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Abstract Background Only a few decades ago, colorful, small‐scale, heterogeneous, and species‐rich hay meadows or extensive pastures were common, but have often been replaced by species‐poor, uniform, large‐scale multicut meadows. Technological advancements and improved efficiency in grassland management have come at the cost of biodiversity. Methods In Germany, 150 grassland plots have been investigated since 2006. Using these extensive data, we propose a new compound index for estimating the site‐specific mowing intensity in order to facilitate assessment of the impact of mowing intensity on biodiversity and ecosystem processes. Our index integrates the various qualitative components of mowing machine type, mowing height and use of a conditioner, with the annual number of cuts. Results The newly proposed index achieves a much finer gradation of mowing intensity compared to the previous quantification based on the number of cuts only. Furthermore, a decrease in plant and arthropod species was observed at higher mowing intensity. Conclusions The proposed mowing intensity index offers enhanced precision in calculations and can easily be integrated into assessments of land‐use intensity in grasslands. Further, it could serve as a basis for providing subsidies to farmers, who adopt low‐impact mowing practices.

In modified production landscapes, biodiversity faces unprecedented pressures from human actions, resulting in significant species declines of plant and animal taxa, including birds. Understanding the underlying mechanisms responsible for such declines is essential to counteract further loss and support practitioners in conserving biodiversity and associated ecosystem function. In this study, we used standardized bird monitoring data collected over 6 years in managed forest and grassland areas across different regions in Germany, Central Europe. We combined these data with morphometric, ecological, behavioral, and acoustic trait data and detailed information on local land use management practices to understand how management decisions affect species and functional diversity, as well as ecological processes shaping local species composition. Our results reveal that the ecosystem and regional context must be considered to understand how management practices affect bird diversity aspects and composition. In forests, regional management decisions related to tree species and stand age affected bird diversity, as well as community and functional composition, and indicated environmental sorting due to ecological and behavioral requirements, biotic interactions, and morphometric constraints. In grasslands, independent of local management practices, increased intensity of land use resulted in an overall loss in bird species richness and functional diversity. Predominantly, constraints due to ecological or behavioral requirements affected bird species assemblage composition. In addition, our results indicated the importance of woody vegetation near managed grasslands and of considering environmental conditions beyond the local scale to support bird diversity and associated ecosystem functions. Our results highlighted that local management decisions can support bird diversity and maintain ecological function. However, this needs a view beyond the local scale of management units. It also demands a joint effort of biologists and land managers to integrate targeted conservation actions into regional management practices and create a network of habitats within production landscapes to protect nature, guard against biotic and functional homogenization, and prevent further degradation of ecosystems in production landscapes.

Bees are the most important group of insect pollinators, but their populations are declining. To gain a better understanding of wild bee responses to different stressors (e.g. land-use change) and conservation measures, regional and national monitoring schemes are currently being established in Germany, which is used here as a model region, and in many other countries. We offer perspectives on how to best design future bee monitoring programs with a focus on evaluating the implementation of conservation measures. We discuss different traditional and novel sampling methods, their efficacy depending on research questions and the life-history traits of target species, and how greater standardization of wild bee sampling and monitoring methods can make data more comparable, contributing to the identification of general trends and mechanisms driving bee populations. Furthermore, the potential impact of bee sampling itself on bee populations is discussed.

Abstract Land‐use management is a key factor causing pollinator declines in agricultural grasslands. This decline can not only be directly driven by land‐use (e.g., habitat loss) but also be indirectly mediated through a reduction in floral resource abundance and diversity, which might in turn affect pollinator health and foraging. We conducted surveys of the abundance of flowering plant species and behavioural observations of two common generalist pollinator species, namely the bumblebee Bombus lapidarius and the syrphid fly Episyrphus balteatus, in managed grasslands of variable land‐use intensity (LUI) to investigate whether land‐use affects (1) resource availability of the pollinators, (2) their host plant selection and (3) pollinator foraging behaviour. We have found that the floral composition of plant species that were used as resource by the investigated pollinator species depends on land‐use intensity and practices such as mowing or grazing. We have also found that bumblebees, but not syrphid flies, visit different plants depending on LUI or management type. Furthermore, LUI indirectly changed pollinator behaviour via a reduction in plot‐level flower diversity and abundance. For example, bumblebees show longer flight durations with decreasing flower cover indicating higher energy expenditure when foraging on land‐use intensive plots. Syrphid flies were generally less affected by local land use, showing how different pollinator groups can differently react to land‐use change. Overall, we show that land‐use can change resource composition, abundance and diversity for pollinators, which can in turn affect pollinator foraging behaviour and potentially contribute to pollinator decline in agricultural grasslands.

IntroductionInteractions between flowering plants and visiting wild bees are crucial for ensuring pollination and subsequent plant reproductive success. However, bee diversity in an area has rarely been recorded in relation to seed set in native plants. In this project, we investigated the effect of local wild bee communities on seed set in four common wild plant species.MethodsPotted plants of Centaurea jacea, Cichorium intybus, Sinapis arvensis, and Salvia pratensis were placed for pollination experiments, in two distinct habitat types that we expected to show distinct bee communities, namely near-natural grassland and perennial wild flower strips, in Germany.ResultsOur results showed that near-natural grassland had a higher bee species richness and an increased seed set compared with flower strips that displayed a higher bee abundance in most study locations. Although we found effects of bee diversity on seed set, we did not detect a significant effect of bee abundance. Furthermore, the seed set in response to wild bee diversity differed across the plant species.DiscussionWe conclude that bee-friendly habitats ensure a high seed set in wild plants, but that the impact varies between different plant species.

