Article

Verges Along Forest Roads Promote Wild Bees

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Abstract

Forests in Germany are occupied with roads, paths, and trails with a density of 5.03 km/km². Their construction and maintenance create a network of verges promoting flowering plants. Whether these verges are visited by bees, which factors are determining their abundance, diversity, and composition, and which flowering resources are used is unknown. We selected 13 verges in the Black Forest (Germany), sweep-netted wild bees along transects, calculated the flowering area of all herbs, and measured the area (hectares) of grassland within 1 km around the transects. To evaluate the resource use of a common bumblebee species, we analyzed the pollen load of common carder bees (Bombus pascuorum) using microscopes. The abundance and diversity of wild bees was positively related to flowering area. With an increasing area of grassland, the abundance of ubiquitous species increased. Wild bee community composition was driven by flowering area. Common carder bees collected pollen from several flower resources but mainly used few species, such as the common hemp nettle (Galeopsis tetrahit L.). As the flowering area influenced wild bee abundance, diversity, and composition, we suggest creating road verges that favor the occurrence of native flowering plants to support wild bees in forest ecosystems. Study Implications: Forest road verges generally have higher light availability than the forest interior and therefore have higher availability of flowering plants. Although the importance of verges for wild bee conservation in agricultural landscapes is known, forest road verges are understudied. Our study demonstrates that forest road verges are important habitats for many ubiquitous bees and that the flowering area on these verges is the key determinant for the abundance and diversity of wild bees. Therefore, creating road verges that favor the occurrence of native flowering plants is key to support bees on these verges.

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1. Wild bees importantly pollinate both crop and wild plants. Yet, in intensive agricultural landscapes, wild bees are rare due to resource limitations of nectar and pollen. Flower strips and hedges are often used as resource enhancements for wild bees to overcome this shortage, but provide floral resources only during specific time periods. To sustain diverse and stable bee communities, bee-attractive flowers need to be available during the entire growing season. This may be achieved by combining flower strips and hedges to complement each other and provide continuous floral resources. 2. Over three subsequent years, we compared the phenology of flower and wild bee communities in perennial flower strips, hedges and improved hedges (complemented with a sown herb layer) in conventional apple orchards in Southern Germany, a pollination-dependent crop system. 3. Hedges provided floral resources in the early season while the flower strips took over later in the season. 4. Bees visited the hedges mostly from March to June, whereas they visited the flower strips from June to August in the first year, and in the second year already from April onwards. Flower strips were visited with an overall higher abundance and species richness than both the hedges and the improved hedges. 5. Synthesis and application. For enhancing wild bees in intensive apple orchards, hedges and perennial flower strips are complementary in providing flower resources. Yet, flower strips bloom more constantly and during periods of flower scarcity, and thus attract a higher diversity of bees than hedges. Perennial flower strips of different age classes should be preferred over annual strips, at best in a network with some well-maintained hedges, as perennial flower strips of different age attract different bee communities and thus potentially a higher bee diversity on the landscape level.
Article
Flower strips are a frequently adopted measure to conserve insects, especially pollinators, and are subsidized as Agri-Environmental Scheme in many regions. They provide a high quantity of flowers, but their flower species composition and phenological development is mostly uniform. This may result in only a fraction of pollinator species being enhanced. Flower-rich semi-natural habitat patches along slopes, fences or ditches, may provide resources for additional species, but they are not politically promoted. In this study, we compare pollinator communities in sown perennial flower strips with existing flower-rich herbaceous semi-natural habitat patches, both located at the edge of conventional apple orchards in Southern Germany. The flower strips attracted a higher pollinator abundance and species richness than the existing habitat patches. However, the bee species composition differed between the two habitat types. The existing habitat patches attracted bee species with different pollen specialization than the sown flower strips. Pollinator abundance and species richness varied between the different existing habitat patches indicating a high heterogeneity of these existing habitats, whereas the flower strips showed consistently high pollinator abundance and richness. We conclude that existing herbaceous habitat patches are attractive for pollinators and should be promoted by policy actions. Flower-rich semi-natural habitat patches develop without sowing next to typical agricultural landscape elements like slopes, fences and ditches with moderate mowing or herbicide application. They generally do not compete with agricultural land use and thus have a high potential to promote pollinator conservation.
