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For exceptions, permission may be sought for such use through Elsevier’s permissions site at: http://www.elsevier.com/locate/permissionusematerial From Suhling, F., Sahlén, G., Gorb, S., Kalkman, V.J., Dijkstra, K-D.B., van Tol, J., 2015. Order Odonata. In: Thorp, J., Rogers, D.C. (Eds.), Ecology and General Biology: Thorp and Covich's Freshwater Invertebrates, Academic Press, 893–932. ISBN: 9780123850263 Copyright © 2015 Elsevier Inc. All rights reserved. Academic Press Author's personal copy Chapter 35 Order Odonata Frank Suhling Institut für Geoökologie, Technische Universität Braunschweig, Braunschweig, Germany Göran Sahlén Ecology and Environmental Sciences, Halmstad University, Halmstad, Sweden Stanislav Gorb Spezielle Zoologie, Universität Kiel, Kiel, Germany Vincent J. Kalkman, Klaas-Douwe B. Dijkstra and Jan van Tol Naturalis Biodiversity Center, Leiden, The Netherlands Chapter Outline Introduction 894 Reproduction 911 Systematic and Phylogenetic Relationships 894 Sexual Dimorphism 911 Zygoptera 894 Mating Systems 911 Anisozygoptera 896 The Mating Process 911 Anisoptera 896 Oviposition 911 Species Numbers 897 Life Cycle 913 General Biology 899 Egg Development 913 External Features of the Larva 899 Larval Development 913 External Features of the Imago 900 Metamorphosis and Emergence 914 Size 902 Seasonal Patterns 915 Egg Structure 902 Prereproductive Period 915 Ultrastructures 903 Adult Life Span 915 Wing Structures 903 Life Cycle Types and Voltinism 916 Coloration 904 General Ecology and Behavior 917 Cuticula 904 Foraging 917 Head Arrester System 904 Larval Foraging 917 Genitalia 905 Adult Foraging Behavior 918 Perception: The Sensory Organs and Dispersal and Migration 918 Neural System 905 Habitats 918 Compound Eyes and Ocelli 906 Generalists versus Specialists 918 Tactile Sensory Organs 907 Habitat Selection 919 Respiration 908 Microhabitat Occupancy by Larvae 919 Larval Gill Systems 908 Lotic Waters 921 Oxygen Demands 909 Temporary Habitats: Coping with or Tracheal System of the Imago 909 Avoiding Drought 921 Physical-Gill Respiration 909 Acidic Ponds and Lakes 922 Thermoregulation 910 Saline Waters 922 Flight 910 Forest and Shade Habitats 922 Thorp and Covich’s Freshwater Invertebrates. http://dx.doi.org/10.1016/B978-0-12-385026-3.00035-8 Copyright © 2015 Elsevier Inc. All rights reserved. 893 Ecology and General Biology, Fourth Edition, 2015, 893-932 Author's personal copy 894 SECTION | VI Phylum Arthropoda Very Small Habitats 922 Range Shifts due to Climate Change 926 Terrestrial Habitats 922 Conservation Status and Biotic Indicators 927 Biotic Interactions 922 Collecting, Culturing, and Specimen Preparation 928 Predation 923 Collecting and Sampling 928 Intraspecific Interactions 924 Culturing 929 Abiotic Limitations and Biotic Interactions 924 Preservation 929 Parasitism and Other Interactions 925 Acknowledgments 930 Distribution and Diversity 925 References 930 Diversity Patterns 925 INTRODUCTION including synonyms and references­ is available on www. odonata.info. Systematic and Phylogenetic Relationships Zygoptera Dragonflies belong to the superorder Odonatoptera, one of the oldest insect radiations to take flight, dating back at The adults (imagines) have a broad head with widely least to the early Carboniferous. This radiation includes the ­separated eyes and a slender abdomen. The fore and hind largest insect that ever lived, the griffenfly Meganeuropsis wings are similar in shape, and most species rest with their permiana Carpenter, 1939, with a wingspan of c. 70 cm. wings closed. The larvae have three (sometimes two) cau- The radiation led to the rise of the order Odonata, with the dal gills for respiration, which can also be used as flippers oldest fossils dating back to the Permian. The present-day for swimming. It is generally agreed that the suborder is Odonata is regarded as a monophyletic group, which is ­monophyletic, and that the superfamily Lestoidea consti- divided into three suborders: Zygoptera or damselflies and tutes the sister group of the remaining 93% of the damsel- Anisoptera or true dragonflies—each with approximately fly species. The monotypic family Hemiphlebiidae from 3000 species—and a small suborder Anisozygoptera (dam- southeastern Australia and Tasmania is the sister group of sel dragons) with four species. Unique features of odonates