Forest birds respond to a diverse set of environmental factors, including those altered by forest management intensity, such as resource and habitat availability in the form of food or nesting sites. Although resource/habitat availability and bird traits likely mediate responses of bird diversity to global change drivers, no study has assessed the direct and indirect effects of changes in forest management and traits on bird assemblages jointly at large spatial scales. In this context the questions remain whether (1) the birds' response to forest management changes through alterations in structural properties and/or food availability, or (2) if birds' eco-morphological traits act as environmental filters in response to environmental factors. We audio-visually recorded birds at 150 forest plots in three regions of Germany and assessed the forest structure (LiDAR) as well as the diversity of the herbaceous layer and diversity and biomass of arthropods. We further assessed eco-morphological traits of the birds and tested if effects on bird assemblages are mediated by changes in eco-morphological traits' composition. We found that abundance and species numbers of birds are explained best by models including the major environmental factors, forest structure, plants, and arthropods. Eco-morphological traits only increased model fit for indirect effects on abundance of birds. We found minor differences between the three regions in Germany, indicating spatial congruency of the processes at the local and regional scale. Our results suggest that most birds are not specialized on a particular food type, but that the size, diversity and species composition of arthropods are important. Our findings question the general view that bird traits adapt to the resources available.

The interaction between bees and flowering plants is mediated by floral cues that enable bees to find foraging plants. We tested floral cue preferences among three common wild bee species: Lasioglossum villosulum, Osmia bicornis, and Bombus terrestris. Preferences are well studied in eusocial bees but almost unknown in solitary or non-eusocial generalist bee species. Using standardized artificial flowers altered in single cues, we tested preferences for color hue, achromatic contrast, scent complexity, corolla size, and flower depth. We found common attractive cues among all tested bees. Intensively colored flowers and large floral displays were highly attractive. No preferences were observed in scent complexity experiments, and the number of volatiles did not influence the behavior of bees. Differing preferences were found for color hue. The specific behaviors were probably influenced by foraging experience and depended on the flower choice preferences of the tested bee species. In experiments testing different flower depths of reward presentation, the bees chose flat flowers that afforded low energy costs. The results reveal that generalist wild bee species other than well-studied honeybees and bumblebees show strong preferences for distinct floral cues to find potential host plants. The diverse preferences of wild bees ensure the pollination of various flowering plants.

Abstract Background Fleshy fruits evolved to be attractive to seed dispersers through various signals such as color and scent. Signals can evolve through different trajectories and have various degrees of reliability. The strongest substrate on which reliable signals can evolve is when there is an inherent link between signal and reward, rendering cheating costly or impossible. It was recently proposed that aliphatic esters in fruit scent may be predictive of sugar content due to their synthesis from products of sugar fermentation. We test this hypothesis on a case study of wild fig species (Ficus tiliifolia) from Madagascar, which relies on seed dispersal by lemurs. Results We found a strong positive correlation between signal (esters) and reward (sugar). We also found that non-esters, including direct fermentation products, in fruit scent do not indicate sugar levels, which implies that this relationship is not simply a product of fruit maturation wherein more mature fruits emit more scent and contain more sugar. Conclusions While based on a single taxon, these results strongly support the hypothesis that a biochemical link between ester synthesis and sugar may render the ester fraction of fruit scent an honest signal for fruit quality, with consequences for animal sensory and feeding ecology, and the evolution of plants in the context of seed dispersal.

Over the past few decades, several investigations around the globe have reported alarming declines in the abundance and diversity of bee species. The success of effective conservation strategies targeting these important pollinators relies heavily on accurate biodiversity assessments. The shortage of taxonomic experts and the escalation of the ongoing biodiversity crisis call for the development of alternative identification tools to implement efficient monitoring programs. The validation of such techniques is crucial to ensure that they provide results comparable to those of traditional morphotaxonomy. Here we performed two double-blind experiments to evaluate the accuracy of a pair of new techniques used for wild bee identification: DNA metabarcoding and in vivo identification in the field. The methods were tested on sets of wild bees from Germany and their results compared against evaluations done by panels of bee experts using traditional morphotaxonomy. On average the congruency of species identification between metabarcoding and morphotaxonomy was 88.98% across samples (N = 10), while in vivo identification and morphotaxonomy were 91.81% congruent (N = 7) for bees considered feasible for in vivo identification in the field. Traditional morphotaxonomy showed similar congruencies when compared to itself: 93.65% in the metabarcoding study and 92.96% in the in vivo study. Overall, these results support both new methods as viable alternatives to traditional microscopy-based assessment, with neither method being error-free. Metabarcoding provides a suitable option to analyze large numbers of specimens in the absence of highly trained taxonomic experts, while in vivo identification is recommended for repeated long-term monitoring, and when working in areas where the sampling of individuals could threaten local populations of endangered wild bee species. Further research is still needed to explore the potential of both techniques for conservation management and wildlife monitoring, as well as to overcome their current limitations as taxonomic tools.

Multi-kingdom community complexity and the chemically mediated dynamics between bacteria and insects have recently received increased attention in carrion research. However, the strength of these inter-kingdom interactions and the factors that regulate them are poorly studied. We used 75 piglet cadavers across three forest regions to survey the relationship between three actors (epinecrotic bacteria, volatile organic compounds (VOCs) and flies) during the first 4 days of decomposition and the factors that regulate this interdependence. The results showed a dynamic bacterial change during decomposition (temperature–time index) and across the forest management gradient, but not between regions. Similarly, VOC emission was dynamic across a temperature–time index and the forest management gradient but did not differ between regions. However, fly occurrence was dynamic across both space and time. The strong interdependence between the three actors was mainly regulated by the temperature–time index and the study regions, thereby revealing regulation at temporal and spatial scales. Additionally, the actor interdependence was stable across a gradient of forest management intensity. By combining different actors of decomposition, we have expanded our knowledge of the holistic mechanisms regulating carrion community dynamics and inter-kingdom interactions, an important precondition for better describing food web dynamics and entire ecosystem functions.

Land use intensification is a major driver of biodiversity change. Here the authors measure diversity across multiple trophic levels in agricultural grassland landscapes of varying management, finding decoupled responses of above- and belowground taxa to local factors and a strong impact of landscape-level land use.