Article
Forest-associated species, which depend on forest habitat for their survival, are among the world's most vulnerable species due to widespread forest loss. However, in many parts of the world, forests are re-growing. Thus, if forest-associated species can persist in young forests their conservation outlook is less bleak. We examined the effects of forest loss and regrowth on bee pollinators in eastern North America using three datasets totaling 36,605 individual specimens. First, we conducted a regional-scale study to identify forest-associated and habitat generalist bee species. Second, we examined how the abundance and richness of each group change with forest area and age, by collecting bees from forests chosen to vary orthogonally in these variables. Lastly, we examined whether the results of our field studies were consistent with long-term, regional trends in the richness of both groups, using a dataset of museum specimens collected between 1872 and 2011, which was a period of reforestation in our study region. We found that the abundance and richness of forest-associated bees increase with forest area, while being relatively insensitive to forest age. By contrast, habitat generalist bees are unaffected by forest area, but decrease in abundance with forest age. Consistent with these results, we found a 16% increase in the richness of forest-associated bees over a 140-year time series as reforestation occurred in eastern North America, but no significant trend for habitat generalists. Overall, our results show that the loss of forest area from a landscape harms forest-associated bees, and that young forests have high conservation value for both forest-associated and habitat generalist bees.
Article
Native pollinator populations across the United States are increasingly threatened by a multitude of ecological stressors. Although the drivers behind pollinator population declines are varied, habitat loss/degradation remains one of the most important threats. Forested landscapes, where the impacts of habitat loss/degradation are minimized, are known to support robust pollinator populations in eastern North America. Within heavily forested landscapes, timber management is already implemented as a means for improving forest health and enhancing wildlife habitat, however, little is known regarding the characteristics within regenerating timber harvests that affect forest pollinator populations. In 2018 and 2019, we monitored insect pollinators in 143 regenerating (≤9 growing seasons post-harvest) timber harvest sites across Pennsylvania. During 1129 survey events, we observed over 9100 bees and butterflies, 220 blooming plant taxa, and collected over 2200 pollinator specimens. Bee and butterfly abundance were positively associated with season-wide floral abundance and negatively associated with dense vegetation that inhibits the growth of understory floral resources. Particularly in late summer, few pollinators were observed in stands >6 years post-harvest, with models predicting five times more bees in 1-year-old harvests than in 9-year-old harvests. Pollinator species diversity was positively associated with floral diversity and percent forb cover, and negatively associated with percent tall (>1 m) sapling cover. These results suggest that regenerating timber harvests promote abundant and diverse pollinator communities in the Appalachian Mountains, though pollinator abundance declined quickly as woody stems regenerated. Ultimately, our findings contribute to a growing body of literature suggesting that dynamic forest management producing a mix of age classes would benefit forest pollinator populations in the Central Appalachian Mountains.
Article
Populations of bumble bees and other pollinators have declined over the past several decades due to numerous threats, including habitat loss and degradation. However, we can rarely investigate the role of resource loss due to a lack of detailed long‐term records of forage plants and habitats. We use 22‐year repeated surveys of more than 262 sites located in grassland, forest, and wetland habitats across Illinois, USA to explore how the abundance and richness of bumble bee food plants have changed over the period of decline of the endangered rusty patched bumble bee (Bombus affinis). We document a decline in abundance of bumble bee forage plants in forest understories, which our phenology analysis suggests provide the primary nectar and pollen sources for foundress queens in spring, a critical life stage in bumble bee demography. By contrast, the per‐unit area abundance of food plants in primarily midsummer‐flowering grassland and wetland habitats has not declined. However, the total area of grasslands has declined across the region resulting in a net loss of grassland resources. Synthesis and applications. Our results suggest a decline in spring‐flowering forest understory plants is a previously unappreciated bumble bee stressor, compounding factors like agricultural intensification, novel pathogen exposure, and grassland habitat loss. These findings emphasize the need for greater consideration of habitat complementarity in bumble bee conservation. We conclude that continued loss of early‐season floral resources may add additional stress to critical life stages of bumble bees and limit restoration efforts if not explicitly considered in pollinator conservation.
