Essential Entomology

ESSENTIAL ENTOMOLOGY
ESSENTIAL
ENTOMOLOGY
SECOND EDITION
George C. McGavin
Honorary Research Associate of the Oxford University
Museum of Natural History and Honorary Principal Research Fellow,
Imperial College
Leonidas-Romanos Davranoglou
Leverhulme Trust Early Career Fellow,
Oxford University Museum of Natural History
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
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Preface
phyla Life began in the seas, and salt water is still home to most of the earth’s phyla.
plural of phylum, the major On land, however, two groups dominate: the plants and the insects. The word
groupings of the animal insect comes from the Latin, insectus, the past participle of insecare, to cut or
kingdom, composed of
superclasses and classes, that incise, and refers to the major divisions of an insect’s body.
is, Mollusca, Arthropoda, Insects have been around for well over 400 million years. They were the
Chordata, etc.
first animals on land and the first to get into the air. They are the most species-
rich and abundant group of multicellular animals the Earth has ever seen.
More than half of all described species and about 75% of all animals are in-
sects. But exactly how many species share the planet Earth with us? The truth
is: we don’t have the foggiest idea. Some early guesses of 30 million or even
100 million have been replaced in the last decade with more realistic esti-
mates of somewhere between 5 and 10 million species. Despite this massive
uncertainty, one thing is indisputable: the majority of Earth’s inhabitants are
insects.
In terrestrial ecosystems, green plants are the primary producers, trap-
ping the sun’s energy and using it to convert carbon dioxide to organic
matter. Without photosynthesis the planet would be a very different place.
Autotrophs like plants form the base of terrestrial and freshwater food chains,
Trophic level but every trophic level above is dominated by insects.
the levels of food or energy For example, it may come as a surprise to many that all the heaving,
transfer in food chains/webs. snorting herds of grazing ungulates on the African savannahs are entirely
Food producers or autotrophs
form the first level, herbivores ‘out-munched’ (perhaps by a factor of ten to one) by myriads of tiny
form the second level, primary mandibles. What about the meat eaters? Again, insects consume many times
carnivores the third level.
Other carnivores form the
more animal flesh than all the vertebrate carnivores put together and ants
fourth and fifth levels. alone are the major carnivorous species in any habitat you could mention.
If this sounds implausible, consider that, although insects are individually
small, there are an awful lot of them.
Insects pollinate the vast majority of the world’s quarter of a million or so
species of flowering plants. Pollination is one of the most essential symbioses
ever to have evolved. This plant–insect version of ‘I’ll scratch your back if
you scratch mine’ has lasted for 100 million years and it has generated a rich
diversity of species. Twenty thousand species of bee are, to a large extent, re-
sponsible for the continued survival of the angiosperms, which includes a
very long list of the things we eat: fruit and vegetables from pumpkins, plums,
and peas to cherries, cucumbers, and cocoa.
Without the protein provided by insects many groups of higher animals
would simply not exist. Insects feature exclusively or very largely in the diets
of a whole beastly bestiary from the aardvark to the zorilla (an African striped
polecat). Birds, the descendants of the dinosaurs, are mostly insectivorous.
A brood of nine great tit chicks will consume around 120,000 caterpillars
while they are in the nest. A single swallow chick may consume upwards
of 20,000 bugs, flies, and beetles before it fledges, and even species such as
vi PREFACE
Insects 56.3%
Arachnids 4.5%
Crustaceans 2.4%
Other Arthropods 1.2%
Molluscs 4.2%
Nematodes 0.9%
Other Invertebrates 4.0%
Vertebrates 2.7%
Plants 14.3%
Fungi 4.2%
Algae 2.4%
Vertical bar chart showing Protozoans 2.4%
the approximate proportions Bacteria and Viruses
of all species on Earth. 0.5%
The image shown here is a

composite of photographs of
individual insects attracted
to a sea holly flower over the
course of 15 minutes. It is
obvious that without insects,
pollination would be
© Ed Phillips
impossible for most plants,

and the world’s terrestrial
ecosystems would collapse.
PREFACE vii
buntings, which are seed feeders as adults, rear their young on a nutritious
diet of insects. The earliest primates were insectivores, and many primates still
are today. In countries where insects are large or very abundant, they form
an important source of protein, vitamins, and minerals for humans. Locusts
and grasshoppers, beetle grubs, caterpillars, and termites feature regularly in
tropical diets and their nutritional value exceeds that of western-style fast
foods.
At least a quarter of all insect species are parasites or predators of other
species of insect. Insects recycle nutrients, enrich soils, and dispose of car-
casses and dung. Apart from these essential ecosystem services, insects
provide humans with useful products such as silk, honey, waxes, medicines,
and dyes. We use them to control pest organisms and as model systems to help
us study many aspects of biology from genetics and behaviour to physiology
and ecology.
Of course, insects have a dark side. Herbivorous insects may eat up to 15–
20% of all crops grown for human consumption, and locally, the percentage
may be much higher. Although, contrast this with the 30% of all food grown
that is wasted by humans. In addition, insects carry the viruses, bacteria,
spirochaetes, rickettsiae, protozoans, roundworms, and fungi that are respon-
sible for innumerable plant, animal, and human diseases. About one in six
human beings alive today is affected by an insect-borne illness of some kind.
Besides stings and allergies, which can be fatal for some, the list of diseases
carried by insects includes plague, typhus, sleeping sickness, river blindness,
Chagas’ disease, yellow fever, Zika virus, epidemic typhus, trench fever, loia-
sis, filariasis, and leishmaniasis. In the early twentieth century, Sir Ronald
Ross showed conclusively that mosquitoes were responsible for transmitting
malaria. Although much more is known about malaria today, it remains a
deadly disease responsible for hundreds of thousands of deaths every year.
To complete the destructive side of their activities, insects cause great dam-
age to wooden structures and a wide range of natural materials and fabrics.
There are even, much to the aggravation of museum curators, beetles whose
larvae have a voracious appetite for collections of dried insects.
Being much smaller, insects may not have the immediate appeal of the
‘furries or featheries’, but if you look closer, you will find that they are a lot
more varied and interesting. What animals other than humans build air-
conditioned condominiums, use an underwater breathing apparatus, con-
struct acoustic equipment, cause explosions, make paper, cultivate gardens,
and farm other animals? The world of insects is endlessly fascinating, and
they have extraordinary relationships with other organisms. Some of their
lifecycles beggar belief. There are moths that suck mammalian blood, flies
that breed in pools of crude petroleum, flies whose larvae develop only in
the tracheal passages of red kangaroos, and lice that solely inhabit the throat
pouches of certain cormorants and pelicans.
Despite all the talk of the need to quantify biodiversity in recent years,
taxonomy and systematics are still less fashionable today than they once were
and there is a lack of funding for basic taxonomic research. How are we going
to save biodiversity, if we do not know what we are trying to save in the first
viii PREFACE
place? Perhaps we have lost sight of the fact that, without a system of classifi-
cation, our investigations can only be a confusion of unconnected facts. We
survive in a complex world by defining what things are, naming them, and
thus identifying them. Imagine the colossal task of having to remember each
and every novel object individually by its unique characteristics. Classifica-
tion is a central process of the human brain, and it, not mathematics, as has
sometimes been suggested, is the foundation and cornerstone of all science.
As British biologist and beetle taxonomist Roy Crowson pointed out, you
need to know what you are going to count or measure before your numbers
are going to make any sense.
Sex, violence, and a cast of trillions—the study of insects is exciting and
intellectually satisfying, but where do you start? With classification of course.
In any study, whether ecological, physiological, or behavioural, the first thing
you need to know is what sort of insect you’re dealing with. Textbooks some-
times emphasize systems biology at the expense of systematics with the result
that students are familiar with the broader picture of insect physiology or
behaviour but are confused when it comes to recognizing a specific insect.
Whether there are 2 or 10 million insect species, you do not need to know
them all individually. At first glance, a beetle may resemble an ant, and some
flies are excellent mimics of bees, but ants do not have wing cases and bees
have two pairs of wings. Within each order there are conspicuous features
that we use to aid correct identification.
The amount of biological information available, especially concerning in-
sects, is increasing rapidly and newcomers to the study of insects can be
put off simply by the enormity of the subject. Some species have received
a huge amount of attention for various reasons. The American Cockroach,
Periplaneta americana, and locust species, such as Schistocerca gregaria, are
enduringly popular as models for research, and fruit flies (Drosophila spp.)
are extraordinarily useful in genetics. Particular researchers have had their
favourites. Wigglesworth’s pioneering work on insect cuticle used a South
American, cone-nosed, reduviid bug, Rhodnius prolixus, and Dethier’s choice
of a particular blow fly, Phormia regina, for his work on insect feeding and
sensory biology resulted from a lunchtime chance observation of a female of
this species laying her eggs on a liverwurst sandwich. These, and a handful of
other species, are rare exceptions. The rest have yet to enjoy their 15 minutes
of fame. The majority of the world’s living insects are completely unknown
and the biology of many of those we know about are obscure. There is still
plenty of work to be done.
But there is a problem. Human activities have brought about a biodiver-
sity crisis. The conversion of more and more of the planet’s natural capital
into producing human beings and serving their needs and desires has come
at an enormous cost. We seem to have developed a serious disconnect with
the natural world and this disconnect may be our downfall. Populations of
major groups of organisms, for example, birds, reptiles, and mammals, have
declined by an average of 68% in the last 50 years. Almost every recent
study undertaken points to a marked slump in insect species’ richness and
abundance. The decreases seen in well-studied insect groups like bees and
PREFACE ix
butterflies are occurring in many other groups. The causes of these declines
are very clear: the accelerating loss of natural habitat, mostly due to agricul-
tural intensification, coupled with the prodigious amounts of pesticides used,
are taking a heavy toll. On top of this we are facing global climate change
caused by our use of fossil fuels, which affects every species and every habitat
on Earth.
It has long been thought that the world’s tropical forests are home to more
than half of all extant species. If these complex habitats continue to be felled
for timber, cash crops, and ranching at even the slowest rate suggested, it will
still only be a matter of a few hundred years before they are completely lost.
It is therefore an inescapable conclusion that our planet could lose more than
half of all its living species in the time it takes for an acorn to become a veteran
oak tree. We know that no species lasts forever. Of all species that have ever
lived, 99% are now extinct and it was the numerous mass extinction events
that paved the way for the appearance of those creatures that led to human be-
ings. The difference is that now we know enough to do something that might
prolong our own survival.
There can be little doubt that humans are the most intelligent and capable
species yet to evolve on Earth. In the very short time since our appearance,
we covered the entire globe and established colonies wherever it was possible
to survive. A few of us have walked on the surface of the moon and visited
the deepest abysses of the oceans. We spend vast sums of money to probe the
very make-up of matter and examine the universe. We want to understand
the science of everything from the infinitesimally small to the astronomically
large but seem to be paying little heed to the health of the very ecosystems
that support us.
The aim of this book is to provide a readable introduction to the most
abundant multicellular life forms on Earth. It is not a field guide but, should
you need to know the essential facts about a particular order of insect (e.g.
where they occur, how many species are known, what makes them different
and interesting from all other orders), this book will provide them and direct
you to specialist texts and sources of information to guide you further along
any particular path. If you really want to understand how the natural world
works, you need to know about insects.
The book is laid out in an uncomplicated fashion. The first section gives
a brief introduction to the insects and covers topics such as the evolution
of the group and the factors that made them such successful organisms. It
discusses the importance of their role in terrestrial ecosystems and outlines
the features of structure, function, and physiology that they share. The sec-
ond section provides a semi-pictorial key to the insect orders. Unambiguous
text, coupled with clear drawings designed to highlight key features, allow the
reader to assign most adult insects to the correct order. The bulk of the book
is devoted to the essentials of the 28 living insect orders. The final section
deals with collecting techniques.
Acknowledgements
We thank the following people for providing the photographs that decorate
the new edition of our book: Nicky Bay, Paul Brock, John Gausas, Martin
Gore, Matt Doogue, Piotr Naskrecki, Ed Phillips, Mike Picker, Hans Pohl,
Anne Riley, Gilles San Martin, John Smit, Zestin Soh, Rupert Soskin, and
Manfred Ulitzka.
Figure Acknowledgments
All illustrations not listed below are © Richard Lewington, 2001 and 2023
Preface page vi Vertical bar chart © Oxford University Press 2001.
Pages 4-7 Diagrams of the arthropod-like groups © Oxford University Press
2001.
Page 13 Diagram from Wilmer, P., Stone, G. and Johnston, I. (2000) Environ-
mental Physiology of Animals. Blackwell Science, Oxford.
Page 15 Diagram from Gullan, P. J. and Cranston, P. S. (2000) The insects: an
outline of entomology, 2nd edition, Blackwell Science, Oxford.
Page 17 Diagrams from Chapman, R.F. (1998) The insects: structure and func-
tion, 4th edition. Cambridge University Press. © After Hepburn, H.R. (1985)
Structure of the integument, in Comprehensive insect physiology, biochemistry
and pharmacology, Vol. 3 (eds G.A. Kerkut and L.I. Gilbert), Pergamon Press,
Oxford.
Page 19 Diagram © Oxford University Press 2001.
Page 20 Diagram redrawn from Kukulova-Peck, J. (1991) The insects of Aus-
tralia, 2nd edition. Melbourne University Press, Australia. © CSIRO Australia
1991.
Page 21 Diagram after Chapman, R.F. (1998) The insects: structure and
function, 4th edition. Cambridge University Press.
Page 22 Diagram © Oxford University Press 2001.
Page 36 Diagram of the phylogeny of hexapods. Adapted from © Hans Pohl
(2014).
Page 286 Diagram © George C. McGavin 2001.
Contents
Glossary xvii
Section 1 Introduction to insect evolution

and biology 1
······························································································
The evolution of the arthropods 3
The origins of insects and other hexapods 7
Five factors in a winning formula 8
Interactions with other organisms 23
Insect structure 28
Conclusion 31
Section 2 The insect orders 35

······························································································
Insect groups that predate the evolution of wings 37
Archaeognatha (bristletails—alternative name:
Microcoryphia) 38
Zygentoma (silverfish and firebrats—alternative name:
Thysanura) 41
The winged insects 44
Division Palaeoptera 45
Ephemeroptera (mayflies) 45
Odonata (dragonflies and damselflies) 51
Division Neoptera 59
Superorder Exopterygota (= Hemimetabola) 60
Polyneoptera 60
Haplocercata 60
Dermaptera (earwigs) 61
Zoraptera (angel insects) 65
Plecoptera (stoneflies) 69
CONTENTS xiii
Orthoptera (grasshoppers and crickets) 73

Dictyoptera 84
Blattodea (cockroaches and termites—alternative
name: Blattaria) 85
Cockroaches 87
Termites 91
Mantodea (praying mantids) 98
Xenonomia 104
Grylloblattodea (ice crawlers—alternative name:
Notoptera) 105
Mantophasmatodea (heel walkers—alternative name:
Notoptera) 108
Eukinolabia 112
Phasmatodea (stick insects and leaf insects) 113
Embioptera (webspinners—alternative name: Embiidina) 118
Paraneoptera (= Acercaria) 122
Psocodea (barklice, booklice, and true lice) 123
Hemiptera (true bugs) 134
Thysanoptera (thrips) 151
Division Endopterygota (= Holometabola) 156

Neuropterida 156
Raphidioptera (snakeflies) 157
Megaloptera (alderflies, dobsonflies, and fishflies) 161
Neuroptera (lacewings, antlions, and mantidflies) 165
Coleopterida 172
Coleoptera (beetles) 173
Strepsiptera (strepsipterans) 189
Antliophora 196
Mecoptera (scorpionflies) 197
Siphonaptera (fleas) 202
Diptera (flies) 210
Amphiesmenoptera 227
Trichoptera (caddisflies) 228
Lepidoptera (butterflies and moths) 235
Hymenoptera (bees, wasps, and ants) 255
xiv CONTENTS
Section 3 Fieldwork 277

······························································································
Collecting 279
Killing methods and data recording 296
Specimen preservation 298
Long-term preservation 299
Subject Index 303

Biographies
George McGavin studied Zoology at Edinburgh University, followed by a

PhD in entomology at Imperial College and the Natural History Museum
in London. After 30 years as an academic, mostly at Oxford University, he
became an award-winning television presenter. George is an Honorary Re-
search Associate of the Oxford University Museum of Natural History and
an Honorary Principal Research Fellow at Imperial College. He is a Fellow of
the Linnean Society an Honorary Fellow of the Royal Society of Biology, and
an Honorary Life Fellow of the Royal Entomological Society. In 2019 George
became the President of the Dorset Wildlife Trust.
Leonidas-Romanos Davranoglou studied Zoology at Imperial College

London. He then undertook his DPhil thesis in entomology at the Depart-
ment of Zoology, University of Oxford, funded by the prestigious Oxford
Natural Motion and the Onassis Foundation scholarships. In 2021, Leonidas
was awarded with both a Leverhulme Trust Early Career Fellowship and the
John Fell OUP Fund at the Oxford University Museum of Natural History to
study the evolution of insect communication. Leonidas will be researching
xvi BIOGRAPHIES
how insects communicate using sounds by examining the morphology and

biomechanics of living species, as well as their extinct relatives preserved in
100-million-year-old Cretaceous amber.
Glossary
Aedeagus The organ of copulation in male insects.