Abstract Background Fruit scent is increasingly recognized as an evolved signal whose function is to attract animal seed dispersers and facilitate plant reproduction. However, like all traits, fruit scent is likely to evolve in response to conflicting selective pressures and various constraints. Two major constraints are (i) phylogenetic constraints, in which traits are inherited from ancestors rather than adapted to current conditions and (ii) developmental constraints, if phenotypes are limited by the expression of other traits within the individual. We tested whether phylogenetic constraints play a role in fruit scent evolution by calculating the phylogenetic signal in ripe fruits of 98 species from three study sites. We then estimated the importance of developmental constraints by examining whether ripe fruits tend to emit compounds that are chemically similar to, and share biosynthetic pathways with, compounds emitted by conspecific unripe fruits from which they develop. Results We show that closely related taxa are not more similar to each other than to very distinct taxa, thus indicating that fruit scent shows little phylogenetic signal. At the same time, although ripe and unripe fruits of the same species tend to emit different chemicals, they tend to employ chemicals originating from similar biosynthetic pathways, thus indicating that some developmental constraints determine ripe fruit scent. Conclusions Our results highlight the complex landscape in which fruit scent has evolved. On one hand, fruit scent evolution is not limited by common ancestry. On the other hand, the range of chemicals that can be employed in ripe fruits is probably constrained by the needs of unripe fruits.

Abstract Background In flowering communities, plant species commonly share pollinators and therefore plant individuals receive heterospecific pollen (HP). However, the patterns of HP transfers can deviate from patterns of plant-pollinator visitations. Although flower-visitor interactions are known to be mediated by floral traits, e.g. floral size or nectar tube depth, the explanatory power of these traits for HP transfer patterns remains elusive. Here, we have explored pollen transfer patterns at three sites in Southern Germany on three dates (early, mid and late summer). At the plant level, we tested whether flower abundance and floral traits are correlated with HP reception and donation. At the community level, we determined whether flower and bee diversity are correlated with network modularity and whether floral traits explain the module affiliation of plant species. We collected the stigmas of flowering plant species, analysed HP and conspecific pollen (CP) loads and measured floral traits, flower and bee diversity. Results Our results show that the degree and intensity of HP reception or donation at the plant level do not correlate with floral traits, whereas at the community level, the module affiliation of who is sharing pollen with whom is well-explained by floral traits. Additionally, variation in network modularity between communities is better explained by plant diversity and abundance than by bee diversity and abundance. Conclusions Overall, our results indicate that floral traits that are known to mediate flower-visitor interactions can improve our understanding of qualitative HP transfer but only provide limited information about the quantity of HP transfer, which more probably depends on other floral traits, flower-visitor identity or community properties.

Abstract The ability to assess food quality is crucial to all organisms. Fleshy fruits are a major source of nutrients to various animals, and unlike most food sources, have evolved to be attractive and to be consumed by animals to promote seed dispersal. It has recently been established that fruit scent—the bouquet of volatile chemicals emitted by ripe fruit—is an evolved communication system between plants and animals. Further, it has been argued that chemicals that are synthesized from sugar and its products may be an honest signal for sugar content and fruit quality. Elephants are important seed dispersers for numerous species and possess an olfactory system that is likely to outperform most other animals. We tested the hypothesis that fruit scent signifies sugar content and that elephants are capable of assessing fruit sugar levels based on scent alone. Using a paired‐choice test of marula fruits (Sclerocarya birrea) by semitame African elephants, we show that elephants are capable of identifying more sugar‐rich fruits based on scent alone and that this is likely based on two chemical compounds: ethanol and ethyl acetate, both downstream products of sugar fermentation. These results shed light on the mechanisms driving elephant feeding ecology, plant signaling, and the coevolutionary process between angiosperms and animal seed dispersers.

Abstract The sharing of the same food source among parents and offspring can be a driver of the evolution of family life and parental care. However, if all family members desire the same meal, competitive situations can arise, especially if resource depletion is likely. When food is shared for reproduction and the raising of offspring, parents have to decide whether they should invest in self‐maintenance or in their offspring and it is not entirely clear how these two strategies are balanced. In the burying beetle Nicrophorus vespilloides, parents care for their offspring either bi‐ or uniparentally at a vertebrate carcass as the sole food source. The question of whether biparental care in this species offers the offspring a better environment for development compared with uniparental care has been the subject of some debate. We tested the hypothesis that male contribution to biparental brood care has a beneficial effect on offspring fitness but that this effect can be masked because the male also feeds from the shared resource. We show that a mouse carcass prepared by two Nicrophorus beetles is lighter compared with a carcass prepared by a single female beetle at the start of larval hatching and provisioning. This difference in carcass mass can influence offspring fitness when food availability is limited, supporting our hypothesis. Our results provide new insights into the possible evolutionary pathway of biparental care in this species of burying beetles.

Pollinators and other insects are currently undergoing a massive decline. Several stressors are thought to be of importance in this decline, with those having close relationships to agricultural management and practice seemingly playing key roles. In the present study, we sampled Bombus lapidarius L. workers in grasslands differing in their management intensity and management regime across three different regions along a north-south gradient in Germany. We analyzed the bees with regard to (1) their cuticular hydrocarbon profile (because of its important role in communication in social insects) and amount of scent by using gas chromatography and (2) the size of each individual by using wing distances as a proxy for body size. Our analysis revealed changes related to land-use intensity and temperature in the cuticular scent profile of bumble bees. Decreasing body size and increasing total scent amount were explained by an interaction of land-use intensity and study region, but not by land-use intensity alone. Thus, land-use intensity and temperature influence intracolonial communication and size, both of which can have strong effects on foraging. Land management and climate are therefore probably detrimental for colony maintenance and the reproductive success of bumble bees.

Male euglossine bees exhibit unique adaptations for the acquisition and accumulation of chemical compounds from “perfume flowers” and other sources. During courtship display, male bees expose perfume mixtures, presumably to convey species-specific recognition and/or mate choice signals to females. Because olfaction regulates both signal production (in males) and signal detection (in females) in this communication system, strong selective pressures are expected to act on the olfactory system, which could lead to sensory specialization in favor of an increased sensitivity to specific chemical compounds. The floral scents of euglossine-pollinated plants are hypothesized to have evolved in response to the preexisting sensory biases of their male euglossine bee pollinators. However, this has never been investigated at the peripheral olfactory circuitry of distinct pollinating genera. Here, we present a comparative analysis using electroantennography (EAG) of males across the phylogeny of 29 euglossine bee species, among them Euglossa and Eulaema species. First, we tested whether antennal responses differ among different euglossine genera, subgenera and species. Secondly, we conducted a comparative phylogenetic analysis to investigate the macroevolutionary patterns of antennal responses across the euglossine bee phylogeny. We found that antennal response profiles are very unique on the species level and differ on the subgenus and the genus level. The differences can be explained by chemical compounds typically found in the floral scent bouquets of perfume flowers and specific compounds of species either pollinated by Euglossa (e.g., ipsdienol) or Eulaema bees (e.g., (−)-(E)-carvone epoxide). Also, we detected a phylogenetic signal in mean antennal responses and found that especially at the species level of our simulation the overall antennal responses exhibit greater disparity relative to a null model of pure Brownian-motion across the phylogeny. Altogether, our results suggest that (1) euglossine bee species exhibit species-specific antennal responses that differ among euglossine genera and subgenera, (2) antennal responses diverge early after speciation events, and (3) scent composition of perfume flowers evolved in response to pollinator-mediated selection imposed by preexisting sensory biases in euglossine bees.

