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Climate change will cause geographic range shifts for pollinators and major crops, with global implications for food security and rural livelihoods. However, little is known about the potential for coupled impacts of climate change on pollinators and crops. Coffee production exemplifies this issue, because large losses in areas suitable for coffee production have been projected due to climate change and because coffee production is dependent on bee pollination. We modeled the potential distributions of coffee and coffee pollinators under current and future climates in Latin America to understand whether future coffee-suitable areas will also be suitable for pollinators. Our results suggest that coffee-suitable areas will be reduced 73-88% by 2050 across warming scenarios, a decline 46-76% greater than estimated by global assessments. Mean bee richness will decline 8-18% within future coffee-suitable areas, but all are predicted to contain at least 5 bee species, and 46-59% of future coffee-suitable areas will contain 10 or more species. In our models, coffee suitability and bee richness each increase (i.e., positive coupling) in 10-22% of future coffee-suitable areas. Diminished coffee suitability and bee richness (i.e., negative coupling), however, occur in 34-51% of other areas. Finally, in 31-33% of the future coffee distribution areas, bee richness decreases and coffee suitability increases. Assessing coupled effects of climate change on crop suitability and pollination can help target appropriate management practices, including forest conservation, shade adjustment, crop rotation, or status quo, in different regions., Competing Interests: The authors declare no conflict of interest.

Patterns of metabolic rate variation have been documented extensively in animals, but their functional basis remains elusive. The membrane pacemaker hypothesis proposes that the relative abundance of polyunsaturated fatty acids in membrane phospholipids sets the metabolic rate of organisms. Using species of tropical orchid bees spanning a 16-fold range in body size, we show that the flight muscles of smaller bees have more linoleate (%18 : 3) and stearate (%18 : 0), but less oleate (%18 : 1). More importantly, flight metabolic rate (FlightMR) varies with the relative abundance of 18 : 3 according to the predictions of the membrane pacemaker hypothesis. Although this relationship was found across large differences in metabolic rate, a direct association could not be detected when taking phylogeny and body mass into account. Higher FlightMR, however, was related to lower %16 : 0, independent of phylogeny and body mass. Therefore, this study shows that flight muscle membrane composition plays a significant role in explaining diversity in FlightMR, but that body mass and phylogeny are other factors contributing to their variation. Multiple factors are at play to modulate metabolic capacity, and changing membrane composition can have gradual and stepwise effects to achieve a new range of metabolic rates. Orchid bees illustrate the correlated evolution between membrane composition and metabolic rate, supporting the functional link proposed in the membrane pacemaker hypothesis., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

Recently there has been considerable concern about declines in bee communities in agricultural and natural habitats. The value of pollination to agriculture, provided primarily by bees, is >$200 billion/year worldwide, and in natural ecosystems it is thought to be even greater. However, no monitoring program exists to accurately detect declines in abundance of insect pollinators; thus, it is difficult to quantify the status of bee communities or estimate the extent of declines. We used data from 11 multiyear studies of bee communities to devise a program to monitor pollinators at regional, national, or international scales. In these studies, 7 different methods for sampling bees were used and bees were sampled on 3 different continents. We estimated that a monitoring program with 200-250 sampling locations each sampled twice over 5 years would provide sufficient power to detect small (2-5%) annual declines in the number of species and in total abundance and would cost U.S.$2,000,000. To detect declines as small as 1% annually over the same period would require >300 sampling locations. Given the role of pollinators in food security and ecosystem function, we recommend establishment of integrated regional and international monitoring programs to detect changes in pollinator communities., (© 2012 Society for Conservation Biology.)

Stingless bees (Meliponini) constitute a diverse group of highly eusocial insects that occur throughout tropical regions around the world. The meliponine genus Melipona is restricted to the New World tropics and has over 50 described species. Melipona, like Apis, possesses the remarkable ability to use representational communication to indicate the location of foraging patches. Although Melipona has been the subject of numerous behavioral, ecological, and genetic studies, the evolutionary history of this genus remains largely unexplored. Here, we implement a multigene phylogenetic approach based on nuclear, mitochondrial, and ribosomal loci, coupled with molecular clock methods, to elucidate the phylogenetic relationships and antiquity of subgenera and species of Melipona. Our phylogenetic analysis resolves the relationship among subgenera and tends to agree with morphology-based classification hypotheses. Our molecular clock analysis indicates that the genus Melipona shared a most recent common ancestor at least approximately 14-17 million years (My) ago. These results provide the groundwork for future comparative analyses aimed at understanding the evolution of complex communication mechanisms in eusocial Apidae., (Copyright 2010 Elsevier Inc. All rights reserved.)