Ametabolous Development without metamorphosis, such as in bristletails
and silverfish.
Apodous Without legs.
Apodemes Internal tendon-like extensions of the cuticle, which serve as
muscle attachment sites.
Apolysis The separation of the old and new cuticles.
Aposematic Having bright colouration to serve as a warning to predators.
Usually red, yellow, and black.
Apterous Lacking wings.
Apterygota (adj. apterygote) Primitively wingless insects. The smaller
subclass of the Insecta.
Arrhenotoky Parthenogenesis where haploid males are produced from
unfertilized eggs.
Asexual Of reproduction without separate sexes. Reproduction occurring
by parthenogenesis.
Autotomy The casting or shedding off of a part of the body, for example,
when under threat.
Batesian mimicry Where an edible or palatable species mimics the colours
and patterns of an inedible species. The mimics are rare relative to the
models.
Biological control Use of natural predators, parasites, or disease producing
organisms to effect a reduction in the populations of pest species.
Biramous Having two branches.
Bisexual Having two separate sexes.
Caste Any group of individuals in a colony of social insects that is
structurally or behaviourally different from individuals in other groups,
that is, soldiers, workers, and reproductives as seen in termites, bees, and
ants.
Cerci (sing. cercus) A pair of terminal abdominal appendages, which
typically have a sensory function.
Chitin A high molecular weight, unbranched, amino-sugar polysaccharide
made up of β (1–4) linked units of N-acetyl-D-glucosamine. Chitin is the
main structural element in arthropod exoskeletons and is also found in
fungal cell walls.
Chrysalis The alternative name for a lepidopteran pupa.
Class A major taxonomic subgroup, above Order and below Phylum.
xviii GLOSSARY
Coelom The main body cavity surrounding the gut in animals such as
annelid worms, starfish, and vertebrates. In arthropods and molluscs, the
coelom is much reduced, and the main body cavity is a haemocoel.
Colony An aggregation of social insects sharing a nest.
Corbiculum (pl. corbicula) The pollen basket of honey bees, which is a
concave, shiny area on the hind tibiae, fringed with stiff hairs.
Cosmopolitan Occurring throughout most of the world.
Coxa (pl. coxae) The first segment of an insect’s leg, joining the rest of the
leg to the thorax.
Crop An area of the foregut in insects where food is stored.
Cuckoo A term used for an insect that uses the food stored by another to
rear their own young.
DDT Dichlorodiphenyltrichloroethane.
Dimorphic Occurring in two distinct forms. The sexes of some insects are
differently coloured or shaped.
Ecdysis The last stage of moulting, which is the casting or sloughing of the
old cuticle.
Elytra (sing. elytron) The rigid front wings of beetles, modified as covers for
the hind wings and not used in flight.
Endite A lobe of a leg segment that is directed inwards towards the midline
of the body.
Endopterygota (adj. endopterygote) Insects where the wings develop
internally. Metamorphosis is complete and there is a pupal stage
(holometabolous).
Entognathous Having eversible mouthparts contained inside the head
within a small pocket.
Epidermis In insects, the single layer of cells that secretes the cuticle.
Exite A lobe of a leg segment that is directed outwards away from the
midline of the body.
Exopterygota (adj. exopterygote) Insects where the wings develop outside
the body. Metamorphosis is incomplete and there is no pupal stage
(hemimetabolous).
Family A taxonomic group below that of Order and comprised of
subfamilies, genera, and species.
Fat body An aggregation of metabolic and storage cells in the haemocoel.
Femur (pl. femora) The portion of an insect’s leg that corresponds to the
mammalian thigh. The order of segments is
coax–trochanter–femur–tibia–tarsus.
Fungivorous Eating fungi.
GABA Gamma-aminobutyric acid. Typically an inhibitory transmitter of
neuromuscular junctions. Gamma is often shown symbolically:
γ-aminobutyric acid.
GLOSSARY xix
Gall An abnormal plant growth caused by a virus, bacterium, fungus, mite,

or insect.
Genitalia The hard parts of insect reproductive systems that engage
between males and females during mating.
Genus (pl. genera) A low taxonomic group composed of species.
Haemocoel The haemolymph-filled internal body cavity of arthropods,
such as insects, which is essentially an enlarged part of the circulatory
system.
Haemolymph A fluid in invertebrate animals that is the equivalent of
blood.
Haltere(s) The greatly modified hind wings of Diptera, which serve as
balancing organs.
Hemimetabolous Developing by incomplete or gradual metamorphosis,
for example, in the Orthoptera and Hemiptera. Immature stages are
called nymphs (see exopterygota).
Hemimetabola Those orders with incomplete metamorphosis.
Holometabolous Developing by complete metamorphosis, for example, in
the Diptera and Lepidoptera. Immature stages are called larvae (see
endopterygota).
Holometabola Those orders with complete metamorphosis.
Homeotic genes Genes involved in the regulation of body plan.
Homology Character states acquired by direct inheritance from an
ancestral form.
Homoplasy Character states acquired by convergent evolution.
Honeydew The carbohydrate-rich excrement of plant sap feeders such as
aphids.
Hyperparasitoid A parasitoid of another parasitoid.
Idiobiont A parasitoid that kills or paralyses its host when laying an egg,
and thus preventing the host from developing any further.
Inquiline A species that lives in the nest, gall, or home of another.
Insect The word insect comes from the Latin, insectus, past participle of
insecare, to cut into, and refers to the major divisions of an insect’s body.
Integument The cuticular and cellular covering of insects and other
arthropods (i.e., the epidermis and cuticle combined).
Koinobiont A parasitoid that does not kill the host when egg laying, thus
allowing the host to develop further.
Lobopods Protoarthropods with antennae, annular body segmentation
(not a jointed exoskeleton), and soft, fleshy appendages bearing claws.
Meconium The accumulated solid wastes of the larval stages of some
insects, which are expelled when the adult stage is reached.
Metamorphosis The transformation between the immature form and the
adult form, with the forms being different.
xx GLOSSARY
Monophagous Restricted to eating a single species plant or type of

foodstuff.
Monophyletic Of organisms that have descended from a single common
evolutionary ancestor or ancestral group.
Mullerian mimicry Where several inedible or unpalatable species converge
in colour pattern. As all the species are non-palatable, mimicry rings, as
they are known, are enhanced by the frequency all ring members and
even distantly related species can be involved.
Mycetocytes Special cells that permanently house intracellular bacterial
symbionts.
Myrmecophilous Greek for ant-loving. Any species that lives with and
depends on ants for food, protection, and care.
Neotenic Retaining features of the immature stages in the adult.
Ocelli Simple light-receptive organs on the head of many insects.
Oligophagous Restricted to eating a small range of related plants or
foodstuffs (see polyphagous, monophagous).
Ommatidia Individual light receptive units in the compound eyes of
insects. Each ommatidium comprises a clear lens system and
light-sensitive retinula cells.
Omnivorous Having variable eating preferences.
Ootheca Protective egg case. Can contain a single egg in some orders, and
up to several hundred eggs in other orders.
Order A taxonomic rank below a class and above a family.
Oviparous Producing young by means of eggs expelled from the body
before they are hatched.
Ovoviviparous The young are produced by means of hatching eggs inside
the body.
Palaeozoic (Paleozoic) The first of the three eras of the Phanerozoic
(having visible life). The Palaeozoic includes the Cambrian, Ordovician,
Silurian, Devonian, Carboniferous, and Permian periods. The Palaeozoic
is followed by the Mesozoic and Cenozoic eras.
Parasite (adj. parasitic) A species living off the body or tissues of another,
but not causing the death of the host.
Paraphyletic Of a group of species that includes some, but not all, of the
species that share a single common ancestor.
Parasitoid A species that develops by consuming the body tissues of a
single host, eventually causing its death (see idiobiont and koinobiont).
Parthenogenesis (adj. parthenogenetic) Reproduction without the need for
fertilization.
Perineurium A layer of glial cells inside the neural lamella, which acts as
the blood–brain barrier.
Pheromone A volatile substance produced by insects to communicate with
others for the purposes of reproduction, aggregation, defence, etc.
GLOSSARY xxi
Phloem The main transport vessels of plants taking nutrients from the
leaves to other parts.
Phyla (sing. phylum) The major groupings of the animal kingdom,
comprised of superclasses and classes, for example, Mollusca,
Arthropoda, Chordata, etc.
Physogastric When the abdomen (usually in females) becomes greatly
distended and membranous, typically after a meal, when pregnant, or
when storing fat or food.
Polyembryony A single egg giving rise to many embryos by subdivision.
Polymorphic Having more than two forms.
Polyphagous Eating a wide range of plants or foodstuffs.
Polyphyletic Of a group of species that have different ancestors. Such taxa
are ‘unnatural’ or incorrect.
Predacious (n. predator) Eating other animals.
Proleg A fleshy abdominal limb of insect larvae.
Pronotum The dorsal cover of the first segment of the thorax.
Pruinescent (colouration) Of a powder, or powdery.
Pterygota (adj. pterygote) Winged insects. The larger subclass of the
Insecta.
Rostrum The prolonged part of the head of weevils and scorpionflies. Also
the tubular, slender, sucking mouthparts of insects such as the Hemiptera
(also known as the labium in the latter).
Saprophagous Eating decayed organic material.
Sclerite A hardened chitinous body plate surrounded by flexible
membranes or sutures (Gk. skleros—hard).
Sister groups Groups of the same taxonomic rank resulting from the
splitting of an ancestral lineage.
Solitary Occurring singly or in pairs, not in social groups.
Somite A single body division of a segmented animal.
Species A group of living organisms consisting of related similar
individuals capable of exchanging genes or interbreeding. A taxonomic
rank below a genus.
Spiracle(s) The breathing holes of insects leading to the tracheae occurring
at the sides of the abdomen and thorax.
Stridulation The act of producing sound, usually by rubbing two parts of
the body together.
Subfamily A taxonomic subdivision of Family.
Subimago Pre-adult winged stage of Ephemeroptera.
Suborder A taxonomic subdivision of Order.
Suture A groove on the outside of an insect body segment that indicates the
fusion of two sclerites.
Symbiosis Different species living in an association that brings mutual
benefit.
xxii GLOSSARY
Synanthropic Associated with humans and their habitations.

Synonymy Where a single species has been described and named more
than once
Tagmata (sing. tagma) A group of body segments that together form a
major functional unit such as the head, thorax, or abdomen.
Tarsus (pl. tarsi) The foot of an insect. It is attached to the end of the tibia
and made up of a small number of subsegments or tarsomeres.
Thelytoky Parthenogenesis where only female offspring are produced.
Tibia (pl. tibiae) The lower leg segment of insects, corresponding to the
mammalian shin.
Tracheae (sing. trachea) The internal airways of insects.
Trochanter The small segment of the insect leg between the coxa and the
femur.
Trophallaxis The exchange of food between colony members in social
insects.
Trophic level The levels of food or energy transfer in food chain. Food
producers or autotrophs form the first level, herbivores form the second
level, and primary carnivores the third level. Other carnivores form the
fourth and fifth levels.
Vector An intermediate host that carries a disease organism in contact with
its target organism.
SECTION 1
Introduction to insect
evolution and biology
3
The evolution of the arthropods

The Earth formed from gas and debris around 4,600 million years ago (mya).
The first traces of life appear as fossils of blue-green algae around 3,600–
radiodonts
Early arthropods. The front 3,400 mya, and yet for more than 2 billion years, nothing more complicated
section of the elongate body than bacteria, algae, and minute, unicellular planktonic organisms existed.
carried a pair of appendages Then evolution went into overdrive. Known popularly as biology’s ‘big bang’,
and a ventral mouth with
distinctive radiating teeth. The the time around the beginning of the Cambrian period (543 mya) saw the
trunk of the segmented body ‘sudden’ appearance in the fossil record of an incredible assemblage of mul-
had lobe- or flap-like
appendages. Some were
ticellular animals, some of which were the ancestors of all species on Earth
predators while others were today. Chief among these animals were early arthropods, such as radiodonts
filter feeders and lobopods.
lobopods Although the Cambrian explosion is characterized by some exceptionally
early arthropods with
antennae, annular body
well-preserved material, such as those found in the Burgess Shale deposits of
segmentation (not a jointed British Columbia, it is often difficult to say exactly which might have been
exoskeleton) and soft, fleshy the direct ancestors of modern forms. The process of fossilization is un-
appendages bearing claws.
Earlier studies suggested that predictable, and fine detail is often recorded only under special conditions.
velvet worms (Onycophora) Evolutionary intermediates may never be found. The segmented, worm-like
and water-bears (Tardigrada)
forebears of these creatures had certainly arisen by the late Precambrian (the
are direct descendants of
lobopods, although recent Vendian), for the early arthropods had already evolved into a range of clearly
works suggest that their recognizable groups with distinct body plans by the early to mid-Cambrian.
similarities are likely only
superficial Arthropods differ from each other chiefly in the number and design of
homeotic genes
their appendages. The discovery of homeotic genes and the function of some
genes involved in the of these genes has provided some insight into the early diversification of the
regulation of body plan group.
We now know how certain homeotic genes may have determined the evo-
lution of segment diversity, as well as how they led to the evolution of the
dazzling range of body plans exhibited by arthropods as a whole.
Just as the invention of simple and then compound microscopes in the sev-
enteenth century advanced the biological techniques of the day, and therefore
made it inevitable that old classifications had to be abandoned and new ones
adopted, molecular techniques are producing data that sometimes contradict
views based on traditional morphology. While the phylogeny (evolutionary
history) of any group may be inferred from morphological or molecular stud-
ies, both approaches can suffer from being unable to distinguish whether
or not particular character states shared by two organisms are due to com-
mon evolutionary descent (acquired by direct inheritance from an ancestral
form—homology, or by convergent evolution—homoplasy). Very different
animals might evolve features that make them look superficially similar (e.g.
moles and mole crickets; see figure on next page), and it is not unusual for
natural selection acting over a long time to make closely related species look
monophyletic very different from each other (e.g. elephants and hyraxes).
(of) a group of species that Arthropods are a monophyletic group, that is, they have a single common
share a common ancestor
ancestor. The staggering morphological diversity found in today’s arthro-
polyphyletic pods led some authors in the past to suggest a polyphyletic origin, where
(of) a group of species having arthropods evolved from four different types of worms that independently
different ancestors. Such taxa
are ‘unnatural’ gained an exoskeleton. This view is now obsolete. We know that arthropods
Essential Entomology. Second Edition. George C. McGavin and Leonidas-Romanos Davranoglou, Oxford University Press.
© George C. McGavin and Leonidas-Romanos Davranoglou (2022). DOI: 10.1093/oso/9780192843111.001.0001
4 SECTION 1 INTRODUCTION TO INSECT EVOLUTION AND BIOLOGY
Front legs of the mole and

the mole cricket,
demonstrating their
similarity. Foreleg of mole Foreleg of mole cricket
originate from a single worm-like ancestor, which gained a chitinous, seg-

mented exoskeleton that was regularly shed to allow the organism to grow
and to heal from injuries. The cousins to all other arthropods, the tardi-
grades or water bears (Tardigrada) and the velvet worms (Onychophora), do
not possess a rigid exoskeleton, but a soft cuticle also made of chitin, which
they regularly shed, therefore giving us an idea of how early arthropods may
have looked. Arthropods, tardigrades, and velvet worms all belong to a likely
monophyletic lineage known as the Panarthropoda.
Features that characterize arthropods

• Possession of a chitinous cuticle, which acts as a rigid exoskeleton for
the internal attachment of striated muscles and is periodically moulted to
allow for growth.
• Segmental paired legs.
• Body segments aggregated and/or fused into tagmata (discrete functional
units) of which the most universal is the head. Besides the head there may
be a trunk, as in the myriapods, or a separate thorax and abdomen, as in
the crustaceans and hexapods.
The arthropod-like groups (Panarthropoda)

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Onycophora
Onycophora (velvet-worms) > 200 species

Onycophorans are soft-bodied, worm-like animals that are confined to
Velvet worm tropical soil and litter.
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Tardigrada
Tardigrades (water-bears) > 1,300 species

Tiny (less than 1 mm from head-to-tail) and typically live in the water films
associated with mosses.
Water bear
NB: Illustrations not to scale.