Catasetum is a speciose Neotropical orchid genus of which male and female flowers emit scents acting both as attractant and reward for their exclusive pollinators, male orchid bees (Euglossini: Apidae). In Catasetum, it is well known that flowers display a remarkably morphological sexual dimorphism. However, it remains poorly investigated whether this is also true for floral scents. Here, we investigated the pollination ecology and floral scent traits (chemistry and total emission) of C. maranhense, a species endemic to the Brazilian N/NE region. Males of Euglossa securigera are the only pollinators of C. maranhense. The floral scent of C. maranhense is composed of 29 volatile compounds, with eucalyptol, indole, (E)-Methyl p-methoxycinnamate, and (Z)-Methyl p-methoxycinnamate accounting for more than 80% of the scent bouquet. No sexual dimorphism was detected in any of the traits investigated. We discuss the ecological and evolutionary significance of our findings to Catasetum species and other unisexual perfume plants.

Insect species richness and abundance has declined rapidly over the last few decades. Various stressors, such as the conversion of natural habitats, climate change, land-use intensification, agrochemicals and pathogens, are thought to be major factors in this decline. We treated female bees of two common pollinator species in Europe, Osmia bicornis and Bombus terrestris, with a field-realistic dose of the neonicotinoid clothianidin. We tested its effects on the foraging behavior of O. bicornis under semi-natural conditions and on the antennal sensitivity of both bee species to common floral volatiles by using electroantennography. Clothianidin negatively affected the foraging behavior in O. bicornis by decreasing the number of flowers visited per foraging flight and by increasing the time per flower visit and the searching time between two flowers. It also decreased the antennal sensitivity to 2-phenylethanol in the two bee species. Thus, clothianidin is clearly a threat for bees via its effects on their foraging behavior and antennal sensitivity and is hence probably detrimental for pollination and the reproductive success of bees.

Oligolectic bees are highly dependent on the availability of the host plants to which they are specialized. Nevertheless, females of Chelostoma rapunculi have recently been monitored occasionally to visit Malva moschata and Geranium sanguineum flowers, in addition to their well-known Campanula spp. hosts. The questions therefore arise which floral cues promote visits to non-host plants. As host-specific floral cues are key attractants for oligolectic bees, we have studied the attractiveness of olfactory and visual cues of the established host Campanula trachelium in comparison to the non-host plants G. sanguineum and M. moschata in behavioral experiments. Chemical and electrophysiological analyses of the floral scent and spectral measurements of floral colors were used to compare and contrast host and non-host plants. The behavioral experiments showed that foraging-naïve bees, in particular, were attracted by olfactory cues of the non-host plants, and that they did not favor the Campanula host scent in choice experiments. Many electrophysiologically active floral volatiles were present in common in the studied plants, although each species produced an individual scent profile. Spiroacetals, the key components that enable C. rapunculi to recognize Campanula hosts, were detected in trace amounts in Geranium but could not be proved to occur in Malva. The visual floral cues of all species were particularly attractive for foraging-experienced bees. The high attractiveness of G. sanguineum and M. moschata flowers to C. rapunculi bees and the floral traits that are similar to the Campanula host plants can be a first step to the beginning of a host expansion or change which, however, rarely occurs in oligolectic bees.

Abstract Plant species with fleshy fruits offer animals rewards such as sugar, protein, and fat, to feed on their fruits and disperse their seeds. They have also evolved visual and olfactory signals indicating their presence and ripeness. In some systems, fruit color serves as a reliable visual signal of nutrient content. Yet even though many volatile chemicals used as olfactory signals derive from nutrients animals seek, it is still unknown whether fruit scent encodes information regarding nutrient content in wild fruits. We examine the relationship between olfactory signals and nutrient rewards in 28 fruiting plant species in Madagascar. We measured the relative amounts of four chemical classes in fruit scent using gas chromatography and mass spectrometry, as well as the relative amounts of sugar and protein in fruit pulp. We found that protein levels are not associated with elevated amounts of chemically related volatile compounds in fruit scent. In contrast, sugar content is strongly associated with the chemical composition of fruit scent. To our knowledge, this is the first research to explore the connection between fruit chemical signals and nutrient rewards. Our results imply that in the case of sugar, fruit scent is predictive of nutrient content and hence an honest signal.

The availability of nesting resources influences the persistence and survival of bee communities. Although a positive effect of artificial nesting structures has frequently been shown for aboveground cavity-nesting wild bees, studies on below ground-nesting bees are rare. Artificial nesting hills designed to provide nesting habitats for ground-nesting bees were therefore established within the BienABest project in 20 regions across Germany. Wild bee communities were monitored for two consecutive years, accompanied by recordings of landscape and abiotic nest site variables. Bee activity and species richness increased from the first to the second year after establishment; this was particularly pronounced in landscapes with a low cover of semi-natural habitat. The nesting hills were successively colonized, indicating that they should exist for many years, thereby promoting a species-rich bee community. We recommend the construction of nesting hills on sun-exposed sites with a high thermal gain of the substrate because the bees prefer south-facing sites with high soil temperatures. Although the soil composition of the nesting hills plays a minor role, we suggest using local soil to match the needs of the local bee community. We conclude that artificial nesting structures for ground-nesting bees act as a valuable nesting resource for various bee species, particularly in highly degraded landscapes. We offer a construction and maintenance guide for the successful establishment of nesting hills for bee conservation.