Neotropical orchid bees (Euglossini) are often cited as classic examples of trapline-foragers with potentially extensive foraging ranges. If long-distance movements are habitual, rare plants in widely scattered locations may benefit from euglossine pollination services. Here we report the first successful use of micro radio telemetry to track the movement of an insect pollinator in a complex and forested environment. Our results indicate that individual male orchid bees (Exaerete frontalis) habitually use large rainforest areas (at least 42-115 ha) on a daily basis. Aerial telemetry located individuals up to 5 km away from their core areas, and bees were often stationary, for variable periods, between flights to successive localities. These data suggest a higher degree of site fidelity than what may be expected in a free living male bee, and has implications for our understanding of biological activity patterns and the evolution of forest pollinators.

The widely accepted idea that bees fuel flight through the oxidation of carbohydrate is based on studies of only a few species. We tested this hypothesis as part of our research program to investigate the size-dependence of flight energetics in Panamanian orchid bees. We succeeded in measuring rates of O(2) consumption and CO(2) production in vivo during hovering flight, as well as maximal activities (V(max) values) in vitro of key enzymes in flight muscle energy metabolism in nine species belonging to four genera. Respiratory quotients (ratios of rates of CO(2) production to O(2) consumption) in all nine species are close to 1.0. This indicates that carbohydrate is the main fuel used for flight. Trehalase, glycogen phosphorylase and hexokinase activities are sufficient to account for the glycolytic flux rates estimated from rates of CO(2) production. High activities of other glycolytic enzymes, as well as high activities of mitochondrial oxidative enzymes, are consistent with the estimated rates of carbohydrate-fueled oxidative metabolism. In contrast, hydroxyacylCoA dehydrogenase, an enzyme involved in fatty acid oxidation, was not detectable in any species. Thoracic homogenates displayed ADP-stimulated oxidition of pyruvate + proline, but did not oxidize palmitoyl l-carnitine + proline as substrates. A metabolic map, based on data reported herein and information from the literature, is presented. The evidence available supports the hypothesis that carbohydrate serves as the main fuel for flight in bees.

The relationship between body size and flight energetics was studied in the clade of tropical orchid bees, in order to investigate energy metabolism and evolution. Body mass, which varied from 47 to 1065 mg, was found to strongly affect hovering flight mass-specific metabolic rates, which ranged from 114 ml CO(2) h(-1) g(-1) in small species to 37 ml CO(2) h(-1) g(-1) in large species. Similar variation of wingbeat frequency in hovering flight occurred among small to large species, and ranged from 250 to 86 Hz. The direct relationship between such traits was studied by the comparative method of phylogenetically independent contrasts (PIC), using a new molecular phylogeny generated from the cytochrome b gene partial sequences. We found wingbeat frequency variation is satisfactorily explained by variation in wing loading, after corrections for body mass and phylogeny. The correlated evolution of mass-specific metabolic rate, wingbeat frequency and wing loading was also revealed after correcting for phylogeny and body mass. Further, the effect of body size on flight energetics can be understood in terms of a relationship between wing form and kinematics, which directly influence and explain the scaling of metabolic rate in this group of bees.

The evolution of metabolic pathways involved in energy production was studied in the flight muscles of 28 species of orchid bees. Previous work revealed that wingbeat frequencies and mass-specific metabolic rates decline in parallel by threefold as body mass increases interspecifically over a 20-fold range. We investigated the correlated evolution of metabolic rates during hovering flight and the flux capacities, i.e. V(max) values, of flight muscle enzymes involved in substrate catabolism, the Krebs cycle and the electron transport chain. V(max) at the hexokinase (HK) step scales allometrically with an exponent almost identical to those obtained for wingbeat frequency and mass-specific metabolic rate. Analysis of this relationship using phylogenetically independent contrasts supports the hypothesis of correlated evolution between HK activity and mass-specific metabolic rate. Although other enzymes scale allometrically with respect to body mass, e.g. trehalase, glycogen phosphorylase and citrate synthase, no other enzyme activities were correlated with metabolic rate after controlling for phylogenetic relatedness. Pathway flux rates were used with enzyme V(max) values to estimate fractional velocities (fraction of V(max) at which enzymes operate) for various reactions to gain insights into enzyme function and how this varies with body mass. Fractional velocity is highly conserved across species at the HK step, but varied at all other steps examined. These results are discussed in the context of the regulation and evolution of pathways of energy metabolism.