THE EVOLUTION OF THE ARTHROPODS 5
The true arthropods (Euarthropoda)

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Trilobita
Trilobita (trilobites) > 20,000 species
Extinct marine, bottom-living, and pelagic arthropods that ranged from
1 mm to 1 m in length with the body divided into three regions: a head or
cephalon, a mid-body or thorax, and a hind-body or pygidium. The head
Trilobite had antennae and eyes, the latter similar to the compound eyes of modern
arthropods.
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Chelicerata
Merostomata (horseshoe crabs) 4 species
Horseshoe crabs are large, marine, bottom-living scavengers. Species of the
genus Limulus, which belong to the Xiphosura, a group little changed since
the Silurian, can reach up to 60 cm in length. The Eurypterida, predatory
giant sea scorpions, were very abundant in the Silurian and up to 2 m long.
Mygalomorph spider Arachnida (spiders, scorpions, harvestmen, ticks, mites, psuedoscorpions,

palpigrades, sunspiders, whipscorpions, whipspiders, and ricinuleids) >
100,000 species
A largely terrestrial group, although some mites and a few spider species
are found in freshwater. Most arachnids are predators, but some mites are
parasites or herbivores.
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Pycnogonida
Pycnogonida (sea spiders) > 1,300 species
Highly specialized, marine, inconspicuous, leggy arthropods which do not
have an obvious trunk or body. Sea spiders are predators of hydroids and
polyzoans.
Sea spider
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Pancrustacea (crustaceans and hexapods)

Cephalocarida (cephalocaridans) 12 species
These are small (< 4 mm), marine, bottom-living, filter-feeding crustaceans
first described in the 1950s.
Krill Branchiopoda (water-fleas, fairy shrimps, and brine shrimps) ~ 1,100 species
Branchiopods are small, filter-feeding, freshwater and marine crustaceans.
Unlike most crustaceans, they have flat, leaf-like body appendages called
phyllopodia.
Oligostraca (mussel shrimps and seed shrimps) > 13,000 species

Ostracods are marine and freshwater crustaceans mostly around 1 mm long
(although extinct species were ten times longer). The entire body is enclosed
in a bivalvate shell or carapace.
Malacostraca (mantis shrimps, crabs, shrimps, krill, lobsters, and hermit crabs)
> 40,000 species
Most living Crustacea, including all the larger shrimp- and crab-like
species, belong to this group. They are primarily marine, although some are
freshwater or terrestrial.
Maxillopoda (copepods, fish lice, and barnacles) > 14,000 species

These small, filter-feeding crustaceans are major elements of marine plank-
ton. Some species are parasites of fish and crabs, and some live in freshwater.
This is a polyphyletic group, and some lineages are related to the Ostracoda.
Remipedia (remipedians) 28 species

The first representative of this marine group (Speleonectes sp.) was found in a
sea cave in 1980. Remipedians are the sister group to the Hexapoda.
··································································································································································································
Hexapoda
Collembola (springtails) > 9,000 species

Springtails are mainly terrestrial, but a few occur on the surfaces of fresh and
saltwater pools. They are mostly detritivores but also eat pollen and algae,
and a few are pests of crops, such as sugarcane. They are < 4 mm in length
and have a special abdominal ‘spring-and-catch’ jumping mechanism.
Protura (proturans) > 800 species

Dragonfly Proturans are soft-bodied, minute (< 2 mm), soil-dwelling, blind, fungus
feeders.
Diplura (diplurans) 800 species

Diplurans are small to quite large (5–50 mm), slender, blind, terrestrial
hexapods. Most species live under stones and in soil and litter. The abdomen
has a pair of conspicuous terminal cerci.
Insecta (insects) > 1,000,000 species

Insects are mainly terrestrial but there are a significant number of aquatic
species. Ranging in size from minute parasitic wasps at around 0.2 mm to
stick insects measuring 30 cm in length, insects have evolved very diverse
lifestyles. The head bears external mouthparts, antennae, and compound
eyes. The thorax bears three pairs of legs and (typically) two pairs of wings.
Insects have a well-developed tracheal system.
THE ORIGINS OF INSECTS AND OTHER HEXAPODS 7
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Myriapoda
Pauropoda (pauropods) > 800 species

Pauropods are soft-bodied, soil-dwelling, blind, scavenging myriapods that
are < 2 mm long.
Diplopoda (millipedes) > 12,000 species

Millipedes are terrestrial, slow-moving, mostly herbivorous, burrowing myr-
iapods with elongate cylindrical bodies. They vary considerably in size (<
Millipede 2 mm to > 25 cm) and are typically round in cross section. The first three
segments of the trunk have no legs, but the remaining segments are fused in
pairs, each bearing two pairs of legs.
Chilopoda (centipedes) 8,000 species

Centipedes are terrestrial, carnivorous myriapods with venom-bearing
claws derived from the first pair of legs. Some are elongate, thread-like,
slow-moving, soil dwellers, while others are long legged and fast running.
Centipedes can range from 5 mm to 20 cm in length. The trunk segments
each bear a single pair of legs.
Symphyla (symphylans) 200 species

Symphylans are typically small (1–8 mm), soft-bodied, blind, soil- or litter-
dwelling, herbivorous myriapods.
NB: Illustrations not to scale.
The origins of insects and other

hexapods
Insects evolved on land from arthropod ancestors that had long before
made their way from the sea. However, the lack of fossil material has
meant that it has been challenging to find details about this sea-to-land
transition.
Most of the fossils of the earliest purported insects from the Devonian
have now been shown to either be an entirely different organism altogether,
or they are too poorly preserved to enable a reliable identification. Rhyniella
praecursor, for example, was initially thought to be a larval insect, and is now
known to be a springtail (Collembola). One of the most important of these
fossils undoubtedly is Rhyniognatha hirsti, which, for more than a decade,
was considered as the earliest example of a flying insect, and consequently
was of essential importance for disentangling the complex origins of flight.
However, a 2017 study suggested that Rhyniognatha more likely represents
an early centipede, confirming previous studies indicating a myriapod origin
for this fossil. It is clear that the deep origin of insects is clouded by much
debate and will likely remain so in the near future unless spectacular new
fossils come to light and change the game.
Based on this consideration, one could be forgiven for taking the rather
pessimistic view of John L. Cloudsley–Thompson, who, in 1988, wrote the
following:
Time spent on insect phylogeny is often wasted. The origins of most insectan
orders is a mystery, and is likely to remain so into the foreseeable future—
certainly until existing fossils have been studied more extensively and further
palaeontological evidence has been collected.
Nevertheless, Cloudsley–Thompson concluded that they evolved from

entognathous terrestrial ancestors, which had antennae, entognathous labiate mouthparts,
having eversible mouthparts probably 14 post-cephalic segments, tracheal respiration, 12 pairs of legs,
contained inside the head
within a small pocket coxal styles, eversible vesicles on most segments, and a pair of terminal cerci.
The validity of his assertions is highly debated, since, as one can imagine, they
cerci (sing. cercus)
a pair of terminal abdominal are highly speculative.
appendages that typically A revolution in our understanding of hexapod evolution came not from
have a sensory function
fossils, but from studying the DNA and structure of extant insects. Phy-
logenomic and morphological studies have convincingly suggested that the
Remipedia, a group of elusive, marine cave-inhabiting crustaceans are the
sister groups sister group of insects and hexapods as a whole. Although we may have not
groups of the same taxonomic
rank resulting from the
yet found the closest relatives of hexapods in the fossil record, these find-
splitting of an ancestral lineage ings clearly demonstrate that insects are nested deeply within crustaceans,
and they could be described as flying prawns!
Five factors in a winning formula

The staggeringly high insect diversity on Earth today is the result of a com-
bination of high rates of speciation and the fact that many insects are very
persistent, that is, they show relatively low rates of extinction. Most insect
species may survive, little changed, for 2 million years or so, but a few species,
found in 35-million-year-old Baltic amber, are morphologically identical to
living species. A number of central factors that have led to the success of the
Insecta can be identified as crucial.
Five factors contributing to the success of insects

• Small size
• A protective cuticle
• An efficient nervous system (the blood–brain barrier and sensory neuro-
motor refinement)
• The evolution of flight
• High reproductive rate
Size
Your size determines how the environment will affect you. The range of sizes
of all organisms covers eight orders of magnitude, from the smallest bacterial
FIVE FACTORS IN A WINNING FORMULA 9
cell at around 0.3 µm (micrometres) long to a blue whale at 30 m long. In the

middle of this range, organisms of around 3 mm—the insects—dominate. In
rain forests, where, intuitively, large species might be expected to abound,
90% of the beetles collected were < 5 mm in length. To really understand the
insect world, you need to imagine how the physical environment affects small
Log species richness

3
A typical relationship
between log body weight –2 –1 0 1 2 3 4 5
and insect species richness. Log body weight (mg)
creatures. Take falling for example. On Earth, the acceleration due to gravity
is a constant, but its effects vary according to the size of the falling object.
Falling pollen grains and similarly small objects, for example, may be blown
around for weeks before they reach the ground, and no matter the height from
which an insect is dropped, it will not be damaged.
J. B. S. Haldane takes up the story:
You can drop a mouse down a thousand-foot mine shaft and, on arriving
at the bottom, it gets a slight shock and walks way. A rat is killed, a man is
broken and a horse splashes.
It is due to the ratio of surface area to volume. Area is proportional to the
square of the length (l2), whereas volume is proportional to the cube of the
length (l3). Drag forces cause the terminal velocity achieved by falling objects
to be proportional to their frontal surface area-to-mass ratio. Thus, a 1-kg
lead ball will attain a much higher terminal velocity than a 1-kg lead sheet,
and a 1-cm diameter stone will attain a higher terminal velocity than a 1-cm
diameter sponge. For most organisms, however, the density of the tissues is
much the same, and volume is thus a good measure of mass. A petit pois has
a much smaller volume (and therefore mass) in relation to its greater surface
area than a ripe water melon, which has a large volume (and therefore rel-
atively greater mass) in relation to its small surface area. If you are still not
convinced, ask yourself which you would rather drop on your kitchen floor.
More massive objects have greater momentum; or put simply, the bigger they
come, the harder they fall.
Small animals are also relatively more powerful than large ones. The cross-
sectional area of insect muscles is relatively large compared to the mass they
are supporting. Humans are able to carry a load roughly equal to their body
weight, whereas the load-carrying ability of an ant is many times its own body
weight, and earwigs are able to move more than fifty times their own weight.
A chrysomelid beetle hanging on to a leaf in a gale may resist a force ten or