Despite increased focus on elucidating the various reproductive strategies employed by orchids, we still have only a rather limited understanding of deceptive pollination systems that are not bee- or wasp-mediated. In Europe, the orchid Neotinea ustulata has been known to consist of two phenologically divergent varieties, neither of which provide rewards to its pollinators. However, detailed studies of their reproductive biology have been lacking. Our study aimed to characterize and understand the floral traits (i.e., morphology, color, and scent chemistry) and reproductive biology of N. ustulata. We found that the two varieties differ in all their floral traits; furthermore, while Neotinea ustulata var. ustulata appears to be pollinated by both bees (e.g., Anthophora, Bombus) and flies (e.g., Dilophus, Tachina), var. aestivalis is pollinated almost entirely by flies (i.e., Nowickia, Tachina). Tachinids were also found to be much more effective than bees in removing pollinaria, and we show experimentally that they use the characteristic dark inflorescence top as a cue for approaching inflorescences. Our results thus suggest that while both N. ustulata varieties rely on tachinids for pollination, they differ in their degree of specialization. Further studies are, however, needed to fully understand the reproductive strategy of N. ustulata varieties.

Abstract The outcomes of human‐elephant conflict range from expensive to fatal to both humans and elephants across Africa and Asia, which has prompted extensive efforts to mitigate it. Previous attempts have focused primarily on physical barriers, plant deterrent compounds, and elephant‐nuisance species. However, the handful of effective approaches are expensive and maintenance‐intensive. Here, we approach the problem from the perspective of the sensory ecology of fear, and demonstrate elephant aversion to olfactory signals of lions, African elephants' top predator. We find that elephants demonstrate aversion to olfactory predator signals, and moreover, to inexpensive synthetic mixtures mimicking these signals. This research illustrates the importance of predator avoidance even in the world's largest land animal, and provides the basis for its practical implication to mitigate a devastating and enduring problem.

Abstract Human–elephant conflict is a persistent problem across elephant home ranges, that results in economic damage to commercial and subsistence farmers, and physical harm and death to humans and elephants. This problem is likely to intensify with increased development, dwindling of natural habitats, and climate change‐driven environmental shifts. Various methods to mitigate human–elephant conflict have been employed, but to date these have been hampered by financial and logistical considerations. Based on the fact that African elephants are predated by lions and possess a remarkable sense of smell, we hypothesize that elephants are strongly averse to olfactory signals of lion presence, and that this can be utilized to create invisible barriers which elephants will not cross. We conducted a series of tests that show that lion dung is an effective deterrent of elephants. We conducted chemical analyses of lion dung and identified the main compounds. We then used synthetic mixtures containing these compounds, and show that they successfully elicit the deterrence effect, even in miniscule concentrations. These results indicate that elephants can be deterred using simple and low‐concentration mixtures based on available commercial products, that can be developed into products that offer a safe, sustainable, and cost‐effective method to mitigate human–elephant conflict.

Climate change is leading to increasing drought and higher temperatures, both of which reduce soil water levels and consequently water availability for plants. This reduction often induces physiological stress in plants, which in turn can affect floral development and production inducing phenotypic alterations in flowers. Because flower visitors notice and respond to small differences in floral phenotypes, changes in trait expression can alter trait-mediated flower visitor behavior. Temperature is also known to affect floral scent emission and foraging behavior and, therefore, might modulate trait-mediated flower visitor behavior. However, the link between changes in flower visitor behavior and floral traits in the context of increasing drought and temperature is still not fully understood. In a wind-tunnel experiment, we tested the behavior of 66 Bombus terrestris individuals in response to watered and drought-stressed Sinapis arvensis plants and determined whether these responses were modulated by air temperature. Further, we explored whether floral traits and drought treatment were correlated with bumblebee behavior. The initial attractiveness of drought and watered plants did not differ, as the time to first visit was similar. However, bumblebees visited watered plants more often, their visitation rate to flowers was higher on watered plants, and bumblebees stayed for longer, indicating that watered plants were more attractive for foraging. Bumblebee behavior differed between floral trait expressions, mostly independently of treatment, with larger inflorescences and flowers leading to a decrease in the time until the first flower visit and an increase in the number of visits and the flower visitation rate. Temperature modulated bumblebee activity, which was highest at 25°C; the interaction of drought/water treatment and temperature led to higher visitation rate on watered plants at 20°C, possibly as a result of higher nectar production. Thus, bumblebee behavior is influenced by the watered status of plants, and bumblebees can recognize differences in intraspecific phenotypes involving morphological traits and scent emission, despite overall morphological traits and scent emission not being clearly separated between treatments. Our results indicate that plants are able to buffer floral trait expressions against short-term drought events, potentially to maintain pollinator attraction.

Social vespid wasps regularly forage on flowers with a generalist pollination system. However, little is known about communication between wasps and their host plants. The aim of this study was to investigate the role of olfactory and visual floral signals of Hedera helix and Heracleum sphondylium, both frequently visited by Vespula and Dolichovespula wasps for the collection of nectar. A combination of chemical, electrophysiological and spectral analyses and behavioral experiments was used to identify attractive floral signals. We identified 37 and 41 EAD-active substances (mainly terpenoids and aromatics) in H. helix and H. sphondylium, respectively. The most abundant floral compounds were 4-oxoisophorone in H. helix and linalool in H. sphondylium, followed by (E)-linalool oxide furanoid in both plants. The olfactory signals were attractive for wasps; however, a combination of olfactory and visual signals made both plants more attractive than olfactory signals alone and, in the case of H. helix, also than visual traits alone. Visual traits were not attractive by themselves. Wasps were also attracted by a synthetic solution resembling the floral scent of H. helix. Our study contributes to a better understanding of the foraging behavior of vespid wasps and describes, for the first-time, floral signals that attract vespine wasps to generalist flowers.

