Abstract

Geographical patterns in body size have been described across a wide range of species, leading to the development of a series of fundamental biological rules. However, shape variables are less well-described despite having substantial consequences for organism performance. Wing aspect ratio (AR) has been proposed as a key shape parameter that determines function in flying animals, with high AR corresponding to longer, thinner wings that promote high manoeuvrability, low speed flight, and low AR corresponding to shorter, broader wings that promote high efficiency long distance flight. From this principle it might be predicted that populations living in cooler areas would exhibit low AR wings to compensate for reduced muscle efficiency at lower temperatures. I test this hypothesis using the riverine damselfly, Calopteryx maculata, sampled from 34 sites across its range margin in North America. Nine hundred and seven male specimens were captured from across the 34 sites (mean = 26.7 ± 2.9 SE per site), dissected and measured to quantify the area and length of all four wings. Geometric morphometrics were employed to investigate geographical variation in wing shape. The majority of variation in wing shape involved changes in wing aspect ratio, confirmed independently by geometric morphometrics and wing measurements. There was a strong negative relationship between wing aspect ratio and the maximum temperature of the warmest month which varies from west-east in North America, creating a positive relationship with longitude. This pattern suggests that higher aspect ratio may be associated with areas in which greater flight efficiency is required: regions of lower temperatures during the flight season. I discuss my findings in light of research of the functional ecology of wing shape across vertebrate and invertebrate taxa.

Highlights

  • Powered flight has evolved independently in four different lineages: the pterosaurs, insects, birds, and bats, allowing animals to exploit novel niches and avoid predators

  • The literature on the functional relevance of insect wing morphology is heavily biased towards theory (Dudley, 2002), laboratory studies (Betts & Wootton, 1988; Marden, 1995) and observations of kinematics (Ruppell, 1989; Wakeling & Ellington, 1997a; Wakeling & Ellington, 1997b; Wakeling & Ellington, 1997c) rather than quantitative data collected from the field

  • The use of geometric morphometrics to analyse shape confirms that changes in aspect ratio constitute the major source of variation between specimens from different sites

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Summary

Introduction

Powered flight has evolved independently in four different lineages: the pterosaurs, insects, birds, and bats, allowing animals to exploit novel niches and avoid predators. There has been speculation that the benefits of high aspect ratio may be reduced or even reversed at the low Reynolds numbers (a measure of aerodynamic turbulence, with lower numbers corresponding to the viscous forces experienced by small objects) experienced by insects (Ennos, 1989; Wootton, 1992). This speculation, along with the difference in the nature of flight—number, structure and locomotory independence of wings—between birds and insects complicates the formation of hypotheses concerning the implications of variation in flight morphology (Betts & Wootton, 1988; Johansson, Soderquist & Bokma, 2009). The literature on the functional relevance of insect wing morphology is heavily biased towards theory (Dudley, 2002), laboratory studies (Betts & Wootton, 1988; Marden, 1995) and observations of kinematics (Ruppell, 1989; Wakeling & Ellington, 1997a; Wakeling & Ellington, 1997b; Wakeling & Ellington, 1997c) rather than quantitative data collected from the field

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