Euglossine bees (Apidae; Euglossini) exclusively pollinate hundreds of orchid species and comprise up to 25% of bee species richness in neotropical rainforests. As one of the first studies of comparative phylogeography in a neotropical insect group, we performed a mitochondrial DNA (mtDNA)-based analysis of 14 euglossine species represented by populations sampled across the Andes and/or across the Amazon basin. The mtDNA divergences within species were consistently low; across the 12 monophyletic species the mean intraspecific divergence among haplotypes was 0.9% (range of means, 0-1.9%). The cytochrome oxidase 1 (CO1) divergence among populations separated by the Andes (N = 11 species) averaged 1.1% (range 0.0-2.0%). The mtDNA CO1 data set displayed homogeneous rates of nucleotide substitution, permitting us to infer dispersal across the cordillera long after the final Andean uplift based on arthropod molecular clocks of 1.2-1.5% divergence per million years. Gene flow across the 3000-km breadth of the Amazon basin was inferred from identical cross-Amazon haplotypes found in five species. Although mtDNA haplotypes for 12 of the 14 euglossine species were monophyletic, a reticulate CO1 phylogeny was recovered in Euglossa cognata and E. mixta, suggesting large ancestral populations and recent speciation. Reference to closely related outgroups suggested recent speciation for the majority of species. Phylogeographical structure across a broad spatial scale is weaker in euglossine bees than in any neotropical group previously examined, and may derive from a combination of Quaternary speciation, population expansion and/or long-distance gene flow.

Pollination deficits in agricultural and natural systems are suggestive of large reductions in pollinator populations. However, actual declines are difficult to demonstrate using census data. Here, we show census data to be misleading because many abundant pollinators exhibit high levels of production of sterile diploid males usually found only in small inbred hymenopteran populations; Euglossa imperialis exhibits high levels of diploid male production induced by low effective population sizes (Ne approximately 15), despite being the most abundant orchid bee in lowland tropical forests in Panama. We caution that although some pollinators appear abundant on the basis of census data, their long-term persistence may be highly tenuous based on genetic evidence. We propose the use of diploid male frequency data as a metric for assessing the sustainability of bee populations.

The self-pollinating African shrub Coffea arabica, a pillar of tropical agriculture, was considered to gain nothing from insect pollinators. But I show here that naturalized, non-native honeybees can augment pollination and boost crop yields by over 50%. These findings, together with world coffee-harvest statistics and results from field studies of organically shade-grown coffee, indicate that coffee plants would benefit from being grown in habitats that are suitable for sustaining valuable pollinators.

Background: Individuals that band together create new ecological opportunities for microorganisms. In vertical transmission, theory predicts a conserved microbiota within lineages, especially social bees. Bees exhibit solitary to social behavior among and/or within species, while life cycles can be annual or perennial. Bee nests may be used over generations or only once, and foraging ecology varies widely. To assess which traits are associated with bee microbiomes, we analyzed microbial diversity within solitary and social bees of Apidae, Colletidae, and Halictidae, three bee families in Panama's tropical forests. Our analysis considered the microbiome of adult gut contents replicated through time, localities, and seasons (wet and dry) and included bee morphology and comparison to abdominal (dissected) microbiota. Diversity and distribution of tropical bee microbes (TBM) within the corbiculate bee clade were emphasized. Results: We found the eusocial corbiculate bees tended to possess a more conserved gut microbiome, attributable to vertical transmission, but microbial composition varied among closely related species. Euglossine bees (or orchid bees), corbiculates with mainly solitary behavior, had more variable gut microbiomes. Their shorter-tongued and highly seasonal species displayed greater diversity, attributable to flower-visiting habits. Surprisingly, many stingless bees, the oldest corbiculate clade, lacked bacterial genera thought to predate eusociality, while several facultatively social, and solitary bee species possessed those bacterial taxa. Indeed, nearly all bee species displayed a range of affinities for single or multiple variants of the "socially associated" bacterial taxa, which unexpectedly demonstrated high sequence variation. Conclusions: Taken together, these results call into question whether specific bacterial associates facilitate eusocial behavior, or are subsequently adopted, or indicate frequent horizontal transmission between perennial eusocial colonies and other social, facultatively social, and solitary bees. 7ptKwZS8umiRRciGf9GNsX Video Abstract [ABSTRACT FROM AUTHOR]

