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      The Allometry of Bee Proboscis Length and Its Uses in Ecology

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          Abstract

          Allometric relationships among morphological traits underlie important patterns in ecology. These relationships are often phylogenetically shared; thus quantifying allometric relationships may allow for estimating difficult-to-measure traits across species. One such trait, proboscis length in bees, is assumed to be important in structuring bee communities and plant-pollinator networks. However, it is difficult to measure and thus rarely included in ecological analyses. We measured intertegular distance (as a measure of body size) and proboscis length (glossa and prementum, both individually and combined) of 786 individual bees of 100 species across 5 of the 7 extant bee families (Hymenoptera: Apoidea: Anthophila). Using linear models and model selection, we determined which parameters provided the best estimate of proboscis length. We then used coefficients to estimate the relationship between intertegular distance and proboscis length, while also considering family. Using allometric equations with an estimation for a scaling coefficient between intertegular distance and proboscis length and coefficients for each family, we explain 91% of the variance in species-level means for bee proboscis length among bee species. However, within species, individual-level intertegular distance was a poor predictor of individual proboscis length. To make our findings easy to use, we created an R package that allows estimation of proboscis length for individual bee species by inputting only family and intertegular distance. The R package also calculates foraging distance and body mass based on previously published equations. Thus by considering both taxonomy and intertegular distance we enable accurate estimation of an ecologically and evolutionarily important trait.

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          Bee foraging ranges and their relationship to body size.

          Bees are the most important pollinator taxon; therefore, understanding the scale at which they forage has important ecological implications and conservation applications. The foraging ranges for most bee species are unknown. Foraging distance information is critical for understanding the scale at which bee populations respond to the landscape, assessing the role of bee pollinators in affecting plant population structure, planning conservation strategies for plants, and designing bee habitat refugia that maintain pollination function for wild and crop plants. We used data from 96 records of 62 bee species to determine whether body size predicts foraging distance. We regressed maximum and typical foraging distances on body size and found highly significant and explanatory nonlinear relationships. We used a second data set to: (1) compare observed reports of foraging distance to the distances predicted by our regression equations and (2) assess the biases inherent to the different techniques that have been used to assess foraging distance. The equations we present can be used to predict foraging distances for many bee species, based on a simple measurement of body size.
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            A global quantitative synthesis of local and landscape effects on wild bee pollinators in agroecosystems.

            Bees provide essential pollination services that are potentially affected both by local farm management and the surrounding landscape. To better understand these different factors, we modelled the relative effects of landscape composition (nesting and floral resources within foraging distances), landscape configuration (patch shape, interpatch connectivity and habitat aggregation) and farm management (organic vs. conventional and local-scale field diversity), and their interactions, on wild bee abundance and richness for 39 crop systems globally. Bee abundance and richness were higher in diversified and organic fields and in landscapes comprising more high-quality habitats; bee richness on conventional fields with low diversity benefited most from high-quality surrounding land cover. Landscape configuration effects were weak. Bee responses varied slightly by biome. Our synthesis reveals that pollinator persistence will depend on both the maintenance of high-quality habitats around farms and on local management practices that may offset impacts of intensive monoculture agriculture. © 2013 Blackwell Publishing Ltd/CNRS.
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              The development and evolution of exaggerated morphologies in insects.

              We discuss a framework for studying the evolution of morphology in insects, based on the concepts of "phenotypic plasticity" and "reaction norms." We illustrate this approach with the evolution of some of the most extreme morphologies in insects: exaggerated, sexually selected male ornaments and weapons, and elaborate social insect soldier castes. Most of these traits scale with body size, and these scaling relationships are often nonlinear. We argue that scaling relationships are best viewed as reaction norms, and that the evolution of exaggerated morphological traits results from genetic changes in the slope and/or shape of these scaling relationships. After reviewing literature on sexually selected and caste-specific structures, we suggest two possible routes to the evolution of exaggerated trait dimensions: (a) the evolution of steeper scaling relationship slopes and (b) the evolution of sigmoid or discontinuous scaling relationship shapes. We discuss evolutionary implications of these two routes to exaggeration and suggest why so many of the most exaggerated insect structures scale nonlinearly with body size. Finally, we review literature on insect development to provide a comprehensive picture of how scaling relationships arise and to suggest how they may be modified through evolution.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                17 March 2016
                2016
                : 11
                : 3
                : e0151482
                Affiliations
                [1 ]Department of Entomology, University of Minnesota, Saint Paul, Minnesota, United States of America
                [2 ]Department of Ecology, Evolution, and Natural Resources, Rutgers University, New Brunswick, New Jersey, United States of America
                [3 ]Departamento Ecología Integrativa, Estación Biológica de Doñana (EDB-CSIC), Avda. Américo Vespucio s/n, Isla de la Cartuja, 41092, Sevilla, Spain
                [4 ]Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore, 117543, Singapore
                [5 ]Department of Entomology, Michigan State University, East Lansing, Michigan, United States of America
                University of North Carolina, Greensboro, UNITED STATES
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: RW. Performed the experiments: GKN JZ. Analyzed the data: DPC IB. Wrote the paper: DPC GKN IB JZ JSA JG RW. Identified the bee specimens: JSA JG.

                Article
                PONE-D-15-35132
                10.1371/journal.pone.0151482
                4795761
                26986000
                a6c53afe-0aef-49e6-b522-d88d6350905b
                © 2016 Cariveau et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 10 August 2015
                : 29 February 2016
                Page count
                Figures: 3, Tables: 2, Pages: 13
                Funding
                This work was supported by the United States Department of Agriculture, Natural Resources Conservation Service Conservation Innovation Grant, NRCS Agreement #69-3A75-10-163 (to RW) (URL: nrcs.usda.gov/wps/portal/nrcs/main/national/programs/financial/cig/); a Fulbright-Nehru Postdoctoral Fellowship (to GN) (URL: cies.org/grantee/geetha-nayak); and a Rutgers University Aresty Undergraduate Research Award (to JZ) (URL: https://aresty.rutgers.edu/our-programs/funding/undergraduate-research-fellowships). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Organisms
                Animals
                Invertebrates
                Arthropoda
                Insects
                Hymenoptera
                Bees
                Biology and Life Sciences
                Physiology
                Physiological Parameters
                Medicine and Health Sciences
                Physiology
                Physiological Parameters
                Biology and Life Sciences
                Ecology
                Community Ecology
                Ecology and Environmental Sciences
                Ecology
                Community Ecology
                Biology and Life Sciences
                Ecology
                Plant Ecology
                Ecology and Environmental Sciences
                Ecology
                Plant Ecology
                Biology and Life Sciences
                Plant Science
                Plant Ecology
                Biology and Life Sciences
                Organisms
                Animals
                Invertebrates
                Arthropoda
                Insects
                Moths and Butterflies
                Biology and Life Sciences
                Evolutionary Biology
                Evolutionary Systematics
                Phylogenetics
                Biology and Life Sciences
                Taxonomy
                Evolutionary Systematics
                Phylogenetics
                Computer and Information Sciences
                Data Management
                Taxonomy
                Evolutionary Systematics
                Phylogenetics
                Biology and Life Sciences
                Behavior
                Animal Behavior
                Foraging
                Biology and Life Sciences
                Zoology
                Animal Behavior
                Foraging
                Biology and Life Sciences
                Plant Science
                Plant Anatomy
                Flowers
                Custom metadata
                The data are found in two places. First, they are part of the supplemental material. Second, they are included in the R package that was created for this manuscript. The R package can be downloaded at: https://github.com/ibartomeus/BeeIT.

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