Article
Concerns about long-term pollinator declines have made assessing bee communities a priority in nonagricultural ecosystems, including managed forests. We assessed wild bee communities in Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) stands one to 15 years after clearcut harvest in western Oregon, USA, testing the hypothesis that bee diversity would be high initially and then decline with time-dependent reductions in floral resources. We captured 2,009 individual bees that represented 67 distinct species/morphospecies in 20 genera and five families. Asymptotic estimators of bee diversity representing Shannon and Simpson diversity were greater in communities during the second half of the early seral period, indicating older early seral stands were less diverse and contained more common and dominant bee species. In addition, observed species richness and bee abundance peaked at approximately three years postharvest and declined thereafter by 20% and 30% per year, respectively. Because floral resources declined in concert with reductions in bee diversity as stands aged, food appears to be a key driver of forest bee communities. Our results indicate that postharvest Douglas-fir stands supported a diversity of bees, including important crop pollinators, but their value to bees was restricted to a relatively short window at the beginning of the early seral period.
Article
Global biodiversity declines are attributed to many factors, including landscape fragmentation and vegetation homogenization. These patterns may be exacerbated by the intensification of management in agroecosystems, as management to meet the increasing demand for food, fuel, and fiber often comes at the cost of biodiversity and subsequent ecosystem functions and services. Conserving biodiversity will be necessary to create sustainable agroecosystems capable of optimizing both production and services such as pollination. We conducted a meta-analysis with 109 studies to examine the relationship between plant species richness and pollinator species richness to determine whether higher plant species richness supports higher pollinator species richness, especially in areas prone to biodiversity losses. We found most groups of insect pollinators, including bees, butterflies, flies, moths, and wasps, responded positively to increasing plant species richness, irrespective of location or land use, suggesting the capacity to increase pollinator richness through management strategies that increase plant species richness. However, we found pollinators in manipulated studies did not consistently respond to increasing plant species richness despite the overall positive relationships in observational and experimental studies, highlighting the importance of plant selection when making management decisions aiming to improve pollinator richness. Additional studies in regions such as Africa and South America will help fill in latitudinal gradients and provide greater coverage necessary to refine patterns. Increasing plant species richness through management changes or restorations will likely increase pollinator richness and be beneficial in agroecosystems to support biodiversity.
Article
1. Understanding the roles of habitat fragmentation and resource availability in shaping animal movement are integral for promoting species persistence and conservation. For insects such as bumble bees, their movement patterns affect the survival and reproductive potential of their colonies, as well as the pollen flow of plant species. However, the understanding of their mobility or the impact of putative barriers in natural environments is limited due to the technical difficulties of studying wild populations. 2. Genetic mark–recapture was used to estimate the foraging distance, resource use, and site connectivity of two bumble bee species in a montane meadow complex composed of open meadows within a matrix of forest. 3. There was no evidence that forests or changes in landcover function as barriers to the fine‐scale movement for either species. Substantially greater colony‐specific foraging distances were found for Bombus vosnesenskii (maximum: 1867 m) compared to Bombus bifarius (maximum: 362 m). Despite this difference in absolute range, both species were detected across putative forest barriers at frequencies expected by uninhibited movement. Siblings separated by greater distances were more likely to be foraging on different floral species, potentially suggesting a resource‐based motivation for movement. 4. These results suggest that bumble bee foraging patterns are influenced by species‐specific differences in movement capacity, with little influence of matrix composition between resource patches. They also support the perspective that habitat conservation for bumble bees should prioritise providing abundant and diverse patches of resources within species‐specific movement radii with less emphasis on matrix composition.