the three other lestoid families. Lestidae comprises 72% are the strongly modified larval labium and the mechanism of the lestoids and, while monophyletic, its main genus of indirect sperm transfer (both described later), which Lestes Leach, 1815, is very heterogeneous and likely to requires a male copulatory organ at the second abdominal be split in the future. While the 150 lestid species occupy segment. Sperm must be transferred externally to this organ mainly open, stagnant,­ and often temporary waters world- before copulation. During the insemination, the male holds wide, the less than 60 other lestoids are localized in mostly the female with his abdominal appendages, behind the head mountain or forest streams. Synlestidae is found in eastern (Anisoptera) or on the prothorax (Zygoptera), while the ­Australia, southern and central Africa, and the tropics of female bends the tip of her abdomen toward the secondary mainland Southeast Asia, while Perilestidae is restricted to genitalia of the male, forming the well-known mating wheel the ­Neotropics. of dragonflies. Platystictoidea with its sole family Platystictidae is the In the past, wing venation was used as the main guide sister group of all Zygoptera excluding Lestoidea. The group for classifying Odonata; but as similar characters evolved is confined to the wettest tropics, being notably absent from multiple times, this is often not a reliable indication of Africa and Madagascar. While it is also present in Central close relationships. Studies incorporating other morpho- and northern South America, 80% of the species occur from logical features, including those of larvae, have helped to Sri Lanka and India to New Guinea. Currently the over 220 overcome this (von Ellenrieder, 2002; Rehn, 2003; Fleck species are placed in only eight genera, a higher ratio than et al., 2008a), as have molecular studies (Bybee et al., in any other family. This is due to the highly polyphyletic 2008; Carle et al., 2008; Fleck et al., 2008b; Dumont nature of the two main genera, Protosticta Selys, 1885, and et al., 2010). Molecular studies have in particular resulted Drepanosticta Laidlaw, 1917, which will undoubtedly be in major changes in odonate taxonomy in recent years split up in the near future. ­(Dijkstra and Kalkman, 2012). For Anisoptera we follow About 58% of all damselfly species belong to the here the taxonomy proposed by Dijkstra et al. (2013); superfamily Coenagrionoidea, which includes three while for Zygoptera the taxonomy of Dijkstra et al. families. Isostictidae is the smallest, containing less than (2014) is used. The inferred phylogeny of extant Odonata 2% of all damselfly species, all of which are confined to is shown in Figure 35.1. A checklist of all dragonflies Australia, New Guinea, New Caledonia, and neighboring Ecology and General Biology, Fourth Edition, 2015, 893-932 Author's personal copy Chapter | 35 Order Odonata 895 FIGURE 35.1 Inferred phylogeny of families of: (a) Zygoptera and (b) Anisozygoptera (Epiophlebiidae) and Anisoptera. Line thickness is indicative of species diversity. The relationships of most families within Zygoptera are unresolved, and this polytomy is here depicted as a paraphyletic assemblage. The numerous monogeneric families within this polytomy are listed to the left. Genera incertae sedis not shown. Figure by VK and KD. islands. Platycnemididae is restricted to the Old World The superfamily name Calopterygoidea has been used and contains over 400 species, most of which are confined for a very diverse assemblage of mostly densely veined to streams and rivers. One of its subfamilies (Disparo- damselflies constituting 27% of the total damselfly diver- neurinae) comprises all Old World species of the former sity. While the other superfamilies are quite well defined, family Protoneuridae. Coenagrionidae is the largest dam- the remaining families are grouped together without much selfly family with over 1200 species. The family includes evidence for their monophyly. Calopterygoidea may ulti- two large lineages that are referred to as the core and the mately prove polyphyletic, requiring the creation of fur- ridge-faced Coenagrionidae, each with about 600 spe- ther superfamilies. With almost a quarter of the species, cies. Although both groups are found worldwide, a rela- Calopterygidae is the largest and most widespread family tively high proportion of