more times greater than its mass because the adhesive structures on its feet
have a relatively huge surface area compared to the rest of the body.
What other consequences are there to being small? The world is a much
finer-grained environment to small animals and offers a lot more places to
live. Insects have simply evolved to occupy many more ecological niches than
larger organisms ever could. An oak tree during the course of its life may
support a few mammal and bird species, but many hundreds of different
insect species. Dozens of species of gall wasp attack the oak’s buds, leaves,
flowers, acorns, and roots. The leaf-mining larvae of flies and moths neatly
graze inside its leaves, secured between the upper and lower epidermal lay-
ers. Scuttling, flat bugs move about under flakes of its bark, feeding on fungal
threads, while many more closely related species insert slender, needle-like
mouthparts into the oak’s xylem or phloem vessels, or systematically empty
the contents of individual cells. The larvae of longhorn, bark, and ambrosia
beetles devour the very fabric of the tree. In the height of summer in north-
ern latitudes, the sound of falling caterpillar droppings can be heard quite
clearly. Each of these herbivorous insects may fall prey to a number of carni-
vores and they all may act as hosts to one or more species of parasitic wasps or
flies. Nests, fungi, lichens, and the rain-filled hollows left when branches have
fallen all support rich and varied communities. When the tree ages and finally
dies, its decaying structure will be alive with insects for months or years to
come. Our oak tree is just one of 600 species of oak found growing in northern
temperate regions of the world, each with a distinctive insect fauna.
That there are no really large insects today does not mean that it would be
impossible for them to exist. It is often stated that the upper limits to the size
of an insect are constrained by the mechanics of moving with an exoskeleton
spiracles and the difficulties of getting enough oxygen to all the tissues via spiracles and
openings to the tracheal diffusion alone, but this is not really why insects are not bigger than they are.
system of insects, located on
thoracic and abdominal Gaseous exchange by means of air-filled tracheae is quite an efficient sys-
segments tem and, in any case, suitable modifications, such as sac-like air pumps,
would most likely evolve. It is now known that caterpillars possess quite a
sophisticated gaseous exchange system equivalent to the vertebrate lung (see
Lepidoptera). If larger, the weight of an insect’s body at moulting might be
greater than the soft cuticle could bear, and being vulnerable to predators,
they would probably have to hide. However, they could get round the me-
chanical problem by moulting in water or by maintaining their body shape
with hydrostatic pressure (although falling any distance at this stage might
be a disaster). Fossil evidence shows that, in the late Carboniferous and early
Permian, most insects were bigger (some very much bigger) than they are to-
day. Meganeura, a carboniferous dragonfly relative, had a wingspan of 75 cm;
the wingspan of the biggest odonate today, Megaloprepus coerulatus (a beau-
tiful damselfly of South and Central America, which oviposits in rain-filled
bromeliad tanks and tree holes), is a quarter of this. Ramsdelepedion schusteri,
a carboniferous silverfish, was a chunky 6 cm long; today, the very biggest sil-
verfish are approximately 1 cm in length. It has been suggested that large size
evolved as a defensive strategy against predators such as proto-dragonflies.
They, in turn, got bigger and the process continued until it was halted, ei-
ther because they were no longer mechanically viable or, more likely, because
of the appearance and rise of vertebrate predators, such as reptiles, who ate
them. From that time on, insects got smaller and smaller to avoid preda-
tion and were thus able to radiate into all the available empty niches. The
lower limit of the insect size range is determined by a number of physical
constraints.
The heaviest Goliath beetles weigh about 60–100 g, making them heavier
than many birds; at only 1.6 g, the bee hummingbird is lighter than an adult
locust. A few butterflies and moths have very large wingspans (up to 280 mm),
but their bodies are not very massive. The longest species, a stick insect, has
a body length of around 30 cm. The smallest insects are species of parasitic
wasp, being a little more than 0.2 mm from head to tail; this is no more than
the thickness of the cuticle of large insect species and about half the size of
some protozoan organisms. The smallest beetle in the world, a tiny feather-
winged ptinid beetle, could sit comfortably on the very claw tip of a Goliath
beetle.
So what are the constraints of size, and how can insects be so small and
still function appropriately?
Oxygen supply All animal cells need a supply of oxygen and organisms
that are very minute might be able to get enough oxygen by means of simple
diffusion. This is because a small body has a proportionally greater surface
area than volume. Active insects need to get oxygen to all their cells and al-
though oxygen diffuses 10,000 times faster in air than in water, it will not pass
through waterproofed cuticle. The tracheal system of insects is a simple and
effective means of gaseous exchange. Pairs of spiracles, the openings of the
system, are located laterally on various body segments and are connected to
air sacs and large tracheae, which then branch into even smaller branches,
getting finer and finer. The finest branches, the tracheoles, are in intimate
contact with the tissues and are especially abundant in tissues where oxygen
demand is high. Interestingly, no matter how big or small the insect, the finest
tracheoles always have a diameter of 0.2 µm. This is because if the tubes were
any narrower, oxygen molecules would simply not diffuse any further along
them. Evolution has produced a tube diameter that is ‘just right’. A small in-
sect can have a small number of tracheoles, but they cannot be any finer than
0.2 µm.
Cell size Another reason why insects may have reached their minimum po-
tential size might be due to the fairly constant size of animal cells. For various
reasons concerning internal transport and operation dictated by relationships
between surface area and volume, animal cells tend to be the same size. From
this, it follows that larger animals are made up of more cells and smaller an-
imals of fewer cells, and there must be a minimum number of cells needed
to make a functioning insect. An adult locust, for instance, may have a mil-
lion or so neurons to serve all its various sensory and motor needs. Although
axons can vary in length greatly, nerve cell bodies tend to be similarly sized.
A flying insect the size of a full stop might be able to fit in between one and
ten thousand neurons, but any fewer would be inadequate for it to function.
To overcome these difficulties, insects have very economical wiring and have
shifted the job of integration from the central nervous system (CNS) to the
peripheral nervous system.
Vertebrate muscles have a single motor neuron innervating each separate
muscle fibre. To increase or decrease the force exerted by a muscle requires
the recruitment of more or fewer motor neurons. Such nervous extravagance
is impossible in tiny creatures. Instead, insects have polyneural, multitermi-
nal innervation. In other words, only a few motor neurons innervate an entire
muscle by branching extensively and by having multiple terminals. There are
three types of motor neuron: fast, slow, and inhibitory. Fast motor neurons
innervate most of the fibres within a muscle and their terminals release the
excitatory transmitter l-glutamate, causing fast ‘twitch’ responses. Slow mo-
tor neurons run to a smaller subset of fibres and also release l-glutamate,
causing slow, graded contractions. Inhibitory motor neurons innervate to all
muscle fibres and their terminals release the neurotransmitter GABA, which
inhibits contraction. In addition, neuromodulatory neurons are used to ‘fine-
tune’ the system by setting the gain and threshold of motor innervation and
muscle response. This arrangement allows simplification of the nervous sys-
tem while still retaining flexibility. In vertebrate animals, the CNS is heavily
involved in the integration of motor patterns. However, in insects, the mo-
tor neurons themselves play an important part in the integration of walking,
flying, and reflex leg movements. There are also ways in which insects can
shift the early stages of sensory integration to their peripheral nervous system.
A good example comes from the African migratory locust; the sensitivity of
chemosensillae on the maxillary palps is affected by the nutritional state of
haemolymph the animal and it is measured by the levels of nutrients in the haemolymph.
the body fluid in invertebrate
animals that is the equivalent In other words, if a locust has fed on carbohydrate-rich food, its sen-
of blood sitivity to carbohydrate is reduced and other nutrients, such as protein,
will be phagostimulatory. The locust can thus regulate its diet selection
in quite a sophisticated manner without the need for high-level CNS
integration.
Temperature regulation The large surface area-to-volume ratio of insects

can make it difficult to keep cool and expensive to keep warm. Insects are
ectothermic, but this is not to say that insects cannot regulate their tem-
perature at all. Bees and moths, for example, need to warm up their flight
muscles before take-off, and, when flying in hot conditions, need to lose heat
by increasing haemolymph flow to the abdomen, where excess heat can be
lost. Even in the hottest of environments, insects can survive. Desert beetles
and ants often have very long legs and run fast to minimize contact with the
scorching sand. The important point to remember, however, is that in any
habitat there will always be a great range of temperatures available. Cooler,
moister microhabitats will exist near plants or under stones, and burrowing
underground, even a small distance, will ameliorate the hottest desert condi-
tions. Small animals like insects are much more able to take advantage of a
thermally heterogeneous and fine-grained environment. An elephant cannot
move from an air temperature of 50◦ C to a much more bearable 30◦ C simply
by hiding under a leaf; put another way, microclimates matter much more to
insects (see figure below).
Shaded leaf
28.2°C
25.7°C
33.0°C Sunlit leaf
29.5°C
31.0°C 32.2°C
Microclimate matters. The 27.5°C

relation between body 10 cm
temperature of a dead
50-mg insect, measured with
28.5°C
an inserted thermocouple,
and position on a plant.
Cuticle
The common analogy of a medieval suit of armour for the protective and wa-
terproof outer covering of insects is as much of a hindrance to understanding
as a help. The only similarities are that both are the interface between the
living animal on the inside and its environment, and, just like armour, the
cuticle provides mechanical protection for the body and its appendages.
A conventional suit of armour was made of a single type of metal that was
tough enough to deflect a mace blow or stop an arrow. However, armour has
progressed somewhat in several hundred years. Nowadays special alloys can
stop knives (but not bullets) and semi-flexible polymer materials can stop bul-
lets (but not knives). Composite tank armour made from many layers of metal
and ceramic sandwiched together has increased the effective range of these
fighting machines by greatly reducing their weight. Although we have realized
the great potential of studying natural designs, no biomimetic product has yet
matched the versatility and effectiveness of the insect cuticle.
integument Unlike man-made armour, the integument is, to a large degree, capable
the cuticle, together with the of self-repair. Mechanical damage to larval cuticle, such as a small tear or
epidermis that produces it
puncture, is quickly sealed.
If penetrated, the basement membrane of the integument closes up and
new epidermal cells are produced on either side, and full repair by the pro-
duction of a completely new cuticle takes place at the next moult. The ability
of most insects to regenerate legs after amputation, known as autotomy, is
well known.
Once thought of as a purely inert mechanical barrier, the integument also
provides protection from the biological warfare of most viruses, bacteria, and
fungal pathogens. In many cases, it is also able to mount an immune response

by producing antibacterial proteins.
The cuticle is not simply protective—it also serves as an exoskeleton that
anchors muscles and allows movement. Muscles are attached to the inside of
apodemes the integument and in places the cuticle is extended to form apodemes.
the internal extensions of the The cuticle also bristles with chemo- and mechanosensory devices of all
cuticle, for the attachment of
muscles kinds and has built-in strain gauges to signal loads and tensions within its
structure.
The outward appearances of insects are much more variable than the in-
sides. Most insects have the same kinds of internal organs, even though they
may differ in size by three orders of magnitude. The cuticle has been moulded
throughout the course of evolution to provide all kinds of devices for running,
burrowing, jumping, flying, swimming, catching food, and so on. Cuticle may
be cryptically or brightly coloured, or it may smooth, spiny, or hairy, but it
can be tough enough to cut through metals and hardwood or so delicate that
gases will diffuse through its surface.
Cuticle structure We need to look closely at the structure of cuticle to un-

derstand just how it serves all its various functions. Cuticle is produced by
the epidermis, a single layer of cells supported by a basement membrane. It
covers the whole of the outside of an insect, also extending inside where it
lines the foregut, the hindgut, and the tracheal system. Cuticle is made up of
two major layers: a chitin-containing procuticle, which can be nearly 0.20 mm
thick (200 µm), and a much thinner outer layer, the epicuticle (< 5 µm thick)
(see Figure).
The procuticle is made up of chitin, a high molecular weight, unbranched
amino-sugar polysaccharide made of β -(1→4)-linked units of N-acetyl-d-
glucosamine. Packs of chitin molecules are formed into microfibrils, and
these are embedded side by side in protein to make sheets. These sheets are
laid down in the plane of the cuticle and successive layers are rotated slightly
so that a helicoidal pattern is produced, giving the cuticle a lamellar appear-
ance in cross-section. This arrangement confers great tensile strength. The
outer part of the procuticle becomes tanned or sclerotized, an irreversible pro-
cess of stiffening caused by protein chains becoming chemically cross-linked
to each other by quinones. This dark, sclerotized layer of the procuticle is now
known as exocuticle, while the thicker, inner membranous layer is known as
endocuticle. Very highly sclerotized cuticle is found in areas demanding great
toughness, such as claws and the mandibles of chewing mouthparts. To give
sclerites flexibility at leg joints and between adjacent body sclerites, the procuticle is
a hardened chitinous body not differentiated and only flexible endocuticle is present.
plate surrounded by flexible
membranes or sutures A special rubber-like protein called resilin is often incorporated into
particular regions, for example, wing hinges, and other structures that re-
quire great elasticity and energy storage, for example, those associated with
jumping (see Orthoptera and Siphonaptera).
The outermost layer of insect integument is the thin, brittle epicuticle (1–
4 µm thick). The epicuticle is composed of distinct inner and outer layers,
collectively known as the cuticulin layer. Cuticulin is a composite made from
quinone-tanned protein and lipids and is covered by two further layers of
Cement layer
Wax layer
Superficial layer
Outer epicuticle
Wax filament Inner epicuticle
Pore canal
Exocuticle
Dermal gland
duct opening
Epicuticle
Exocuticle
Procuticle
Endocuticle
Formation
zone
Epidermis
Basement membrane
Cross section through a Dermal
typical insect cuticle. gland
hydrocarbon wax or lipid. The molecules of the inner wax layer are closely
packed as a highly orientated monolayer, while those of the thicker, outer
wax layer are not. It is the waxy epicuticle that is largely responsible for limit-
ing water loss (of major importance in terrestrial environments). Minute pore
canals run at right angles through the cuticle from the epidermal cells to the
surface at densities that can be as high as a million per mm2. These canals,
whose function is to transport waxes (lipids) from the epidermis to the sur-
face, branch into even finer wax canals as they travel through the epicuticle.
The outermost layer in the complicated sandwich that comprises an insect’s
integument is a thin coating of lacquer-like cement.
Moulting As insects grow, they need to periodically replace the rigid ex-
oskeleton. Some insects moult as few as three or four times during their life,
while others may moult 50 times or more. There are dangers in moulting
because, for a short time, the protective function of the cuticle is compro-
mised. Moulting is a complex process controlled by hormones secreted by
the brain and neuroendocrine glands, which act on the cells of the epider-
apolysis mis and on the nervous system. Moulting involves two major processes. First,
the separation of the old and the old cuticle separates from the underlying epidermal cells. This is called
new cuticles apolysis.
ecdysis Between this and ecdysis (the action of shedding the old cuticle), unscle-
the last stage of moulting
being the casting or sloughing
rotized parts of the old cuticle are digested and the new cuticle is formed (see
of the old cuticle figure below).
Stages in moulting
1. When moulting begins, the cells of the epidermis draw back from the in-
ner surface of the old cuticle and increase in number, or, in some cases, in
size. The new epidermal surface, having a greater area, is wrinkled. This is
important as the surface area of the epidermis ultimately determines the
surface area of the cuticle it will secrete.
2. Next, moulting fluid, a mixture of protein- and chitin-dissolving enzymes,
is secreted into the space between the epidermis and the old cuticle. The
enzymes are not activated until the epidermis has secreted the new cuti-
culin layer (inner and outer epicuticle). The outer epicuticle is made first,
then the inner epicuticle, with its closely packed wax molecules. This layer
will protect the growing procuticle beneath.
3. Once activated, the enzymes of the moulting fluid digest the old unscle-
rotized endocuticle, but do not affect the old sclerotized exocuticle. The
digested material, comprising the bulk of the old cuticle, is reabsorbed.
4. The epidermal cells secrete the procuticle under the new cuticulin layer.
The production of new cuticle may take place for many days and, in some
species, endocuticle may continue to be produced many days after the old
cuticle is shed.
5. When the old endocuticle has been fully digested and the new procu-
ticle has formed, ecdysis occurs. As a result of muscular activity and/or
increased haemolymph pressure (caused by the insect swallowing air or
water), the old cuticle, now composed only of the thin epicuticle and the
brittle exocuticle beneath, fractures along lines of weakness called sutures.
The main lines of weakness occur along the dorsal midline of the body and
as a Y-shaped suture on the head.
6. Once free of the old cuticle (which includes the lining of the fore- and
hindgut and the tracheae), the insect must expand the new, soft, wrinkled
cuticle before the processes of sclerotization irreversibly change the outer
layer of procuticle into exocuticle. In the case of insects with grub-like
larvae, only the head capsule may be sclerotized and most of the cuticle
remains soft and membranous. If the moult has been from the last imma-
ture stage to the adult of a winged species, the soft, crumpled wings must
be fully expanded.
7. Soon after moulting, dermal glands associated with the epidermis secrete
a cement-like substance, which travels to the surface through large dermal
gland ducts and then covers the surface of the epicuticular wax.
(a) Mature cuticle (b) Apolysis

Epicuticle
Exocuticle
Endocuticle
Exuvial space
Oenocyte–maximal Epidermis
Dormal gland Oenocyte development
(c) New epicuticle produced (d) Endocuticle digested Endocuticle–

partially digested
Molting fluid–
Molting fluid– active
inactive Ecdysial membrane
New epicuticle New procuticle
Oenocyte–
discharged
(e) Molting fluid resorbed (f) New cuticle after ecdysis
Old exocuticle
Ecdysial
membrane
Undifferentiated
procuticle
The stages of moulting.
The nervous system and the blood–brain barrier

