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      Pheromone production, male abundance, body size, and the evolution of elaborate antennae in moths

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          Abstract

          The males of some species of moths possess elaborate feathery antennae. It is widely assumed that these striking morphological features have evolved through selection for males with greater sensitivity to the female sex pheromone, which is typically released in minute quantities. Accordingly, females of species in which males have elaborate (i.e., pectinate, bipectinate, or quadripectinate) antennae should produce the smallest quantities of pheromone. Alternatively, antennal morphology may be associated with the chemical properties of the pheromone components, with elaborate antennae being associated with pheromones that diffuse more quickly (i.e., have lower molecular weights). Finally, antennal morphology may reflect population structure, with low population abundance selecting for higher sensitivity and hence more elaborate antennae. We conducted a phylogenetic comparative analysis to test these explanations using pheromone chemical data and trapping data for 152 moth species. Elaborate antennae are associated with larger body size (longer forewing length), which suggests a biological cost that smaller moth species cannot bear. Body size is also positively correlated with pheromone titre and negatively correlated with population abundance (estimated by male abundance). Removing the effects of body size revealed no association between the shape of antennae and either pheromone titre, male abundance, or mean molecular weight of the pheromone components. However, among species with elaborate antennae, longer antennae were typically associated with lower male abundances and pheromone compounds with lower molecular weight, suggesting that male distribution and a more rapidly diffusing female sex pheromone may influence the size but not the general shape of male antennae.

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          Relationships between body size and abundance in ecology.

          Body size is perhaps the most fundamental property of an organism and is related to many biological traits, including abundance. The relationship between abundance and body size has been extensively studied in an attempt to quantify the form of the relationship and to understand the processes that generate it. However, progress has been impeded by the under appreciated fact that there are four distinct, but interrelated, relationships between size and abundance that are often confused in the literature. Here, we review and distinguish between these four patterns, and discuss the linkages between them. We argue that a synthetic understanding of size-abundance relationships will result from more detailed analyses of individual patterns and from careful consideration of how and why the patterns are related.
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            The role of chemical communication in mate choice.

            Chemical signals are omnipresent in sexual communication in the vast majority of living organisms. The traditional paradigm was that their main purpose in sexual behaviour was to coordinate mate and species recognition and thus pheromones were conserved in structure and function. In recent years, this view has been challenged by theoretical analyses on the evolution of pheromones and empirical reports of mate choice based on chemical signals. The ability to measure precisely the quantity and quality of chemicals emitted by single individuals has also revealed considerable individual variation in chemical composition and release rates, and there is mounting evidence that prospecting mates respond to this variation. Here, we review the evidence for pheromones as indicators of mate quality and examine the extent of their use in individual mate assessment. We begin by briefly defining the levels of mate choice--species recognition, mate recognition and mate assessment. We then explore the degree to which pheromones satisfy the key criteria necessary for their evolution and maintenance as cues in mate assessment; that is, they should exhibit variation across individuals within a sex and species; they should honestly reflect an individual's quality and thus be costly to produce and/or maintain; they should display relatively high levels of heritability. There is now substantial empirical evidence that pheromones can satisfy all these criteria and, while measurements of the actual metabolic cost of pheromone production remain to some degree lacking, trade-offs between pheromone production and various fitness-related characters such as growth rate, immunocompetence and longevity have been reported for a range of species. In the penultimate section, we outline the growing number of studies where the consequences of chemical-based mate assessment have been investigated, specifically focussing on the reported direct and genetic benefits accrued by the receiver. Finally, we highlight potential areas for future research and in particular emphasise the need for interdisciplinary research that combines exploration of chemical, physiological and behavioural processes to further our understanding of the role of chemical cues in mate assessment.
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              The seven deadly sins of comparative analysis.

              Phylogenetic comparative methods are extremely commonly used in evolutionary biology. In this paper, I highlight some of the problems that are frequently encountered in comparative analyses and review how they can be fixed. In broad terms, the problems boil down to a lack of appreciation of the underlying assumptions of comparative methods, as well as problems with implementing methods in a manner akin to more familiar statistical approaches. I highlight that the advent of more flexible computing environments should improve matters and allow researchers greater scope to explore methods and data.
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                Author and article information

                Journal
                Ecol Evol
                Ecol Evol
                ece3
                Ecology and Evolution
                Blackwell Publishing Ltd (Oxford, UK )
                2045-7758
                2045-7758
                January 2012
                : 2
                : 1
                : 227-246
                Affiliations
                [1 ]simpleDepartment of Zoology, University of Melbourne Victoria 3010, Australia
                [2 ]simpleCentre for Integrative Ecology, School of Life and Environmental Sciences, Deakin University Burwood, Victoria 3125, Australia.
                Author notes
                Matthew R. E. Symonds, School of Life and Environmental Sciences, Deakin University, 221 Burwood Highway, Burwood, Victoria 3125, Australia. Tel: +61 3 9251 7437; Fax: +61 3 9251 7626; E-mail: matthew.symonds@ 123456deakin.edu.au

                Funded by a Discovery Project grant from the Australian Research Council (DP0987360).

                Article
                10.1002/ece3.81
                3297191
                22408739
                0b184798-6819-44de-872e-c612527a44fb
                © 2011 The Authors. Published by Blackwell Publishing Ltd.

                This is an open access article under the terms of the Creative Commons Attribution Non Commercial License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes.

                History
                : 13 September 2011
                : 23 October 2011
                : 26 October 2011
                Categories
                Original Research

                Evolutionary Biology
                phylogenetic generalized least squares,forewing length,lepidoptera,antennal morphology,sex pheromone

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