Aphid predators
By Graham E. Rotheray and J. C. Rotheray
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About this ebook
Aphids and their colonies are excellent arenas in which to observe predators in action. A range of insects come to eat or parasitise the aphids or to drink their honeydew.
9781784275105 and 9781784275167 are digital reprints of 9780855462697 (1989).
Graham E. Rotheray
As a child Graham Rotheray was intrigued by flies buzzing around his parents’ vegetable garden, especially the hoverflies and their mimicking of wasps in order to evade birds. For his PhD he devised both laboratory and field experiments to investigate parasitic insects that attacked the larval stages of hoverflies. In 1980 he applied his expertise to a study of parasites of a major pest species in the eastern United States, the introduced gypsy moth, in an attempt to devise a biological means of controlling their spread. On returning to Britain, Graham was appointed Curator of Insects at the National Museums of Scotland in Edinburgh. He was charged with developing the collections of Diptera. He developed an interest in ancient Scottish woodlands and the hoverflies that occur there, such as the rare Callicera rufa.
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Aphid predators - Graham E. Rotheray
Editors’ preface
Students at school or university, and others without a university training in biology, may have the opportunity and inclination to study local natural history but lack the knowledge to do so in a confident and productive way. The books in this series offer them the information and ideas needed to plan an investigation, and the practical guidance to carrry it out. They draw attention to regions on the frontiers of current knowledge where amateur studies have much to offer. We hope readers will derive as much satisfaction from their biological explorations as we have done.
An aphid colony is the scene of a lively drama, that runs day and night. A range of insects come to eat or parasitise the aphids or to drink their honeydew. Other Naturalists’ Handbooks deal with the natural history of two major groups of aphid predators, hoverflies and ladybirds. This book complements them by focussing on the behavioural interactions among these and other visitors to a colony of aphids, interactions that offer limitless opportunities for observation and experimental investigation.
S.A.C.
R.H.L.D.
January 1987
Acknowledgements
I am very grateful to the editors, particularly S. A. Corbet, for their patience, help and encouragement during preparation of this book, and for their invaluable suggestions on an earlier draft. My wife, Joanne, gave particular help and gave assistance with the colour plates. Dr. Francis Gilbert and Mrs. Kathleen Davidson gave many helpful comments. I thank my typists, Mrs. Anne Rotheray and Mrs. Kathleen Davidson, for their patience and expertise.
1 Introduction
A predator is an animal which kills and eats other animals. Predation is important in nature because it is one of the three basic ways in which energy passes between living organisms, the other two being herbivory (feeding on plants) and parasitism (feeding on, but not usually killing, the host). Predation is a common process; almost all animals have predators. For many, predation is a constant threat and life can be seen as a succession of attempts to escape attacks. Through the long interplay between predators and their prey a great diversity of prey-capture and anti-predator devices have evolved.
Part of the aim in studying predation is to understand what these devices are and how they function. Are they effective? How do predators capture and eat prey and why do they do it in the way they do? Can prey animals co-exist with their predators? If so, how do they avoid being eaten? What impact do predators have on the size and distribution of prey populations? Can predators regulate prey numbers? Is there anything in common between various predators, such as lions, sharks and ladybirds? Because aphids are easy to find and slow to move aphid colonies offer an excellent arena in which to observe predators in action.
Apart from these basic questions there is considerable interest in using predators as a natural means of controlling pests. This is biological control. Unlike chemical sprays, predators do not contaminate the environment, and if sufficiently effective, they can offer a better long term solution to most pest control problems. One of the earliest successes came in 1889 with the introduction to California of an Australian predator, the ladybird Rodolia cardinalis, which reduced the scale insect, Icerya purchasi, from a major pest on citrus trees to one of minor importance. Since then, predators have been used widely in biological control projects and many are being evaluated for use in the future. Biological evaluation is necessary because only by understanding the behaviour and ecology of potentially useful predators can the most effective species be chosen.
Aphid predators have often paved the way forward in research on predation. They can be reared and handled with ease, are common and can be observed in the field. They offer a rich variety of study topics and they behave well under laboratory conditions. Much is still unknown and puzzling about them. In this book you will find enough information to plan and carry out your own investigations on aphid predators. The book explains how predators and aphids can be found and identified. It also explains essential concepts involved in predator-prey behaviour and ecology. Finally, it provides an introduction to the literature on predation against which your own investigations should be planned.