Dung beetles provide crucial ecosystem services and serve as model organisms for various behavioural, ecological and evolutionary studies. However, dung beetles have received little attention as consumers of large cadavers. In this study, we trapped copronecrophagous dung beetles on above-ground exposed piglet cadavers in 61 forest plots distributed over three geographically distinct regions in Germany, Central Europe. We examined the effects of land use intensity, forest stand, soil characteristics, vascular plant diversity and climatic conditions on dung beetle abundance, species richness and diversity. In all three regions, dung beetles, represented mainly by the geotrupid species Anoplotrupes stercorosus and Trypocopris vernalis, were attracted to the cadavers. High beetle abundance was associated with higher mean ambient temperature. Furthermore, A. stercorosus and T. vernalis were more abundant in areas where soil contained higher proportions of fine sand. Additionally, an increased proportion of forest understorey vegetation and vascular plant diversity positively affected the species richness and diversity of dung beetles. Thus, even in warm dry monocultured forest stands exploited for timber, we found thriving dung beetle populations when a diverse understorey was present. Therefore, forestry practices that preserve the understorey can sustain stable dung beetle populations and ensure their important contribution to nutrient cycles.

Visual and olfactory floral signals play key roles in plant-pollinator interactions. In recent decades, studies investigating the evolution of either of these signals have increased considerably. However, there are large gaps in our understanding of whether or not these two cue modalities evolve in a concerted manner. Here, we characterized the visual (i.e., color) and olfactory (scent) floral cues in bee-pollinated Campanula species by spectrophotometric and chemical methods, respectively, with the aim of tracing their evolutionary paths. We found a species-specific pattern in color reflectance and scent chemistry. Multivariate phylogenetic statistics revealed no influence of phylogeny on floral color and scent bouquet. However, univariate phylogenetic statistics revealed a phylogenetic signal in some of the constituents of the scent bouquet. Our results suggest unequal evolutionary pathways of visual and olfactory floral cues in the genus Campanula. While the lack of phylogenetic signal on both color and scent bouquet points to external agents (e.g., pollinators, herbivores) as evolutionary drivers, the presence of phylogenetic signal in at least some floral scent constituents point to an influence of phylogeny on trait evolution. We discuss why external agents and phylogeny differently shape the evolutionary paths in floral color and scent of closely related angiosperms.

Research into vibrational communication is becoming increasingly important as we discover more species using vibrational signals in different types of behavior. Males of the solitary red mason bee, Osmia bicornis , are known to use vibrations in female choice, where the females not only evaluate a male’s fitness through their vibrational signal but also use them to distinguish between males of different origins. This was shown conclusively via bioassays, in which females from Germany rejected English males unless they were imbued with an artificial German signal and vice versa. However, an investigation into which parameters of the signal might differ between species and populations has been lacking so far. We therefore recorded O. bicornis as well as O. cornuta males from Germany, England, and Denmark using a laser vibrometer and analyzed the recordings using the software Spike to measure frequency, modulation range, and average pulse duration in each signal. Our results clearly showed significant differences in the signal between O. bicornis and O. cornuta males in all three parameters measured. O. bicornis populations from Germany and Denmark surprisingly also differed significantly in frequency and modulation range, with England lying in between the two. We believe that the females probably use another signal parameter that we have not evaluated yet to make their mate choice. This leaves us with the question of why the observed signal parameters differ between Germany and Denmark. From our knowledge about the system to date, we believe that we are looking at the first steps of speciation in this system and further study could help us with new insights into evolutionary processes in the future.

The plant genus Ficus is a keystone resource in tropical ecoystems. One of the unique features of figs is the diversity of fruit traits, which in many cases match their various dispersers, the so-called fruit syndromes. The classic example of this is the strong phenotypic differences found between figs with bat and bird dispersers (color, size, presentation, and scent). The 'bird-fig' Ficus colubrinae represents an exception to this trend since it attracts the small frugivorous bat species Ectophylla alba at night, but during the day it attracts bird visitors. Here we investigate day to night changes in fruit scent as a possible mechanism by which this 'bird-fig' could attract bats despite its fruit traits, which should appeal solely to birds. Analyses of odor bouquets from the bat- and bird-dispersal phases (i.e. day and night) differed significantly in their composition of volatiles. We observed a significant increase in relative amounts of sesquiterpene and aromatic compounds at night while relative amounts of two compounds of the fatty acid pathway were significantly higher during day. This finding raises the question whether Ficus colubrinae, a phenotypically classic 'bird-fig', might be able to attract bat dispersers by an olfactory signal at night. Preliminary observations from feeding experiments which indicate that Ectophylla alba is capable of finding ripe figs by scent alone point in this direction. However, additional behavioral experiments on whether bats prefer the 'night-bouquet' over the 'day-bouquet' will be needed to unequivocally answer this question.

Parents are faced with the dilemma whether to invest in their current offspring, or potential future young. Here, Engel et al. show that nutritionally-dependent young induce temporary infertility in female burying beetles, which in turn is signalled to the male partner via a hormone-related anti-aphrodisiac.

Intensification of anthropogenic land use is a major threat to biodiversity and thus to essential ecosystem services provided by insects. Rove beetles (Coleoptera: Staphylinidae), which react sensitively to habitat changes, are species-rich colonizers of vertebrate cadavers and contribute to the important ecosystem service of carrion decomposition. The unveiling of anthropogenic and environmental drivers that modify carrion-associated rove beetle communities should improve our understanding of the plasticity of cadaver decay. We report the presence of 80 rove beetle species on 65 decomposing piglet cadavers at forest sites characterized by a gradient of management intensity across three geographic regions in Germany. Local and landscape drivers were revealed that shape beetle abundance, diversity, and community composition. Forest management and regions affect rove beetle abundance, whereas diversity is influenced by local habitat parameters (soil pH, litter cover) and regions. The community composition of rove beetles changes with management intensification by promoting generalist species. Regarding single species, Philonthus decorus and Anotylus mutator are linked to unmanaged forests and Ontholestes tessellatus to highly used forest stands. The spatial information provided about carrion-associated rove beetle communities in German forests is not only of carrion-ecological but also of forensic entomological interest.