Most animals have similar kinds of internal systems: a digestive system, a re-
productive system, a circulatory system, and so on. Of primary importance,
the nervous system integrates information about the environment outside
the body and the physiological conditions inside. The internal organs of in-
sects are constantly immersed in haemolymph, whose composition varies
with food intake and environmental conditions. However, to work properly
the nervous system needs an environment that is chemically stable within a
narrow tolerance band. Other tissues can cope quite well with fluctuations
in osmotic pressure and ionic concentrations, and normal homeostatic pro-

cesses are able to maintain conditions within acceptable limits. A conspicuous
feature of the CNS of insects and other terrestrial arthropods (and verte-
brates) is the presence of a clearly defined blood–brain barrier. The nervous
system needs a private pool, not communal bathing.
The insect CNS The CNS of an insect is made up of a dorsally located brain
connected on two sides to a large ventral ganglion lying beneath the oesopha-
gus (the suboesophageal ganglion). From the suboesophageal ganglion, a pair
of stout connectives (the paired ventral nerve cord) run backwards along the
body cavity and join a series of thoracic and abdominal ganglia.
The brain, ganglia, ventral nerve cords, and the major peripheral nerves
are sheathed by a supportive, corset-like layer of connective tissue called
the neural lamella (or neurolemma) inside which the perineurium, made up
of glial cells, acts as the blood–brain barrier (see Figure below). The de-
gree to which this sheath reaches down the peripheral nerves varies, but,
even beyond it, some sort of barrier layer exists in the form of accessory
cells. Glial cells have different sorts of connections or junctions with the
other cells around them. Some of these junctions are adhesive and struc-
tural, bonding the glial cells in the blood–brain barrier to each other and
to the neural lamella above. Neighbouring glial cells communicate metabol-
ically by means of gap junctions, but the most important sort of junction
between glial cells in the perineurium are tight junctions that prevent ions and
molecules in the haemolymph from diffusing in and out of the CNS. These
tight junctions ionically isolate the fluid in the tiny extracellular ‘private pool’
immediately surrounding the CNS from the rest of the haemolymph. Active
transport mechanisms pump ions in and out of the CNS to keep the ionic
concentrations of this fluid constant. The nervous system can thus function
efficiently at all times, unaffected by fluctuations in ionic concentrations in
the haemolymph. Tracheae for gaseous exchange penetrate deep inside the
CNS and branch so extensively that the finest branches, the blind-ending
tracheoles, are never located more than 5 µm from any neuron.
Within the CNS, other sorts of glial cells, which greatly outnumber nerve
cells, intimately wrap each nerve cell. These glial cells electrically insulate
the nerve cells from each other, play an important role in the development
of the nervous system, and provide metabolic support. Imagine the modern
world without electrical insulation. Very careful routing of bare wires would
be needed to avoid the continual shorting-out of circuits. If it were not for
the insulating properties of the glial cells, electrical signals passing through
the million or so neurons that make up an insect’s nervous system would be
grossly degraded.
A blood–brain barrier is found in other terrestrial arthropod groups, for
instance, scorpions and spiders, whereas crustaceans only have a partial bar-
rier. Myriapods do have a barrier of some sort, but there is no evidence yet
that it is especially ion-tight. Though of variable complexity within insects, the
possession of a highly efficient blood–brain barrier is a major contributory
factor to their huge radiation on land and in the air, allowing them to
successfully colonize even the driest and most inhospitable habitats.
Ion–tight junctions
‘Rind’ of nerve between glial cells
cell bodies
Neural lamella
(connective tissue sheath)
A'
Perineurium (blood-brain
barrier)
Fibre Nerve cell body

tracts
Neuropile (core of nerve cell
Part of
axons + dendrites)
CNS
Section through ganglion at A'
(i) (ii) (iii)

Diagram of the blood–brain
barrier (perineurium).
Flight
The origin of wings The power of flight, another major chapter in the suc-
cess story of insects, would have never been possible without lightweight,
rigid cuticle. While there might be arguments about exactly what sort of
arthropod first walked over dry land, there is no doubt that insects were
the first animals to take to the air. Insects evolved wings in the Carbonif-
erous around 300 mya, and today, only a minute percentage of the world’s
extant insect species belong to the orders (Archaeognatha and Zygentoma)
that are ancestrally wingless. Insects with developed wings appear relatively
suddenly in the fossil record, and as there are no fossils that show intermediate
stages, there is a great deal of speculation as to how they might have evolved.
One additional source of controversy is the debate around the purpose of
the first protowings. One view has been that outgrowths from the top of the
thorax would have enabled insects to have a longer, more stable glide path.
This ‘paranotal theory’ envisages early insects climbing plants and launching
themselves into the air as a more efficient, and perhaps safer, means of getting
around or avoiding predators. These lobes might have had other functions,
and experimental evidence suggests that, when small, they would have had
a significant thermoregulatory function. Other ideas have involved small in-
sects being carried around in air currents or larger insects jumping from the
ground and taking to the air. There is a major problem with ideas that involve
paranotal lobes and gliding, which is that, at some point in time, these fixed
structures need to have developed hinge mechanisms and muscle systems to
make them flap up and down. It is much easier to imagine structures that
were already hinged, tracheated, and muscled fulfilling some prior function
and then eventually taking on an aerodynamic role. The ‘endite–exite theory’
suggests that protowings developed from the fusion of inner and outer ap-
endite
a lobe of a leg segment that is
pendages (endites and exites, respectively) of basal leg segments, which were
directed inwards towards the already articulated and might have been under some sort of muscular control
midline of the body (see Figure below).
exite The latest phylogenomic and developmental studies, as well as a critical
a lobe of a leg segment that is
directed outwards away from
reinterpretation of wing development in fossil insects, suggest that the ori-
the midline of the body gin of wings is more nuanced than previously thought. A dual model, which
combines the contribution of both thoracic (paranotal) and leg structures for
Notum
Proto wing
Epicoxa
Expicoxa
Proto wing
Subcoxa
Exites Femur
Sternum
Coxa Tibia
Prefemur
Trochanter
Tarsus
Paranotal Theory Endite–Exite Theory

Comparison of paranotal
theory and endite–exite Wing develops from Wing develops from exites
theory. outgrowths of nota of primitive leg segments
the development of wings, is now the most strongly supported theory. Dur-
ing their development, insect wings express a set of genes that otherwise form
tergal (the dorsal surface of the thorax) and leg structures. Importantly, these
sets of genes exist in Archaeognatha and Zygentoma, insects that predate the
evolution of wings, as well as crustaceans, which demonstrates that they are
part of two distinct ancient sets of gene networks involved in the formation of
the dorsal body wall and appendages, respectively. The dual model suggests
that these two distinct regulatory networks at some point collaborated to form
an entirely novel structure: the insect wing. Indeed, in insects with incom-
plete metamorphosis, wings originate from primordial cells that are part of
primitive leg segments, which, during early development, move upwards to
the tergum. They then receive a signal from thoracic genes and become flat-
tened and expand outwards to form early wing pads. Also, the direct muscles
that are involved in the movement and articulation of the wings are originally
leg muscles, further highlighting the participation of the latter in wing for-
mation. Future developmental studies, and perhaps new fossil data, will help
clarify the complex origins and timing of insect wings. However, we might
never find the exact developmental intermediate steps and the evolutionary
pressures that led to the evolution of protowings.
Flight certainly allowed insects to disperse, to colonize new habitats, and

to escape from their enemies and the vagaries of the natural environment.
Which of these aspects might have been of primary significance in the
evolution of wings is a matter of debate, but predator avoidance is an ex-
cellent candidate—life in the Palaeozoic was dangerous! Pre-eminent among
predator avoidance mechanisms must have been increasingly sophisticated
early-warning systems composed of delicate vibration- or wind-sensing hairs.
Abdominal cerci coupled to giant interneurons running to thoracic ganglia
and the brain are common in some insect orders today and also existed in
early insects (in most fossils, of course, the neural connections are educated
conjecture). These systems elicit an escape response of rapid running or
jumping. It has been suggested that this escape response, development of pro-
towings, and a gradual rewiring of thoracic and abdominal ganglia might have
been the route by which insects achieved powered flight so quickly. It is easy
to see how being airborne for longer and/or being further away from danger
would have conferred a major selective advantage.
Viscosity As things get smaller, the effect of the density of the medium they
are in becomes more important. Viscous forces become greater than the iner-
tial forces. The Reynolds number is a ratio of these forces. At high Reynolds
numbers (100 and above), flow of liquids or gases across solid structures is
turbulent and chaotic. At Reynolds numbers of 10 and below, flow is laminar
The wings of a fairyfly.
(follows the surface contours). At very low Reynolds numbers, the air to a
very small insect is much more like a liquid than a gas, and not surprisingly,
the wings of minute insects differ greatly from their much larger relatives. The
wings are reduced to slender, hair-fringed paddle-like struts with which the
tiny insect ‘rows’ through the relatively sticky air (see Figure).
Dragonfly Pleural Notal

Notal process hinge Pleural
process
hinge Wing
Point of action
Dorsal longitudinal of basalar muscle
muscle
Dorso-ventral
Dorso-ventral muscle–
muscle– Basalar relaxed
contracted muscle–
Basalar muscle–
relaxed
contracted
Fly
Tergum–depressed Tergum–raised
Notal Pleural Notal
hinge process hinge Pleural
process
Dorsal longitudinal Dorsal longitudinal

Dorso-ventral muscle–contracted
muscle–relaxed muscle–
contracted
Dorso-ventral
muscle–
relaxed
Diagram of two types of flight
motor. Dragonfly: direct
muscle for downstroke—
neurogenic rhythm. Fly:
indirect muscle for
downstroke—myogenic
rhythm.
Flight mechanics Flying is a key adaptation of insects, but as insects get

smaller, the faster they have to flap their wings to fly. There are limits to the
rate at which muscles can contract to bring about wing movement. The lim-
its are not set by the muscles’ ability to contract but by the inability of motor
neurons to fire above a certain rate. In large insects with long wings, the flight
muscle can be attached directly to the wing bases and speed of firing the motor
neurons is sufficient to flap the wings at a suitable speed. The rhythm pro-
duced by the ‘flight motor’ is termed neurogenic. Smaller insects have smaller
wings and the frequency at which they need to beat is far above the speed at
which motor neurons can fire. This problem has been solved in two ways.
Firstly, indirect flight muscles are attached to a highly elastic, box-like tho-
rax. Distortion of the thorax and the way in which the wings are joined and
articulated to the latter generate the up and down stroke, as well as the angle
of attack. Secondly, the flight muscles can be asynchronous and react rapidly
to being stretched by contracting. The net result is that a high wing-beat fre-
quency can be produced without the need for a similar firing rate in the flight
muscle motor neurons. The rhythm produced by this type of flight motor is
termed myogenic (see figure above).
We now have a clearer understanding of the way in which insect wings
generate lift. For many years, insect flight was explained by comparing in-
sect wings to simplistic models such as fixed-wing aerofoils and propellers.
This led to confusion and to patently absurd statements such as ‘according
to the laws of aerodynamics, bumble bees should not be able to fly’. In real-
ity, bees fly very well; the problem has been how additional, unpredicted lift
is generated. Research has demonstrated that the leading edge of an insect’s
wing generates a strong helical airflow along its length during the downstroke.
Vortices (rotating cylinders of air) move along the length of the wings and are
shed from the wing tip (see Figure). We can now dispense with explanations
involving steady-state aerodynamics and begin the work of discovering ex-
actly how these vortices are generated and maintained along the wing. There
is still much to do when we consider the flying abilities of the hover fly or the
difficulty in swatting a house fly.
Diagram of vortices being

shed from wings.
Once wings evolved, there is no reason for them to then disappear un-
less having them incurred any kind of selective disadvantage. Secondary
flightlessness is not uncommon. Loss of wings has occurred several times in
INTERACTIONS WITH OTHER ORGANISMS 23
insects, most dramatically among parasitic species. It is likely that the major
radiation of groups, such as lice and fleas, took place only after the loss of
wings. Wing loss seems to be more common in habitats such as mountains,
deserts, and caves, which are very stable over long periods of time, or cold
and windy habitats where flight would be very costly. There is also strong ev-
idence of a trade-off between flight and fecundity. Wings and their muscles
are developmentally and energetically costly appendages. By not developing
wings, wingless individuals might be able to produce more offspring over a
longer period of time than winged individuals of the same species, as they
allocate their resources into developing larger reproductive organs instead of
wings.
Reproduction
Many insects have remarkably high reproductive rates. This, coupled with
typically short generation times, enable them to evolve faster, and to adjust
to environmental changes much more rapidly than slower-breeding animals.
Consider the folk tale about an old man who helped a vain King (the story
likely originates in India, with variants told in European and Islamic cultures).
The King had the man brought to court and told him that he could have any-
thing he wanted as a reward. The old man asked for a chessboard. The King
naturally agreed but, as he had wanted to show off his great wealth, demanded
that the man asked for something else more valuable. The old man then asked
for one grain of wheat to be put on the first square of the chessboard, two
grains on the second square, four grains on the third, and so on, until the
sixty-fourth square. Still not very pleased, the King agreed. It was not until
they got to the last few squares that the King realized the trouble he was in.
On the sixty-fourth square would rest more than 9 × 101⁸ grains—probably
more than has been cultivated in the 6,000 years of human history. It is also,
incidentally, nine times more than the estimated number of individual insects
living on Earth.
A pair of small bruchid beetles can produce about eighty offspring with
equal numbers of male and females every 21 days or so. Theoretically, the
progeny from the second generation would number 3,200, and the third gen-
eration would number 12,800 beetles. By the eighteenth generation (a mere
432 days) the beetles would number 1.4 × 102⁹ individuals and occupy a vol-
ume equal to that of the Earth. This kind of population explosion is never
seen due to a variety of regulatory factors, including weather conditions, dis-
ease, predation, and food availability, but it does illustrate the potential for
population growth. Insect outbreaks are favoured by the coincidence of two
or more of these factors, often in agricultural or other non-natural systems.
Interactions with other organisms