Article
Bumble bees ( Bombus) are unusually important pollinators, with approximately 260 wild species native to all biogeographic regions except Africa, Australia, and New Zealand. As they are vitally important in natural ecosystems and to agricultural food production globally, the increase in reports of declining distribution and abundance over the past decade has led to an explosion of interest in bumble bee population decline. We summarize data on the threat status of wild bumble bee species across biogeographic regions, underscoring regions lacking assessment data. Focusing on data-rich studies, we also synthesize recent research on potential causes of population declines. There is evidence that habitat loss, changing climate, pathogen transmission, invasion of nonnative species, and pesticides, operating individually and in combination, negatively impact bumble bee health, and that effects may depend on species and locality. We distinguish between correlational and causal results, underscoring the importance of expanding experimental research beyond the study of two commercially available species to identify causal factors affecting the diversity of wild species. Expected final online publication date for the Annual Review of Entomology, Volume 65 is January 7, 2020. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
Article
Increasing the proportion of unmanaged forests in multi-functional forest landscapes is a primary goal of international and national conservation strategies aiming at restoring natural properties in structurally simplified forests. However, the development of structural features and associated habitat suitability for forest species is largely unknown and even controversially discussed, as the development of newly established reserves is uni-directional and passes through dense maturation stages. This may negatively affect open forest species in the first phase after reserve designation. We evaluated the effects of management cessation on key habitat characteristics of four mountain forest bird species indicative of different structural components: Capercaillie (Tetrao urogallus), Hazel grouse (Bonasa bonasia), Three-toed woodpecker (Picoides tridactylus) and Pygmy owl (Glaucidium pas-serinum) across four mountain regions in Central Europe. Habitat suitability was modelled based on 300 forest sites selected independently of their management status, and predicted to an independent dataset of 42 strictly protected forest reserves in the same regions. We then compared forest reserves to managed forests with species presence or absence with regard to habitat suitability and key habitat structures and related both to the time since reserve designation. For all model species, except Pygmy owl, habitat suitability in forest reserves was significantly higher than in managed forests with species' absence, but not different from managed forests with species presence. For the species associated with open forest structures (Capercaillie, Hazel grouse, Pygmy owl) habitat suitability was significantly related to the "reserve age": reserves in the first three decades after management cessation showed a significant decrease in suitability, which increased afterwards up to the maximally recorded time of 100 years. No such correlation was found for the Three-toed woodpecker associated with deadwood and barkbeetle infestations following temporally unpredictable disturbance events. Structural characteristics varied greatly in abundance and distribution, with open structures being related to the time since reserve designation. We therefore recommend focusing on mature, near-natural and structurally diverse forests when designating new strict forest reserves.
Article
Managed conifer forests in temperate regions are critical for supplying wood products, but little is known about their potential for pollinator conservation. We hosted a workshop for Pacific Northwest managers and biologists to identify perceived information gaps regarding pollinators in managed conifer forests; we also undertook a literature review on this topic to assess gaps in the primary literature. The most important gaps identified by workshop participants were a need for baseline data on pollinators in managed conifer forests, and for determining how forest management influences pollinators. Our literature review found a dearth of pollinator studies in managed conifer forests, which were limited to few regions and a subset of taxa. Given these findings, we developed a research agenda that targets identified knowledge gaps, including the need for documenting fundamental aspects of pollinator ecology in managed conifer forests and testing how pollinators and their habitats are influenced by management activities.
Article
Pollinator communities exhibit variable responses to changing landscape composition. A general expectation is that a decreasing cover of semi-natural habitats negatively affects pollinator reproduction, population size and pollination services, but few studies have investigated the simultaneous effects of landscape complexity on different aspects of pollinator communities and functioning. In 20 agricultural landscape plots the size of an average Dutch farm, we studied how changing landscape complexity affected wild bee abundance, species richness and reproduction. To measure pollination, we placed potted strawberry plants as phytometers in landscapes. Landscape complexity was characterized as the area of semi-natural habitats. In addition, we estimated floral resource abundance in each landscape plot. We expected that i) bee species richness, reproduction and pollination would be positively related to area of semi-natural habitats and flower abundance, and that ii) species richness and reproduction would be positively related to pollination. An increase in semi-natural habitats in landscapes increased both the abundance of cavity-nesting bees colonizing trap nests, and the growth rates of experimental Bombus terrestris L. colonies, but not the species richness of wild bees measured by pan traps. There was only a tendency for higher pollination levels of strawberry plants with higher cover of semi-natural habitats. There was no relationship between species richness and bee reproduction in a landscape and the pollination services. Estimated flower abundance in landscape had a positive effect on bumblebee colony growth only and not on the other variables. Our results suggest that, by improving habitat quality on their farms through establishing more semi-natural habitats or enhancing the flower availability in semi-natural habitats, farmers can promote reproduction of a number of functionally important bee species and the pollination services they provide. Bee species richness, however, seems to be more difficult to enhance and requires more than just creating more of the same type of habitats or flowers.