During decomposition, vertebrate carrion emits volatile organic compounds to which insects and other scavengers are attracted. We have previously found that the dung beetle, Anoplotrupes stercorosus, is the most common dung beetle found on vertebrate cadavers. Our aim in this study was to identify volatile key compounds emitted from carrion and used by A. stercorosus to locate this nutritive resource. By collecting cadaveric volatiles and performing electroantennographic detection, we tested which compounds A. stercorosus perceived in the post-bloating decomposition stage. Receptors in the antennae of A. stercorosus responded to 24 volatiles in odor bouquets from post-bloating decay. Subsequently, we produced a synthetic cadaver odor bouquet consisting of six compounds (benzaldehyde, DMTS, 3-octanone, 6-methyl-5-hepten-2-ol, nonanal, dodecane) perceived by the beetles and used various blends to attract A. stercorosus in German forests. In field assays, these beetles were attracted to a blend of DMTS, 3-octanone, and benzaldehyde. Generalist feeding behavior might lead to the super-dominant occurrence of A. stercorosus in temperate European forests and have a potentially large impact on the exploitation and rapid turnover of temporally limited resources such as vertebrate cadavers.

In some insect species, females may base their choice for a suitable mate on male odor. In the red mason bee, Osmia bicornis, female choice is based on a male's odor bouquet as well as its thorax vibrations, and its relatedness to the female, a putative form of optimal outbreeding. Interestingly, O. bicornis can be found as two distinct color morphs in Europe, which are thought to represent subspecies and between which we hypothesize that female discrimination may be particularly marked. Here we investigated (i) if these two colors morphs do indeed represent distinct, reproductively differentiated populations, (ii) how odor bouquets of male O. bicornis vary within and between populations, and (iii) whether variation in male odor correlates with genetic distance, which might represent a cue by which females could optimally outbreed. Using GC and GC-MS analysis of male odors and microsatellite analysis of males and females from 9 populations, we show that, in Denmark, an area of subspecies sympatry, the two color morphs at any one site do not differ, either in odor bouquet or in population genetic differentiation. Yet populations across Europe are distinct in their odor profile as well as being genetically differentiated. Odor differences do not, however, mirror genetic differentiation between populations. We hypothesize that populations from Germany, England and Denmark may be under sexual selection through female choice for local odor profiles, which are not related to color morph though which could ultimately lead to population divergence and speciation.

Anthropogenic land use causes global declines in biodiversity. Despite the knowledge that animal carrion is the most nutrient-rich form of dead organic matter, studies on landscape and local scales determining whether and the means by which land use intensity influences the diversity of the carrion-associated insect fauna are globally scarce. We investigated the effects of land use intensity and abiotic and biotic environmental factors on the abundance, species richness, and diversity of the important ecosystem-service-providing silphid beetle taxon (carrion beetles) in three regions of Germany. In 61 forest stands distributed over three geographically distinct regions in Central Europe, we trapped silphid beetles on exposed piglet cadavers during late summer. In all three regions, higher ambient temperatures and higher fine sand contents were associated with the abundance of the silphid beetle taxa. The carrion community silphid diversity was negatively affected by an increase in mean ambient temperature in all three regions. Although management intensity in forests did not affect the overall abundance of Silphidae, the abundance of Nicrophorus humator decreased significantly with higher forest management intensity across all three regions. Unmanaged and age-class forests showed a higher abundance of N. humator compared with extensively managed forest stands. These findings indicate that N. humator has potential as an indicator species for anthropogenic disturbances in forests. Overall, the direct responses of the silphid beetle community to diverse soil characteristics underline soil as an important factor determining the abundance and diversity of necrophagous carrion beetles in Central Europe. To protect these valuable ecosystem-service providers, forest-management-induced soil modifications need to be paid close attention.

Queen signals are known to regulate reproductive harmony within eusocial colonies by influencing worker behavior and ovarian physiology. However, decades of research have resulted in the identification of just a few queen signals, and studies of their mode of action are rare. Our aim was to identify queen recognition signals in the halictid bee Lasioglossum pauxillum and to analyze caste differences in the olfactory perception of queen signals in L. pauxillum and the closely related species L. malachurum. We performed chemical analyses and bioassays to test for caste differences in chemical profiles and worker behavior influenced by queen-specific compounds in L. pauxillum. Our results indicated that caste differences in the chemical profiles were mainly attributable to higher amounts of macrocyclic lactones in queens. Bioassays demonstrated a higher frequency of subordinate behavior in workers elicited by queen-specific amounts of macrocyclic lactones. Thus, macrocyclic lactones function as queen recognition signals in L. pauxillum, as in L. malachurum. Using electrophysiological analyses, we have demonstrated that queens of both tested species lack antennal reactions to certain macrocyclic lactones. Therefore, we assume that this is a mechanism to prevent reproductive self-inhibition in queens. Our results should stimulate debate on the conservation and mode of action of queen signals.

Nectar-feeding bats depend mainly on floral nectar to fulfil their energetic requirements. Chiropterophilous flowers generally present strong floral scents and provide conspicuous acoustic echoes to attract bats. While floral scents are assumed to attract bats over long distances, acoustic properties of flower structures may provide detailed information, thus supporting the localization of a flower at close ranges. So far, to our knowledge, there is no study trying to understand the relative importance as well as the combination of these generally coupled cues for detection (presence) and localization (exact position) of open flowers in nature. For a better comprehension of the significance of olfaction and echolocation in the foraging behaviour of nectar-feeding bats, we conducted two-choice experiments with Leptonycteris yerbabuenae. We tested the bats' behaviour in three experimental scenarios with different cues: (i) olfaction versus echolocation, (ii) echolocation versus echolocation and olfaction, and (iii) olfaction versus echolocation and olfaction. We used the floral scent of the bat-pollinated cactus Pachycereus pringlei as olfactory cue and an acrylic paraboloid as acoustic cue. Additionally, we recorded the echolocation behaviour of the bats and analysed the floral scent of P. pringlei. When decoupled cues were offered, bats displayed no preference in choice for any of the two cues. However, bats reacted first to and chose more often the coupled cues. All bats echolocated continuously and broadcast a long terminal group before a successful visit. The floral scent bouquet of P. pringlei is composed of 20 compounds, some of which (e.g. methyl benzoate) were already reported from chiropterophilous plants. Our investigation demonstrates for the first time to our knowledge, that nectar-feeding bats integrate over different sensory modes for detection and precise localization of open flowers. The combined information from olfactory and acoustic cues allows bats to forage more efficiently.