Apart from their obviously successful design characteristics, insects have a
vast number of interactions with a multitude of other organisms from viruses,
bacteria, and protozoa to fungi, plants, themselves, and other animals. A full
description of these interactions, which can be facultative or obligatory, sim-
ple or complex, and beneficial to one partner or both, is well outside the scope
of this volume. However, here we touch on relations involving some bacteria
and fungi, and deal with the enormous subject of plant–insect interactions in
slightly more depth.
Wolbachia are a widespread group of bacteria that are inherited
cytoplasmically and cause a range of reproductive changes in arthropods,
especially insects. Distortions of sex ratio such that mostly females are pro-
duced have been found in ladybirds, parasitic wasps, and many other taxa.
Another effect of Wolbachia and related bacterial infections is reproductive
incompatibility. If mating occurs between two individuals of strains or geo-
graphic isolates of the same or closely related species that do not carry the
same infection, embryos will not develop. The effects are typically unidi-
rectional but can be bidirectional. As yet, little is known about the actual
mechanisms by which these microbes alter the reproduction of their hosts
to their own advantage. A survey of 154 insect species in Panama found that
nearly 17% were infected with Wolbachia. Given this prevalence, and the
number of insects species in the world, Wolbachia must be very abundant
organisms, and it is likely (although not yet conclusively proved) that they
are a significant potential mechanism for speciation in insects by preventing
the reproduction of otherwise compatible populations.
A widespread phenomenon in insects is the possession of symbiotic,
non-pathogenic bacteria, protozoan, yeasts, or fungi. In some insects, these
organisms are extracellular and may be found in the gut epithelium or free
inside the gut. A much more intimate association and one pointing to co-
mycetocytes
special cells that permanently
evolution, concerns intracellular bacterial symbionts permanently housed
house intracellular bacterial inside special cells called mycetocytes, which are usually found within the
symbionts fat body.
fat body These symbionts are thought to have evolved from extracellular bacte-
an aggregation of metabolic
and storage cells in the
ria. Mycetocytes may be grouped into bodies or organs called mycetomes.
haemocoel The presence of symbionts is strongly associated with insects whose diet is
poor or lacking in particular nutrients, such as blood-feeding species (e.g.
cimicid bugs and lice), wood-feeding beetles (e.g. cerambycid beetles and
woodworm), and sap-sucking bugs (e.g. planthoppers and aphids). In the case
of bacterial symbionts, studies using antibiotics have shown that the insects
benefit greatly, as the former metabolize amino acids and vitamins of essential
importance to the latter. Extracellular symbionts are transferred from gen-
eration to generation by the young eating either the adults’ excreta or their
own eggshells, which have been daubed with symbionts. Intracellular sym-
bionts are transferred by transovarial transmission, that is, internally from
the mycetocytes to the ovaries, and then to the eggs.
Insect–fungal interactions are numerous and varied. Fungi are eaten di-
rectly by many insects; leaf-cutter ants and termites culture fungal species as
food. Wood-boring beetles, such as bark beetles (Scolytidae) and ambrosia
beetles (Platypodidae), carry fungal species with which they infect dead or
dying wood. The growth of the fungus ‘pre-digests’ the wood, making it more
edible by the beetles and their larvae. Female Sirex wood wasps (Siricidae)
carry spores of rot-producing fungus in their reproductive tract, which are
passed into the wood at oviposition. Although the cuticle is an effective bar-
rier to most species, at least 700 known species of entomopathogenic fungi
are able to penetrate its defensive layers by being able to produce a number of
cuticle-degrading proteinase, chitinase, and lipase enzymes. Besides killing
and consuming insects, many fungal pathogens have interesting effects on
their hosts. Entomopthora muscae, for example, infects house flies and yel-
low dung flies and kills very large numbers annually. Male house flies are
sexually attracted to fungus-infected corpses, perhaps because the swollen
abdomen of the infected flies might look like a fecund female with which
they can mate. Oddly though, the males prefer fungus-infected corpses to
similarly sized non-infected corpses, so there must be more to the story. In
yellow dung flies the Entomopthora fungus changes the perching behaviour
of the fly such that it dies in a suitable elevated position to ensure maximum
spread of the released fungal spores. Within the Basidiomycota, species of a
large genus, Septobasidium, are found only in association with armoured scale
insects (Diaspididae), and, unlike many insect–fungus relationships, the in-
teraction between the fungus and the scale insect is mutually beneficial. The
fungus grows on the bark of trees and shrubs, such as citrus and tea, where
it forms structured colonies, known as ‘houses’, inside which the scale insects
live, feed, and reproduce. Only a certain fraction of the scale insects become
infected by the fungus, and hyphae growing from their bodies join other hy-
phae and transfer nutrients from the fluids of the insect’s body cavity to the
main body of the fungus. The infected scale insects do not become adult or re-
produce, although they may actually live longer than non-infected scales. The
fungus receives food and the uninfected scale insects benefit from protection
from the weather and natural enemies.
As more than half of all insect species are herbivorous, their relationship
with plants must be considered the biggest single type of biological inter-
action on this planet. Initially saprophagous, some early insects might have
avoided competition and predation by moving up to plants to feed on dead
or decaying parts, or nutrient-rich structures. Insects faced several problems
along the evolutionary path to become true herbivores. One of the difficulties
of feeding on aerial parts of plants is that holding on requires special adapta-
tions. Living above ground level also increases the risk of desiccation. Of these
two obstacles, the first was overcome by the evolution of specialized tarsal and
pretarsal structures. Sharp claws and adhesive hair pads allow modern leaf-
feeding insects to cling to the shiniest leaves. Waxy cuticle, spiracular control,
and highly efficient physiological regulation of water balance, together with
the ability to use microclimatic variation in relative humidity, has allowed
insects to tolerate very dry environments. Some herbivorous species have
avoided desiccation problems by feeding internally as leaf miners, stem bor-
ers, or gall formers. The biggest problem of being herbivorous stems from the
fact that plant tissue is not very nutritious, having low protein, sterol, and vi-
tamin contents. Although some spring growth may, for short periods, have
higher levels of nitrogen, the dry weight nitrogen content of most plant tissue
lies at 3–4%. Leaf-chewing insects with dry weight nitrogen contents of 7–

15% have a much harder time than their carnivorous cousins. Sap feeders
fare even worse because phloem sap has a dry weight nitrogen content of no
more than 0.6%, and xylem sap of no more than 0.1%, and mostly less than
0.01%. It should be remembered that the total nitrogen content of plant tissue
is higher than the soluble nitrogen content, that which is actually available to
herbivorous insects. There are a number of solutions to a nutrient-poor diet.
Insects, such as sap-sucking bugs, can consume large amounts of liquid food
and void the excess carbohydrate present. Insects can be very selective in their
feeding, attacking only the most nutritious parts and timing their life cycles to
coincide with rich resources. They can develop slowly or use symbiotic micro-
organisms to boost levels of essential nutrients. Cicadas, whose nymphs feed
on xylem sap from roots, are famous for their long development: species can
take 5–17 years to develop.
The relationship between herbivorous insects and the plants that they eat is
often termed co-evolutionary, and what we see today is the result of an ‘arms
race’ that began in the late Devonian or early Carboniferous. Adaptations in
one species to overcome defence mechanisms evolved by another species are
countered by novel defences in the first, and so on. Working on the principle
of damage limitation, plants have evolved a number of defences to herbivores.
Physical properties, such as shiny or waxy surfaces, hairs, and sharp or sticky
trichomes make holding on difficult or dangerous, but are not much defence
against internal feeders, such as miners or borers, once they are inside. In-
corporated silica grains, or leaf toughness caused by high levels of lignin or
cellulose, will deter most chewers but will not affect sap suckers much.
Of more interest are the chemical defences that plants have evolved to de-
ter or poison insects. There are a very large number of these secondary plant
compounds or allelochemicals, and more are being discovered. They fall into
a number of classes, of which alkaloids, terpenoids, and flavonoids are the
biggest, with more than 10,000 different active compounds between them.
We should be grateful for the huge diversity of insect–plant interactions, be-
cause many of the compounds we value most highly, and may be addicted
to, such as nicotine, caffeine, and tannins, as well as many useful drugs, are
nothing more than plant-produced anti-insect compounds.
It is now clear that elegant tritrophic relationships have evolved between
plants, insect herbivores, and the wasps that parasitize them. Locating suit-
able resources, whether plant tissue in the case of the herbivore or insect tissue
in the case of the parasitoid, is of primary importance. Plant volatile chem-
icals may act as attractants or repellents allowing insects to orientate from
some distance. Plants damaged by herbivores can produce volatile chemi-
cals that attract natural enemies—predators and parasitoids. One study using
corn seedlings and caterpillars of the beet armyworm showed that certain oral
secretions of the caterpillars act as triggers for the release of plant volatiles—
mechanical damage alone does not have the same effect. The production of
these volatiles, induced through a pathway known as the octadecanoid path-
way, can also be stimulated by applying jasmonic acid (jasmonates are a class
of fatty acids). It remains unclear as to why herbivores should stimulate clear
signals that alert enemies to their presence, but chemicals such as that found
in the oral secretions of the armyworm caterpillars must be advantageous
to the caterpillars in other ways. Natural selection acting on the plants and
the response of natural enemies to volatiles has produced a winning strat-
egy (as far as the plants are concerned). The stage may now be set for ‘stealth
herbivores’ to evolve.
Insect–plant relationships are not one sided. Mutualisms abound, and at
the other end of the spectrum, there are carnivorous plants, which trap and
digest insects. Even here, things are not clear cut; some pitcher plants harbour
many larvae of many flies that live unharmed on the plant’s digestive juices. It
turns out that these larvae feed on the thin soup of water and decaying insects
and provide the plant with nutrients in a readily assimilable form.
Even the shortest discussion of insect and plants should deal with polli-
nation. Using insects as ‘flying genitalia’ is a marked improvement over wind
pollination. Winds are unpredictable in speed and direction, there is much
wastage of pollen, and rare plants would stand little chance of ever being
fertilized. Insects may have visited plants to feed on spores as early as the De-
vonian, and it is likely that the first step towards true pollination came about
incidentally as a result of herbivorous feeding. Flying insects would have car-
ried spores attached to their bodies, and any plant that, by chance, produced
a structure, odour, or some other cue that attracted insects would be at a se-
lective advantage. Plants evolved structures, such as carpels, to protect their
seeds from being eaten by herbivores, while at the same time ensuring that
pollinators could locate them and would continue to visit through the pro-
duction of rewards such as nectar. Flying insects need a carbohydrate fuel for
energy and nectar is typically rich in sucrose, glucose, and fructose. Pollen is
a protein-rich food, but plants that are pollinated by insects unable to utilize
pollen directly (e.g. butterflies) produce nectar with amino acids and proteins.
Since the first flowering plants appeared in the Cretaceous, co-evolutionary
processes have produced a wealth of flower size, shape, and colour, and the
appearance of groups of insects that are tied to them, to a greater or lesser de-
gree. Many groups of insects visit flowers, in particular, beetles, flies, moths
and butterflies, and bees. Mutualistic adaptations are numerous. Plants that
evolved long, tubed flowers were tracked by species that evolved similarly
elongated mouthparts to cope. In this manner, non-specialist flower visitors
are defeated to the mutual advantage of plant and pollinator. The end result
of evolution in plants that ensure extreme pollinator constancy is that tightly
co-evolved groups of plants and pollinators occur. Fig wasps (Agaonidae)
and fig trees are totally dependent on each other for survival. The trees rely
on the wasps for pollination and the wasps, which have complex life cycles,
can only reproduce inside figs (see hymenopteran superfamilies). Bees (see
Apoidea) are considered the most sophisticated plant pollinators and feed
their young on a diet of nectar and pollen. Occasionally, the plants seem
to benefit more than the pollinators do. Orchids of the genus Ophrys have
evolved flowers that mimic the shape and sexual odour of certain female
bees. Males attempting to copulate with the flowers get a pollinium or pollen
packet stuck to their backs. When the bee is pseudo-copulating with another
flower, the pollen in transferred. Very complex relationships exist between