Finding aphid colonies
Aphids, sometimes called greenfly or blackfly, belong to the insect order Hemiptera, the true bugs, which includes cicadas, froghoppers and scale insects. Hemipterans may be known by their mouthparts, which are modified to form a tube used for piercing and sucking (fig. 1). These range from the beak-like rostrum with which bed bugs suck blood to the long thread-like stylets of aphids. Aphids feed on plant sap, tapping the phloem tubes by inserting their mouthparts deep into a plant. Most aphids spend nearly all their time feeding and so are relatively immobile. Often many individuals feed close together forming densely packed aggregations or colonies. This habit makes them easier to find. Colonies (pl. 2) form underneath leaves, on stems, tree trunks, flowers or roots, and are particularly common on the growing tips of plants. Some species are hidden within leaf curls or galls. The aphid faunas of some common plants are described in chapter 4. If the foliage of these plants is carefully examined at the right time of year, aphid colonies should soon be found.
Fig. 1. Adult rose aphid Macrosiphum rosae.
Insects associated with aphid colonies
Many species of insect visit aphid colonies for a variety of purposes. A large group come to feed on the honeydew. Honeydew is a clear sticky fluid periodically ejected as droplets from the anus of a feeding aphid. It consists of excess sugars from the plant sap imbibed by the aphid and some waste products from its digestive system. Honeydew production can be observed easily by watching a feeding aphid for a few minutes. Often honeydew falls on leaves below the colony giving the leaves a wet, shiny appearance. When searching for aphids, this is a useful sign that they are close by. Possibly attracted by honeydew odour, many flies, bees, ants and wasps visit aphid colonies to feed on this sugary liquid. Throughout the day and night insects constantly arrive, feed and depart. Aphid colonies are probably just as attractive as flowers to many insects. Sooty moulds and fungi tend to grow in any uneaten honeydew, giving leaf surfaces a black, powdery look. This is another clue that aphids are close by.
Other visitors are aphid parasitoids. A parasitoid is an insect whose larva develops parasitically on or inside the body of its host insect. Unlike a true parasite, such as a liver fluke, a parasitoid normally consumes and kills its host. The adult parasitoids of aphids are black, wasp-like insects about the size of their hosts. Often the females are seen running excitedly around aphids tapping them with their antennae and probing them with the ovipositor or egg- laying tube. The parasitoids become still only when the ovipositor is pushed inside the aphid to deposit an egg. In the later stages of development, parasitised aphids can be recognised by their rounded, straw-coloured or dark appearance. After spending its larval and pupal period inside the aphid, the new parasitoid emerges by biting its way out leaving a characteristic hole in the dried husk of the aphid.
Sometimes a regular association will occur between an aphid colony and a nearby nest of ants. A worker ant will follow a trail from the nest to the colony, where she will shepherd the aphids, harvest their honeydew and return to the nest with her abdomen swollen with her load. Sometimes ants protect colonies by driving away predators and parasitoids and by carrying aphids to a new plant if the old one begins to wilt or die. There are circumstances, however, in which ants will give up their protective role and will eat aphids. Further details of ant - aphid relationships are discussed in chapter 2.
One of the largest groups of insect visitors tied to aphids are the predators. Aphid predators are of two types. Firstly there are the obligatory or aphid-specific predators, which depend on aphids for food. Included in this group are ladybirds, many hoverflies, aphid midges, lacewings, flower bugs and some solitary wasps and other flies. In many of these groups it is only the larval stages which eat aphids but in some, adults are predators too (see chapter 2). The second group are facultative or polyphagous (feeding on many different types of food) predators in which aphids are just part of a much wider diet. This group includes ground beetles, social wasps, birds, earwigs, and various groups of predatory flies. This book deals with examples from both groups.
Finding aphid predators
There are several methods by which aphid predators can be obtained for detailed study. When an aphid colony is found, carefully examine that part of the plant containing the aphids. Search next to and among the aphids. Avoid breathing on the aphids as they may fall off. Many predators are cryptically coloured (camouflaged) and difficult to spot, especially when they are young and small. Plate 1 shows the general appearance of