hypothetical invasion of the varroa bee mite (Varroa destructor) into Australia and the negative flow-on effects that it would have on pollination by reducing honey bee populations, giving rise to a loss of pollination services, reduced crop yields, and additional production costs. If the mite were to continue to be prevented from entering the country over the next 30 years, we estimate that the economic costs avoided would be U.S. $16.4-38.8 million (Aus $21.3-50.5 million) per year. We suggest that current invasion response funding arrangements in Australia, which do not acknowledge these avoided damages, require amendment.