Stingless bees form perennial colonies of honey-making insects. The >600 species of stingless bees, mainly Neotropical, live throughout tropical latitudes. Foragers influence floral biology, plant reproduction, microbe dispersal, and diverse ecosystem functions. As tropical forest residents since the upper Cretaceous, they have had a long evolutionary history without competition from honey bees. Most stingless bees are smaller than any Apis species and recruit nest mates to resources, while their defense strategies exclude stinging behavior but incorporate biting. Stingless bees have diversified ecologically; excel in nesting site selection and mutualisms with plants, arthropods, and microbes; and display opportunism, including co-opting plant defenses. As their biology becomes better known, applications to human endeavors are imposing selective pressures from exploitation and approaches to conservation that entail colony extraction from wildlands. Although some meliponines can adjust to new conditions, their populations shall require tropical diversity for survival and reproduction. [ABSTRACT FROM AUTHOR]

Understanding the effect of body size on flight costs is critical for the development of models of aerodynamics and animal energetics. Prior scaling studies that have shown that flight costs scale hypometrically have focused primarily on larger (>100 mg) insects and birds, but most flying species are smaller. We studied the flight physiology of 13 stingless bee species over a large range of body sizes (1–115 mg). Metabolic rate during hovering scaled hypermetrically (scaling slope = 2.11). Larger bees had warm thoraxes, while small bees were nearly ecothermic; however, even controlling for body temperature variation, flight metabolic rate scaled hypermetrically across this clade. Despite having a lower mass-specific metabolic rate during flight, smaller bees could carry the same proportional load. Wingbeat frequency did not vary with body size, in contrast to most studies that find wingbeat frequency increases as body size decreases. Smaller stingless bees have a greater relative forewing surface area, which may help them reduce the energy requirements needed to fly. Further, we hypothesize that the relatively larger heads of smaller species may change their body pitch in flight. Synthesizing across all flying insects, we demonstrate that the scaling of flight metabolic rate changes from hypermetric to hypometric at ∼58 mg body mass with hypermetic scaling below (slope = 1.2) and hypometric scaling (slope = 0.67) >58 mg in body mass. The reduced cost of flight likely provides selective advantages for the evolution of small body size in insects. The biphasic scaling of flight metabolic rates and wingbeat frequencies in insects supports the hypothesis that the scaling of metabolic rate is closely related to the power requirements of locomotion and cycle frequencies. [ABSTRACT FROM AUTHOR]

Plants and pollinators are linked but their dynamics are scarcely known. Chemical monitoring of male "orchid bees" at two sites revealed 75% of species were stable or increasing. Forest bees of 33 species, with live sighting at Pipeline Road (PR), and trapping on Barro Colorado Island (BCI), included 132,000 individuals. No species or community changes occurred in strong El Niño‐Southern Oscillation (ENSO) climate events, which lasted 145 total months during more than 70 bee generations. Parasite and host bees fluctuated in unison. A few very common species, adapted to relatively disturbed habitat, diminished over time at PR, while sightings and traps revealed stable abundance ranks in Euglossa 40 years on BCI. Bee abundance and biomass were stable but 50% of species had few records. Orchid bees appear more stable in older forest, they were evidently insulated from ENSO disturbance, and probably benefit from the increasing abundance of flowering lianas and vines. [ABSTRACT FROM AUTHOR]

GLO:69VH/01jul21:bes21905-fig-0002.jpg PHOTO (COLOR): . gl Photo 2: Woodpecker or other predator damage to Cecropia trunks where bee and ant colonies live. The first known bees nesting in a preformed plant cavity (an ant-plant providing cavities with no known bee inhabitants or defensive ants) were found in the western Amazon. Their evolution as I Cecropia i bees coincided with the evolution of ant plants and remained prominent in two species of a tree clade often defended by highly aggressive I Azteca i ants against herbivores and competitors. [Extracted from the article]