Article
Intensification of agriculture has been one of the major drivers for biodiversity loss in recent decades. Pollinators, which serve an important role in pollinating crops as well as wild plants, have shown a decline in species richness. Flower strips can be used to support pollinators in agro-ecosystems, however the question remains as to how their design can be optimized in order to best benefit pollinators. Increasing plant species diversity has been shown to be beneficial for pollinators, and it is often suggested that functional traits are driving this relationship. Therefore, increasing plant functional diversity could be a tool to support pollinator abundance and diversity. As experimental evidence on this relationship is scarce, we developed a field study with experimental sown flower strips with four functional diversity levels, based on multiple flower traits and with equal plant species richness. We monitored vegetation development, as well as the flower-visiting pollinator community and their interaction networks with flowers. We were able to create a functional diversity gradient while controlling for plant species richness and evenness. However, in contrast to our expectations, pollinator species richness and evenness were not influenced by functional diversity, and increasing functional diversity even resulted in lower flower visitation rates. Network stability metrics showed no effect or negative relationships with functional diversity. We conclude that increasing functional diversity was not the key for supporting pollinators in wildflower strips. Our results also suggest that, for a constant amount of flower resources, increasing plant functional diversity and thus decreasing redundancy of potential pollinator feeding niches, decreases the amount of flower resources present per feeding niche. As pollinator species tended to have less overlap in their feeding niches in flower strips with increased functional diversity, this may lead to a reduction of flower resources available for pollinator species with a more specialized feeding niche.
Book
Dieses Lehrbuch gibt einen Überblick über das komplexe, in zahlreiche Fachdisziplinen aufgesplitterte Gebiet der Ökologie. Im Fokus steht dabei die Ökologie der Baumarten und der Wälder in Mitteleuropa, einem vegetationsgeografisch eigenständigen Naturraum, der sich von der Ostsee bis zu den Südalpen und von Ostfrankreich bis nach Polen und in die Slowakei erstreckt. Die Ausführungen zu den Strukturen und Prozessen in den natürlichen Waldformationen werden ergänzt durch Forschungsergebnisse zu den ökologischen Auswirkungen der Forstwirtschaft. In eigenen Kapiteln werden auch die globalen Problemfelder Biodiversität, Klimawandel und Waldschäden behandelt. Biologie, Geografie, Landschaftsplanung, Landschaftsökologie, Agrar- und Forstwissenschaften sind Studiengänge, in denen ökologische Themen seit jeher einen wichtigen Teil der Lehrinhalte einnehmen. Hinzu kamen neue Studiengänge für Ressourcenmanagement, Ökosystemmanagement, Natur- und Umweltwissenschaften. Diesen Fachrichtungen bietet das Lehrbuch eine Einführung in die ökologischen Zusammenhänge. Hierbei folgt es einem hierarchischen Aufbau von den Bäumen über die Baumpopulation und das Waldökosystem bis zur Landschaft. Die umfassende Berücksichtigung aktueller und spezieller Literatur macht das Lehrbuch auch zu einem Handbuch und Nachschlagewerk für Masterstudenten, Doktoranden und Dozenten.