certain South American orchids and about 200 species of orchid bees (Apidae:
Apinae: Euglossini). Only the males of these long-tongued, fast-flying bees
visit the orchids. The bees collect the orchid’s fragrance using pads on their
front legs, and the scented oils are then passed back to the rear tibiae, which
are filled with spongy storage tissue. While this is occurring, the bee gets a
pollinium stuck to a part of its body. The exact location of the pollinium is im-
portant, and several closely related species of orchids might be reproductively
isolated by this simple mechanism. It is not yet clear exactly how the bees
use the scents, as they attract only males; perhaps females are attracted to
aggregations of males and then select a mate on other criteria.
Insect structure
For all their diversity, insects are remarkably similar in overall design, both
inside and out. The basic plan has been reworked many times over to produce
a multitude of variants based on three main tagmata or body regions: the
head, thorax, and abdomen.
The head is formed from six fused segments, and carries the compound
ocelli eyes, secondary, light-receptive organs called ocelli, the antennae, and the
simple, light-receptive, mouthparts.
non-image forming organs on
the head of many insects The mouthparts may be modified according to diet, allowing the sucking
or lapping of liquids, or the chewing and grinding of solid foods. The tho-
rax is the powerhouse of the insect and is made up of three segments, each
bearing a pair of legs. The posterior two thoracic segments usually each bear
a pair of wings. The legs, which are each made up of a number of articulated
components—coxa, trochanter, femur, tibia, and tarsus—can be greatly mod-
ified to serve a variety of functions. The abdomen, which is usually made up of
eleven segments, contains the digestive and reproductive organs, and carries
the internal and external genitalia. The latter are often species specific and,
therefore, of great value to taxonomists.
On the inside, the major systems are the nervous system, the circulatory
system, the digestive system, the respiratory system, and the reproductive
system.
Sense organs
From the CNS, described earlier, arises the peripheral nervous system. This
comprises sensory nerves bringing information from sensory receptors into
the CNS, motor nerves that control the action of muscles, and the stomato-
gastric nervous system, which innervates the gut. The compound eyes are the
main visual sense organs of adult insects. The eyes are made up of individual
ommatidia light receptive units called ommatidia, and there may be anything from only
individual light receptive units one or two in some ants to more than 10,000 in dragonflies.
making up the compound
eyes of insects Each ommatidium comprises a clear lens system and light-sensitive ret-
inula cells. In day-flying insects, the image received by the eye as a whole is
INSECT STRUCTURE 29
made up of a mosaic of spots of differing light intensity from all the sepa-
rate ommatidia. The more ommatidia present, the greater will be the insects’
visual acuity. The eyes of night- or dusk-flying insects have a different internal
construction and sacrifice visual acuity in favour of light sensitivity. Colour
vision has been shown to occur in all orders of insects, and some can detect
the plane of polarized light. In general, insects ‘see better’ at the blue end of
the spectrum than at the red end, and, in some groups, the range of sensitivity
carries on into the ultraviolet.
Hairs on the cuticle surface are responsive to vibrations, touch, and sound.
Special hearing structures called tympanal organs may be present on various
parts of the body (legs, wings, thorax, abdomen, and antennae). Depending
on the species, these organs are responsive to sound frequencies ranging from
less than 100 Hz to 240 kHz. Hair beds and arrays of small hair-like sensilla on
the cuticle at joints of legs and segments provide information about the rela-
tive position of body parts. Strain gauges called campaniform sensilla respond
to stresses in the cuticle, and internally, stretch receptors sense distension of
muscles and parts of the digestive tract. Chemical sense organs or chemore-
ceptors are present on the mouthparts, antennae, tarsi, and other parts of the
body. Insects smell by means of olfactory sensilla. These structures are found
mainly on the antennae and can be present in very large numbers to detect
remarkably low odour concentrations. In some cockroaches, there may be
up to 100,000 of these sensilla. Many insects have temperature and humidity
sensors, and some can detect magnetic fields and infrared radiation. Closely
associated with the nervous system is a complex array of hormonal organs.
Together, the nervous and hormonal systems regulate insect behaviour and
physiology.
Circulatory system
The circulatory system is quite simple. The internal organs are bathed in
haemolymph, which is moved around the body cavity by means of a com-
bined heart and aorta lying just under the dorsal body surface. This dorsal
organ has a number of openings called ostia along its length. In very sim-
ple terms, body fluids enter through the ostia and, as muscular contractions
proceed from back to front, the fluids are passed towards the head end.
The haemolymph does not move back through the ostia because they act
as one-way valves. The main function of the haemolymph is concerned with
nutrient and waste transport, although it also has a role in temperature con-
trol, protection against parasites and diseases, and as a hydrostatic skeletal
element.
Digestive system
The digestive system of insects is essentially a long, open-ended tube with
special regions for grinding and storing food, the production of enzymes,
and the absorption of nutrients. The fore- and hindguts, being invagina-
tions of the epidermis, are lined with cuticle. The unlined midgut is the main
area for enzyme production and nutrient absorption. The nutrients (carbo-
hydrate, fats, proteins, vitamins, etc.) needed by insects are essentially the
same as for all other animals. The midgut epithelium of insects secretes a
mucopolysaccharide layer called the peritrophic membrane, which envelops
the food as it enters from the foregut. The membrane prevents bacteria, and
perhaps parasites, from contacting the epithelial cells directly. However, it
mainly works as a filter to prevent the passage of large molecules and food
particles through its pores. Digestion takes place in a series of stages. The
first stage occurs inside the peritrophic membrane, where large molecules are
broken to smaller ones. The second stage occurs in the space between the per-
itrophic membrane and the gut wall, where small molecules are broken down.
Lastly, final digestion and absorption of nutrients takes place on the surface
of the midgut epithelial cells. The peritrophic membrane is produced con-
tinuously and eventually passes out of the anus with food residues. In some
beetles, it continues to be produced after feeding has stopped just prior to pu-
pation. It is issued from the anus as a thread-like ribbon and the larva uses it
to make a cocoon. In most animals, the removal of the toxic waste products
of metabolism requires their solution in water. For small terrestrial animals
like insects, loss of water is critical, and a different system is employed. At
the junction of the mid- and hindgut, there are a number of long, slender,
blind-ended ducts called Malpighian tubules. These tubules extract the waste
products from the haemolymph in which all the internal organs are bathed.
In effect, they continually produce the insect’s urine, which is moved to the
hind gut. Valuable water and salts are reabsorbed by the hindgut and rec-
tum, and the excrement is voided mainly as non-toxic and insoluble uric acid
granules.
Respiratory system
The respiratory system of insects takes the form of an incredible network of
branched tubes, which take gases directly to the cells of the body. The open-
ings of this tracheal system, called spiracles, are located on the sides of each
abdominal segment and on the thorax. No insect has more than ten pairs of
spiracles, and the number is often reduced. In the aquatic nymphs of dragon-
flies, damselflies, mayflies, and stoneflies, which have tracheal gills, there are
no functional spiracles. Spiracles can be closed by means of valves to reduce
water loss. The main tracheae are spirally thickened like the flexible hose of
a vacuum cleaner, to prevent them from collapsing. The tracheae are often
swollen into air sacs, and insects can set up a flow of air through their tra-
cheal system by means of body movements and the rhythmic flattening of
these sacs. The tracheae branch continuously, getting finer and finer, until
they become very fine intracellular tubes known as tracheoles, which are < 1
µm in diameter. Gases are exchanged by diffusion of oxygen into cells and
carbon dioxide out of the cells. Several studies have shown that caterpillars
have a special system analogous to the lungs of vertebrates (see Lepidoptera).
Aquatic insect species show many interesting adaptations to life underwater,
such as gills, air stores, and breathing siphons.
CONCLUSION 31
Reproductive system
The reproductive system of insects consists of a pair of ovaries in the female
and a pair of testes in the male. Each ovary is joined by its own lateral oviduct
to a main or common oviduct. The common oviduct leads to the single vagina
or genital chamber. Each testis is joined by means of a duct, known as the vas
deferens, to a common ejaculatory duct. This common duct takes the sperm
to the external genitalia, of which the aedeagus (or penis) is the most impor-
tant component. Eggs are fertilized by sperms stored inside the body of the
female. The fertilization of eggs by sperm is internal and requires the male
to introduce sperm by means of an aedeagus or penis via the female’s vagina.
Sperm is deposited in or near a sperm-storage organ, known as the spermath-
eca, and is subsequently used to fertilize the eggs. In some insects, a single
mating will provide the female with all the sperm she will ever need, while in
others, sperm cannot be stored indefinitely and the insects mate repeatedly
to maintain a fresh supply.
Conclusion
Before we end this brief introduction, we would like to answer a question of-
ten asked by students: Why are there no truly marine insects? To our eyes,
the sea can seem a dangerous and inhospitable place, but reasonably constant
temperatures (certainly less of a range than found on land), coupled with
fairly constant ionic concentrations, make the seas (physiologically speak-
ing) about as benign an environment as there is on Earth. Life in freshwater
is much more difficult osmotically, yet only a small fraction of 1% of all in-
sect species live in marine habitats. Many insect orders contain a few species
with marine associations, but the majority of species that spend at least part
of their life cycle around the seashore, in brackish water as well as on the sur-
face of the ocean, belong to the orders Hemiptera, Coleoptera, Diptera, and
Trichoptera, among others. A few species of small predatory bugs with the
evocative common name of Coral-treaders belong to the exclusively southern
hemisphere family, the Hermatobatidae. The detailed biology of only one or
two species is known, but they hide in coral and rock crevices and only emerge
to feed at low tide. The most remarkable insects to have colonized saltwater
are a few species of caddisfly species, whose larval and pupal stages are found
in rock pools along the coast of New Zealand and south-east Australia (see
Trichoptera).
Of the various types of marine environment, the least populated by in-
sects are pelagic and coastal water habitats. Members of the hemipteran
family Gerridae (pond skaters) are predacious bugs adapted to life on the
surface of water. The vast majority live on freshwater, but five genera con-
tain marine species. Of special interest are the 50 or so wingless species
of ocean strider belonging to the genus Halobates. Most of these are con-
fined to mangroves and coastal regions, but at least five species are only
found far from land on the open oceans of the southern hemisphere. They
are pelagic in as much as they live independently of land, but they live on
saltwater, not in it. Lice in the anopluran family Echinophthiriidae are par-
asitic to seals, walruses, and sea lions. Although reproduction takes place
when their hosts are on land, the lice are well adapted to long periods (many
weeks or months) of feeding on their hosts while at sea. The lice are verti-
cally transmitted (from mother to offspring) before the pups go to sea and,
like all parasitic lice, there is no stage where they are independent of their
hosts.
The greatest numbers of marine-associated insects are found in intertidal
zones, salt marshes, mangroves, and mudflats, where they are phytophagous
on salt-tolerant plants, predators, or scavengers. However, these inhospitable
habitats are as much extensions of the terrestrial environment as of the ma-
rine. The question ‘why don’t insects live in the sea?’ is easily answered. There
is no physiological reason why insects could not live in saline water; in fact,
it could make things a bit easier. Simply, most of the features that have made
insects so successful on land would be of no great advantage relative to ex-
isting marine arthropods (the Crustacea), whose member species already fill
every available niche. Put simply: insects just don’t have a chance.
A final word must be on biodiversity. An increasing interest in catalogu-
ing the Earth’s biological diversity made it fashionable, for a while, to talk
about larger and ever-larger numbers of undiscovered species in terrestrial
and marine habitats. Some of the estimates for the total number of extant in-
sect species, based on tropical forest samples, even reached the 100 million
mark. Larger estimates might seem more impressive, but there came a point at
which the brakes were applied. A number of studies suggest that there might
not actually be quite as many species as originally thought. Undoubtedly,
there are more species not described than described, but the figure proba-
bly lies somewhere in the range of 3–8 million, not 30–80 million. All species
do not have an equal chance of being described. Small, rare species are much
less likely to be named than larger common species. Not all things collected
are ever described, and, if they are, the time from collection to description is
getting longer and longer. Apart from the serious problem of the dwindling
number of trained taxonomists capable of contributing to the immense task,
or the lack of funds available to train new ones, one of the major difficulties in
biodiversity studies is synonymy. This is where a species has been described
and named more than once, sometimes by different authors and sometimes
by the same author. Taxonomic revisions have shown that, in some taxa,
species may have more than a dozen different names. Overall, it is estimated
that about 20% of all species names are not valid. Studying the patterns of
species description in different groups, especially the very large ones, would
provide interesting insights. One such study on the Geometridae, a large
and widely distributed family of moths, suggests that the actual number of
species is nowhere near the estimation of an order of magnitude greater than
the number of described species. Despite this, there are still plenty of places
around the globe where every other species collected is likely not to have been
described.
CONCLUSION 33
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origins. Current Biology 27: 263–9.
Regier, J., Shultz, J., Zwick, A., Hussey, A., Ball, B., Wetzer, R., Martin, J. W., and Cun-
ningham, C. W. (2010). Arthropod relationships revealed by phylogenomic analysis of
nuclear protein-coding sequences. Nature 463: 1079–83.
Seimann, E., Tilman, D., and Haarstad, J. (1996). Insect species diversity, abundance and
body size relationships. Nature 380: 704–06.
Simpson, S. J. and Douglas, A. E. (eds) (2014). R. F. Chapman—The insects: structure and

function, 5th ed. Cambridge University Press, Cambridge.
Walker, S. M. and Taylor, G. K. (2021). A semi-empirical model of the aerodynamics of
manoeuvring insect flight. Journal of the Royal Society Interface 18: 20210103.
SECTION 2
The insect orders

36
All six-legged animals belong to a single superclass, the Hexapoda of which

the Insecta forms the largest class (circa 1 million spp). The three non-insect
classes are the Protura (~ 800 spp.), Diplura (~ 800 spp.), and the Collem-
bola (> 9,000 spp.). These three classes are known as ‘entognathous hexapods’,
which have jaws inside an eversible pouch in the head. The figure below shows
the subdivisions of the Hexapoda, and the latest insect tree of life.
Protura
Other
Collembola Hexapoda
Diplura
Archaeognatha
Zygentoma
Odonata Palaeo-
Ephemeroptera ptera
Zoraptera
Dermaptera
Plecoptera
Orthoptera
Polyneoptera
Mantophasmatodea
Grylloblattodea
Embioptera
Phasmatodea
Mantodea
Blattodea
Isoptera
Thysanoptera
Pterygota
Paraneoptera
Insecta
Sternorrhyncha
Neoptera
Heteroptera
Coleorrhyncha
Auchenorrhyncha
Psocodea
Hymenoptera
Raphidioptera
Megaloptera
Holometabola
Neuroptera
Strepsiptera
Coleoptera
Trichoptera
Lepidoptera
Siphonaptera
Mecoptera
Diptera
Phylogeny of hexapods as
resolved from the most For practical purposes, the most commonly used level of classification
comprehensive
phylogenomic study (Misof is the order. These correspond roughly to familiar English names for dif-
et al., 2014) at the time of ferent types of insects, flies, beetles, butterflies and moths, earwigs and so
writing this book. Note that
the position of Psocodea as
on, but also include less well-known groups like bristletails or ones without
sister to the Holometabola is vernacular English names like the Strepsiptera. The 28 recognized insect or-
probably artefactual, and that ders are described here in a sequence that reflects their taxonomic grouping
termites (Isoptera) are
included within Blattodea. (i.e., Archaeognatha, Zygentoma, Palaeoptera, Neoptera, and so on). The clas-
Adapted from © Hans Pohl. sification in this book is based on the latest molecular and morphological
systematic studies, which are provided as further reading at the end of each
order.
37
Insect groups that predate the

evolution of wings
··································································································
Ancestrally wingless insects, also sometimes known collectively as the
apterygotes, do not undergo any sort of metamorphosis, that is, they are
ametabolous. The young stages resemble the adults in all respects except size
and sexual maturity and moulting continues until death. These are the most
ancestral of all insects alive today, and the only descendants of a much larger
array of wingless species that once existed. They have persisted in special-
ized cryptic microhabitats and have retained several ancestral features that
likely date back to the Carboniferous times (300 million years ago, mya).
The only two surviving orders are superficially similar in overall appearance
but are in fact independent monophyletic lineages with subtle morphological
differences.
38
Archaeognatha
(bristletails—alternative name: Microcoryphia)
Common name Bristletails Metamorphosis Ametabolous

Derivation Gk. archaeo–ancient; Distribution Worldwide
gnathos–jaw Number of families 2
Size Body length 7–25 mm Known world species 550 (0.05%)
Maxillary palp
Large compound eye
Long multi segmented

Humped thorax
antenna
Style
Style
Cercus
Central filament
Petrobius maritimus — Archaeognatha
ARCHAEOGNATHA 39
Key features
• ancestrally wingless
• elongate, cylindrical body, covered with small scales
• jumping ability
• scavengers
• in litter, soil, among coastal rocks and cliffs, some in dry regions
© Rupert Soskin
Jumping bristletails are the
most evolutionarily
primitive of all living insects.
Archaeognathans are elongate and cylindrical in cross-section with a notice-

ably humped thorax and an exoskeleton covered with scales. The head has
a pair of long, multi-segmented antennae, large compound eyes that touch
on top of the head, and three ocelli (single-faceted, simple eyes). Beneath
the elongate, seven-segmented maxillary palps there are downwards-facing
mouthparts. The jaws, which have a single or two points of articulation, oper-
ate with a vertical, auger-like action. Single-point (monocondylic) jaws were
considered as a primitive feature that defined Archaeognatha and distin-
guished them from all other insects, including the Zygentoma, all of which
have two points of articulation (dicondylic). It is now known that many Ar-
chaeognatha have dicondylic jaws as well, which pushes the appearance of
complex jaws even earlier in insect evolution. In addition to the three pairs of
thoracic legs, there are accessory appendages called styles, which support the
abdomen. These structures, which have their own muscles, are present on ab-
dominal segments 2–9 and assist when running by supporting the abdomen
over uneven or steep surfaces. There are also one or two pairs of eversible vesi-
cles located on the underside of abdominal segments 1–7, which are involved
in absorbing water and assisting with moulting. The abdomen has a pair of
multi-segmented cerci and a much longer central filament.
Bristletails in the family Machilidae are often nocturnal, eating algae,
lichens, and plant debris. Large populations can be found living in cliffs and
crevices in old stone walls. The Meinertellidae are diurnal and scavenge some-
times in very dry environments, for example, deserts and dunes. For only a
handful of species is a detailed life history well known. Petrobius maritimus,
40 SECTION 2 THE INSECT ORDERS
a common bristletail species living on rocky shores in the northern hemi-

sphere, grazes at night on lichens and algae-growing rocks and is active even
in near-freezing conditions.
Immature archaeognathans may moult up to ten times and may take up to
two years to reach sexual maturity after which they may live for another two
years. Some species have been known to moult more than 50 times during the
course of their lives and many species use their adhesive coxal vesicles to stick
themselves safely to vertical or inclined surfaces during moulting. Courtship
and mating, which is quite simple, may involve some sort of dance. Sperm
transfer is indirect but achieved in different ways. In the Meinertellidae the
male deposits a packet of sperm on a silken thread or stalk attached to the
ground during courtship. The male may manoeuvre the female into a position
such that she can pick it up. In Machilidae, sperm transfer is one step nearer to
being direct in that the male actually places sperm on the female’s ovipositor,
while in others the male deposits numerous small droplets of sperm on a silk
thread, which he holds stretched. The female picks up the sperm by means
of her ovipositor. Female bristletails lay batches of 10–15 soft, round eggs in
cracks and crevices. Good egg-laying sites may contain +100 eggs.
Despite being descendants of the first insect lineages, the evolutionary ori-
gins of archaeognathans are particularly challenging to pinpoint, particularly
due to their sparse fossil record. Putative Archaeognatha have been found as
far back as the Late Devonian (more than 380 mya). Confirmed archaeog-
nathans have been described from exceptionally preserved fossils from the
Early Triassic (240 mya), although the origin of the order is undoubtedly
much older.
Key reading
Adis, J. and Sturm, H. (1987). On the natural history and ecology of Meinertellidae.
(Archaeognatha, Insecta) from dry land and inundation forests of central Amazonia.
Amazoniana 10(2): 197–218.
Blanke, A., Machida, R., Szucsich, N.U., Wilde, F., and Misof, B. (2014). Mandibles with
two joints evolved much earlier in the history of insects: dicondyly is a synapomorphy of
bristletails, silverfish and winged insects. Systematic Entomology 40: 357–64.
Edwards, J. S. (1992). Adhesive function of coxal vesicles during ecdysis in Petrobius
brevistylis Carpenter (Archaeognatha: Machilidae). International Journal of Insect Mor-
phology and Embryology 21: 369–71.
Mendes, L. F. (2018). Biodiversity of the Thysanurans (Microcoryphia and Zygentoma). In:
Insect biodiversity: science and society, volume II. Foottit, R. G. and Adler, P. H. (eds), pp.
153–98. John Wiley & Sons Ltd., Hoboken.
Montagna, M., Haug, J., Strada, L., Haug, C., Felber, M. and Tintori, A. (2017). Central ner-
vous system and muscular bundles preserved in a 240-million-year-old giant bristletail
(Archaeognatha: Machilidae). Scientific Reports 7: 46016.
Sturm, H. (1986). Aspects of the mating behavior in the Machiloidea (Archaeognatha,
Insecta). Braunschweiger Naturkundliche Schriften 2(3): 507–18.
41
Zygentoma
(silverfish and firebrats—alternative name: Thysanura)
Common name Silverfish, firebrat Distribution Worldwide but