Several neotropical orchid genera have been proposed as being sexually deceptive; however, this has been carefully tested in only a few cases. The genus Telipogon has long been assumed to be pollinated by male tachinid flies during pseudocopulatory events but no detailed confirmatory reports are available. Here, we have used an array of methods to elucidate the pollination mechanism in Telipogon peruvianus. The species presents flowers that have a mean floral longevity of 33 days and that are self-compatible, although spontaneous self-pollination does not occur. The flowers attract males of four tachinid species but only the males of an undescribed Eudejeania (Eudejeania aff. browni; Tachinidae) species are specific pollinators. Males visit the flowers during the first few hours of the day and the pollination success is very high (42% in one patch) compared with other sexually deceptive species. Female-seeking males are attracted to the flowers but do not attempt copulation with the flowers, as is usually described in sexually deceptive species. Nevertheless, morphological analysis and behavioural tests have shown an imperfect mimicry between flowers and females suggesting that the attractant stimulus is not based only on visual cues, as long thought. Challenging previous conclusions, our chemical analysis has confirmed that flowers of Telipogon release volatile compounds; however, the role of these volatiles in pollinator behaviour remains to be established. Pollinator behaviour and histological analyses indicate that Telipogon flowers possess scent-producing structures throughout the corolla. Our study provides the first confirmed case of (i) a sexually deceptive species in the Onciidinae, (ii) pollination by pre-copulatory behaviour and (iii) pollination by sexual deception involving tachinid flies.

Social insects are well known for their high level of cooperation. Workers of the primitively eusocial bumblebee Bombus terrestris are able to produce male offspring in the presence of a queen. Nonetheless, they only compete for reproduction, in the so-called competition phase, when the workforce is large enough to support the rearing of reproductives. So far, little is known about the proximate mechanisms underlying the shift between altruism and selfish behaviour in bumblebee workers. In this study, we have examined the influence of chemical cues from the nest wax on the onset of worker reproduction. Chemical analyses of wax extracts have revealed that the patterns and amounts of cuticular lipids change considerably during colony development. These changes in wax scent mirror worker abundance and the presence of fertile workers. In bioassays with queen-right worker groups, wax affects the dominance behaviour and ovarian development of workers. When exposed to wax from a colony in competition phase, workers start to compete for reproduction. We suggest that wax scent enables workers to time their reproduction by providing essential information concerning the social condition of the colony.

Oligolectic bees collect pollen from a few plants within a genus or family to rear their offspring, and are known to rely on visual and olfactory floral cues to recognize host plants. However, studies investigating whether oligolectic bees recognize distinct host plants by using shared floral cues are scarce. In the present study, we investigated in a comparative approach the visual and olfactory floral cues of six Campanula species, of which only Campanula lactiflora has never been reported as a pollen source of the oligolectic bee Ch. rapunculi. We hypothesized that the flowers of Campanula species visited by Ch. rapunculi share visual (i.e. color) and/or olfactory cues (scents) that give them a host-specific signature. To test this hypothesis, floral color and scent were studied by spectrophotometric and chemical analyses, respectively. Additionally, we performed bioassays within a flight cage to test the innate color preference of Ch. rapunculi. Our results show that Campanula flowers reflect the light predominantly in the UV-blue/blue bee-color space and that Ch. rapunculi displays a strong innate preference for these two colors. Furthermore, we recorded spiroacetals in the floral scent of all Campanula species, but Ca. lactiflora. Spiroacetals, rarely found as floral scent constituents but quite common among Campanula species, were recently shown to play a key function for host-flower recognition by Ch. rapunculi. We conclude that Campanula species share some visual and olfactory floral cues, and that neurological adaptations (i.e. vision and olfaction) of Ch. rapunculi innately drive their foraging flights toward host flowers. The significance of our findings for the evolution of pollen diet breadth in bees is discussed.

The necrophagous burying beetle Nicrophorus vespilloides reproduces on small carcasses that are buried underground to serve as food for their offspring. Cadavers that are too large to bury have previously been postulated to be important food sources for newly emerged beetles; however, the attractiveness of distinct successive stages of decomposition were not further specified. Therefore, we investigated the potential preference of newly emerged N. vespilloides females for odour bouquets of piglet cadavers at specific stages of decomposition. Analyses of walking tracks on a Kramer sphere revealed a significantly higher mean walking speed and, consequently, a higher mean total track length when beetles were confronted with odour plumes of the decomposition stages 'post-bloating', 'advanced decay' or 'dry remains' in comparison with the solvent control. Such a change of the walking speed of newly emerged N. vespilloides females indicates a higher motivation to locate such food sources. In contrast to less discriminating individuals this behaviour provides the advantage of not wasting time at unsuitable food sources. Furthermore, in the advanced decay stage, we registered a significantly higher preference of beetles for upwind directions to its specific odour plume when compared with the solvent control. Such a change to upwind walking behaviour increases the likelihood that a large cadaver will be quickly located. Our findings are of general importance for applied forensic entomology: newly emerged N. vespilloides females on large cadavers can and should be regarded as potential indicators of prolonged post mortem intervals as our results clearly show that they prefer emitted odour bouquets of later decomposition stages.

Social parasites exploit the colony resources of social insects. Some of them exploit the host colony as a food resource or as a shelter whereas other species also exploit the brood care behavior of their social host. Some of these species have even lost the worker caste and rely completely on the host's worker force to rear their offspring. To avoid host defenses and bypass their recognition code, these social parasites have developed several sophisticated chemical infiltration strategies. These infiltration strategies have been highly studied in several hymenopterans. Once a social parasite has successfully entered a host nest and integrated its social system, its emerging offspring still face the same challenge of avoiding host recognition. However, the strategy used by the offspring to survive within the host nest without being killed is still poorly documented. In cuckoo bumblebees, the parasite males completely lack the morphological and chemical adaptations to social parasitism that the females possess. Moreover, young parasite males exhibit an early production of species-specific cephalic secretions, used as sexual pheromones. Host workers might thus be able to recognize them. Here we used a bumblebee host-social parasite system to test the hypothesis that social parasite male offspring exhibit a chemical defense strategy to escape from host aggression during their intranidal life. Using behavioral assays, we showed that extracts from the heads of young cuckoo bumblebee males contain a repellent odor that prevents parasite males from being attacked by host workers. We also show that social parasitism reduces host worker aggressiveness and helps parasite offspring acceptance.