Article
Anthropogenic disturbance of habitat is considered a contributing factor of pollinator declines, but some disturbances such as silviculture, may have positive implications for pollinator communities. Silviculture is a key source of disturbance in the eastern USA and thus, developing a better understanding of its ramifications for these keystone species is important for effective ecosystem conservation. We sampled bees in 30 forest openings created by group selection harvest as well as 30 sites in adjacent mature forest to examine the extent to which small forest openings support bees, to identify environmental variables influencing bee abundance and diversity, and to gauge their potential to augment bee populations in adjacent unmanaged forest. Bees were significantly more abundant and diverse in forest openings than in mature forest, but species composition did not differ. There was no relationship between opening size and abundance or diversity of bees in openings or adjacent mature forest. Both abundance and diversity were generally positively related to the amount of early-successional habitat on the landscape. Within openings, overall abundance and diversity decreased with vegetation height and increased with a metric representing floral richness and abundance. Notably, social, soft-wood-nesting, and small bees exhibited the opposite pattern in adjacent forest, increasing with vegetation height in openings and decreasing with greater floral richness and abundance within openings. Our results suggest that the creation of small forest openings helps to promote bees both in openings and adjacent mature forest, but this pattern is not consistent for all guilds.
Article
Bees are herbivorous insects, consuming nectar and pollen throughout their life cycles. This paper is a brief review of the chemistry of these two floral resources and the implications for bee nutrition. Nectar is primarily an energy source, but in addition to sugars contains various minor constituents that may, directly or indirectly, have nutritional significance. Pollen provides bees with the protein, lipids, vitamins and minerals that are essential for larval rearing. Chemical analyses of pollen have tended to focus on the protein component of bee-collected pollens as an index of nutritional quality. However, the substantial nectar content of such samples (~ 50% dry mass) should not be ignored, especially in view of current interest in measuring the nutritional quality of floral resources for bees.
Article
Recent studies have shown that the diversity of flowering plants can enhance pollinator richness and visitation frequency and thereby increase the resilience of pollination. It is assumed that flower traits explain these effects, but it is still unclear which flower traits are responsible, and knowing that, if pollinator richness and visitation frequency are more driven by mass-ratio effects (mean trait values) or by trait diversity. Here, we analyse a three-year data set of pollinator observations collected in a European grassland plant diversity experiment (The Jena experiment). The data entail comprehensive flower trait measurements, including reward traits (nectar and pollen amount), morphological traits (height, symmetry, area, colour spectra), and chemical traits (nectar-amino acid and nectar-sugar concentration). We test if pollinator species richness and visitation frequency of flower communities depend on overall functional diversity combining all flower traits within a community, single trait diversities (within trait variation) and community-weighted means of the single traits, using Bayesian inference. Overall functional diversity did not affect pollinator species richness, but reduced visitation frequency. When looking at individual flower traits separately, we found that single trait diversity of flower reflectance and flower morphology were important predictors of pollinator visitation frequency. Moreover, independent of total flower abundance, community-weighted means of flower height, area, reflectance, nectar-sugar concentration and nectar-amino acid concentration strongly affected both pollinator species richness and visitation frequency. Our results, challenge the idea that functional diversity always positively affects ecosystem functions. Nonetheless, we demonstrate that both single trait diversity and mass-ratio effects of flower traits play an important role for diverse and frequent flower visits, which underlines the functionality of flower traits for pollination services. This article is protected by copyright. All rights reserved.
Article
Understanding patterns of species diversity at different spatial scales is important for adapting management and conservation efforts. We have therefore studied wild bee and wasp (Hymenoptera: Aculeata) diversity structure in forest landscapes and evaluated the importance of conservation management at the local and landscape levels. Surveys were conducted at 32 clearcuts in eight landscapes in a managed boreal forest region. We assessed the influence of local habitat and landscape composition on species richness patterns and the effect of prescribed burning and landscape affinity on species composition for all bees and wasps as well as ecological and functional groups. The relative contribution of alpha and beta diversity on the regional level was assessed by diversity partitioning and the beta diversity between landscapes further partitioned into components of species turnover and nestedness. Bee and wasp species richness increased with high flower richness and clearcut size, and species composition differed between burned and unburned sites. Thus, flower-rich early-successional sites in boreal forest landscapes are important habitats for wild bees and wasps. To support this fauna, openness should be maintained for extended periods by delaying or avoiding tree plantation at flower-rich spots on clear-cuts. Beta diversity between landscapes accounted for the greatest proportion of the total regional gamma diversity, and over 70% of this diversity was due to species turnover. This implies that in order to maintain regional diversity, conservation efforts should be spatially dispersed, i.e. applied to every landscape of a few hundred square kilometers and adapted to the local species assemblages.