Derivation Gk. zyg–bridge; predominantly tropical
entoma–insect Number of families 4
Size Body length 1.4–20 mm Known world species 600 (0.05%)
Metamorphosis Ametabolous
Long multisegmented
antenna
Small compound
eye
Flattened body
Scales (Lepismatidae only)
Cercus
Central filament
Lepisma saccharina — Zygentoma
Key features
• ancestrally wingless
• elongate, flattened body, often with silvery scales
• fast running but do not jump
• scavengers
• in soil, litter, burrows, trees, ant nests, sometimes in buildings
The preference of the

firebrat (Thermobia
domestica) for high
temperatures and starchy
© Paul Brock
foods led it to become a
common resident of
bakeries.
Many people will recognize these insects as the small, shiny, silver creatures
that scuttle rapidly for shelter when disturbed after dark in a larder or bath-
room, and some species damage books and papers and can be minor pests
in kitchens. They are, however, quite diverse in shape and appearance and
occupy a greater number of habitats than buildings. They can be elongate or
oval, flattened or slightly convex, and may have body scales. Like the bristle-
tails, the head has a pair of long, multi-segmented antennae but the eyes,
when present, are small and never touch each other on top of the head. There
may be up to three ocelli, although some species have none. The maxillary
palps are shorter than those of the Archaeognatha with only five or six seg-
ments and the downwards-facing jaws, although still of a primitive design,
have two points of articulation with the head and act in the transverse plane.
Accessory walking appendages or styles may be present on abdominal seg-
ments 2–9 but usually on fewer segments (7–9). Pairs of water absorbing,
eversible vesicles usually occur on abdominal segments 2–7, although in some
species they are lacking altogether. The end of the abdomen carries three ter-
minal filaments but unlike the Archaeognatha, they are all more or less the
same length.
Zygentoma are mostly nocturnal and omnivorous scavengers and can
be found free-living in litter or under bark, underground, in caves, in the
burrows of certain mammals, or in association with ant and termite nests.
Atelura formicaria, a species found in association with ants, is able to fol-
low the pheromone trails laid down by foraging workers of Lasius species
and it has been suggested that this behaviour is a dispersal mechanism. Some
ZYGENTOMA 43
species can survive in habitats with little free water by being able to absorb
atmospheric water though their rectum.
Species of the cosmopolitan family Lepismatidae are widespread in many
habitats, including litter and in trees. Some lepismatids are almost exclusively
found in human habitations. One of these, the silverfish, Lepisma saccharina,
prefers damp microhabitats such as those found in cellars and bathrooms,
while the firebrat, Thermobia domestica, can be found living in the warm con-
ditions of kitchens and bakeries. Feeding studies on the firebrat have shown
that gut cellulases are produced by the silverfish itself and not by symbiotic
micro-organisms as in the guts of more advanced insects like the termites. The
domestic species feed on a wide range of starchy materials, such as spilled
flour, starched damp textiles and clothing, as well as silk, book bindings,
wallpaper, and photographs.
Courtship may consist of a simple, synchronized pas de deux, while one
species reproduces parthenogenetically, that is, without fertilization by a
male. Just before mating males secrete silk threads, usually between two sur-
faces, onto which they deposit sperm droplets or packets. Receptive females
take hold of the sperm using their ovipositor and draw it inside. Individuals
may live for three or four years and, as in bristletails, moulting continues until
death.
Due to their small size and elusive nature, silverfish are remarkably scarce
in the fossil record. As a consequence, their deep evolutionary origins re-
main obscure. As with Archaeognatha, reports exist of tentative, difficult-
to-identify fossils of Zygentoma from the Devonian. Confirmed relatives
of modern representatives have been found in Cretaceous amber (100–110
mya), indicating that the current families were well established long ago.
Key reading
Christian, E. (1994). Atelura formicaria (Zygentoma) follows the pheromone trail of Lasius
niger (Formicidae). Zoologischer Anzeiger 232: 213–216.
Mendes, L. F. (2018). Biodiversity of the Thysanurans (Microcoryphia and Zygentoma). In:
Insect biodiversity: science and society, volume II. Foottit, R. G. and Adler, P. H. (eds.), pp.
153–98. John Wiley & Sons Ltd., Hoboken.
Mendes, L. F. and Poinar, G. (2008). A new fossil silverfish (Zygentoma: Insecta) in Mesozoic
Burmese amber. European Journal of Soil Biology 44: 491–4.
Picchi, V. D. (1972). Parthenogenic reproduction in the silverfish Nicoletia meinerti (Thysa-
nura). Journal of the New York Entomological Society 80(1): 2–4.
Sturm, H. (1988). The mating behaviour of Thermobia domestica (Packard) (Lepismatidae,
Zygentoma, Insecta). Braunschweiger Naturkundliche Schriften 2(4): 693–712.
Sturm, H. and Bach de Roca, C. (1993). On the systematics of the Archaeognatha (Insecta).
Entomologia Generalis 18: 55–90.
Treves, D. S. and Martin, M. M. (1994). Cellulose digestion in primitive hexapods: Effect
of ingested antibiotics on gut microbial populations and gut cellulase levels in the fire-
brat Thermobia domestica (Zygentoma, Lepismatidae). Journal of Chemical Ecology 20:
2003–20
44
The winged insects

··································································································
Insects with wings had evolved by the middle of the Carboniferous and their
winged descendants make up 99.9% of all extant insect species. Wings now
as then allow rapid escape from enemies, efficient mate-finding and dispersal
to new feeding, mating, or egg-laying sites. Nevertheless, wings have been
lost or reduced in many taxa for various reasons. The most common cause
of secondary wing loss is due to the development of a specialized lifestyle. In
particular, external parasites like fleas and lice are always wingless. Species in
other orders that have parasitic relationships with other species, either living
in the nests of termites or ants or feeding on vertebrate blood, are also often
wingless or short-winged. Habitat stability over long periods, such as those
found in caves, soil, and certain montane environments is another cause of
secondary winglessness. In these cases, dispersal is no longer essential to the
survival of the species, so not having functional wings and associated muscles
is likely to increase fecundity and thereby select for winglessness. The cost of
having wings and thus the number of flightless insect species is also greater
in habitats where extremes of cold, hot, or wind prevail.
The infraclass Pterygota is divided into two unequal divisions. The
mayflies (Ephemeroptera), comprising 0.3% of all insect species, and the
dragonflies and damselflies (Odonata), comprising 0.5% of all insect species,
are unable to fold their wings back along the body. This condition is known
as palaeopterous (from Greek palaeos = old, ancient; pteron = wing). This
characteristic wing morphology has given mayflies and dragonflies the name
Palaeoptera, which assumes that the two orders are sister groups. This classi-
fication remains debated to this day, as the two orders are dissimilar in other
regards, especially the occurrence in mayflies of a winged sub-adult stage.
Some studies confirm the validity of Palaeoptera, whereas others suggest that
both orders represent their own distinct division. The last, and by far the
largest, subdivision of the Pterygota includes 99% of all insect species. This
division is known as the Neoptera (from Greek neos = new: pteron = wing)
and contains 24 orders.
45
Division Palaeoptera
··································································································································
Ephemeroptera
(mayflies)
Common name Mayflies Metamorphosis Incomplete (egg, nymph,

Derivation Gk. ephemeros–lasting a adult)
day; pteron–a wing Distribution Worldwide but especially
Size Body length 2–50 mm. temperate regions
Wingspan up to 50 mm Number of families 42
Known world species 3,666 (0.32%)
Short antenna
Front leg
Large mesothorax Large compound eye
Large, triangular
front wing
Small, rounded Soft-bodied

hind wing
Slender abdomen
Long cercus
Long tail filament
Ephemera danica — Ephemeroptera
Key features
• aquatic nymphs
• separate winged subimago and adult stages
• two pairs of wings held vertically over body at rest
• adults short-lived and non-feeding
• mating swarms often seen over water
• essential component of freshwater food chains
© George McGavin
Mayflies are unique in that

they go through two moults
once they have reached the
adult stage. Ephemera
danica, UK.
The Ephemeroptera, together with the Odonata, are the oldest groups of
winged insects on Earth today. Ephemeroptera are unique in having a pre-
adult winged stage called the subimago—they are the only insects that moult
after having developed functional wings. This habit was probably much more
common in several extinct Carboniferous and Permian orders, where imma-
ture stages had wing-like structures and moulted them throughout their lives.
In two extant mayfly families, the Polymitarcyidae and the Palingeniidae, the
role of reproductive female has been taken over by the subimago.
Mayflies are soft-bodied and relatively delicate insects with nearly cylin-
drical bodies, longish legs, and two pairs of wings that cannot be folded back
along the body. The head has a pair of short antennae, a pair of large, com-
pound eyes, and three ocelli. Adult mayflies, which are solely concerned
with the purpose of reproduction, do not feed and have very reduced, non-
functional mouthparts. The lifecycle is dominated by the nymphal stages and
adults live for a very short time, mostly much less than a day, and in a few
EPHEMEROPTERA 47
species, only a matter of minutes. All adults are fully winged, and the front
pair of wings are large and triangular, while the hind wings are smaller and
more rounded. The end of the abdomen bears a pair of elongate cerci and,
usually, a single, long tail filament between them. The order used to be divided
into two suborders: the Schistonota (sometimes called split-back mayflies)
and the Pannota (fused-back mayflies). Although this system is no longer
used, the terms Pannota–Schistonota are found in most studies (for historical
reasons), and it is useful to know them. It is easy to tell mature nymphs of the
two suborders apart. Schistonotan nymphs have their wing pads free along
the midline whereas, in pannotan nymphs, the wing pads are fused along the
midline of the body. However, the higher-level classification and phylogeny
of mayflies is complex and highly debated and will likely undergo consider-
able changes as more representatives of the order are studied with molecular
methods.
The aquatic nymphs moult many (10–50) times and can take up to
three years to reach adulthood. Mayflies can be found in aquatic habitats of
all kinds, from ditches, ponds, and lakes to slow-flowing rivers or fast-flowing
streams. Some families, for example, the Baetidae, are found at higher alti-
tudes and latitudes than any others. Different mayfly species are characteristic
of different sorts of habitats. Some species are found in still waters, some crawl
on gravely bottoms, while others live in burrows that they excavate in mud
and silt. Habitat choice is often reflected in nymphal body shape. For instance,
nymphs adapted for life in fast-flowing streams are streamlined, flattened, and
may have special adhesive gill holdfast organs.
Widespread mayfly families
Family Nymphal type: feeding Examples of genera

habits
Baetidae* Active swimmers: Baetis, Centroptilum, Cloeon,
saprophagous or herbivorous Pseudocloeon
·········································································································································································
Ephemeridae* Burrowers: saprophagous Ephemera, Hexagenia
·········································································································································································
Heptageniidae* Poor swimmers or sedentary: Ecdyonurus, Epeorus,
saprophagous Heptagenia, Rhithrogena
·········································································································································································
Leptophlebiidae* Bottom crawlers: Habrophlebia, Leptophlebia,
saprophagous or omnivorous Paraleptophlebia, Traverella
·········································································································································································
Caenidae‡ Crawlers: omnivorous Brachycerus, Caenis
·········································································································································································
Ephemerellidae‡ Crawlers: saprophagous, Attenella, Ephemerella,
omnivorous, carnivorous Serratella
* Schistonota; ‡ Pannota
Mayfly nymphs have a closed tracheal system and instead breathe through
lateral abdominal gills; they also possess a pair of feathery cerci and a termi-
nal filament at the tip of the abdomen. They have chewing mouthparts and
feed on a wide range of submerged plant and animal matter. Most species are
detritivorous or herbivorous, but a few are carnivorous. Nymphs of Ametropis
neavei (Ametropodidae) employ a unique method of suspension feeding:
the front legs, which have modified setae, are used to create a vortex inside
specially made pits in sandy sediments. The vortex concentrates algae and
fine organic particles suspended in the water at the bottom of the pit, where
they are eaten. Vortices are also used as digging tools by other mayfly lar-
vae. By adopting a particular body posture, the larva of Pseudiron centralis
(Pseudironidae) is able to change the current flow locally to produce spiralling
eddy currents which excavate the sediment. As the sand grains are washed
away, small invertebrate animals become exposed and are swiftly consumed
by the larva.
Commensal relationships have been shown to exist between the larvae of
some chironomid midges and mayfly nymphs. The midge larvae, which feed
on minute particles of detritus, cling to the mayfly nymph’s gill filaments and
may move to other locations such as the legs and thorax as they get older. The
cerci of mayfly nymphs are used to detect the presence of foraging predators,
such as stoneflies, and help them to take avoiding action, although, oddly, in
laboratory experiments the removal of cerci in some species has not led to an
increase in the chance of being eaten by stonefly nymphs and other predators.
There is clear evidence that mayfly nymphs can perceive chemical cues from
predators and will take evasive or defensive action when the physical pres-
ence of a predator is felt. Mayfly nymphs tend to moult during the day when
predators like stonefly nymphs are least active.
When the nymphs are fully grown, they rise to the surface of the wa-
ter where they moult into the dull-winged subimago, which then flutters to
nearby vegetation. The nymphs of large species may crawl to the water’s edge
and moult to the subimago while clinging to plant stems or marginal stones.
The final moult to the shiny-winged adult form can take place in less than
an hour or after several days.
When conditions are right, large, sometimes massive, swarms of adults ap-
pear that have emerged simultaneously. In rare instances these have caused
accidents, the bodies of dead and dying adults piling deep enough on roads
to cause vehicles to skid. These mating swarms, which have a characteristic
rising and falling flight pattern, take place often at dawn or dusk, over wa-
ter, in clearings, or near obvious features of the landscape such as trees or
bushes called swarm markers. Males of many species are ‘bifocal’—they have
their eyes divided into upper and lower portions; the upper portion of the
eye is sometimes raised on short stalks. Enlarged facets in the upper region
help the males see potential mates above them in a swarm. Males are able
to recognize swarms of their own species and females of the same species,
who recognize the same sorts of makers and enter the swarms to mate. In
some species males jostle each other within the swarm but do not interfere
with other males when they are actually copulating. There is evidence that
males with bigger wings have more success at mating and this might be due
to them being able to jostle more effectively. Jostling does not occur in all
species but there is some evidence that larger wing size is attractive to females.
Males use their long front legs to hang on their chosen partner, engaging
their tarsi into special regions on the sides of the female’s thorax. Uniquely
among the insects, male mayflies have paired reproductive organs which are
EPHEMEROPTERA 49
inserted into paired genital openings in the female. When locked together,
copulating pairs gradually sink out of the mating swarm. Mated females ei-
ther lay hundreds or thousands of eggs by dropping them singly, in small
groups, or all at once in packets into the water. Some species release their
eggs as they dip the end of their abdomen under the water’s surface. The eggs
of many species have thread-like extensions that unravel on contact with wa-
ter and serve to keep the egg from being washed away. A few species retain
the eggs within their bodies and give birth to live nymphs. Up to half of all
ephemeropteran species may be parthenogenetic, either obligately, faculta-
tively, or accidentally—perhaps as an insurance strategy for times when males
have died too early—that is, before they met and mated with the females.
Mayflies are an extremely important component of freshwater food chains
and are eaten by most predatory species, both vertebrate and invertebrate.
As they make up a major part of the diet of trout and many other fish, it is
not surprising that adult and nymphal mayflies are extensively used by an-
glers as models for tying fishing flies. In the world of fly fishing, subimagos
are called duns and the adults are called spinners and mayfly ‘artificials’ have
names like the Greendrake, the Lake Olive, Lunn’s Particular, and Greenwell’s
Glory. Many different elements come together in fly fishing. A well-tied ar-
tificial is important but knowing the behaviour of the fish, as well as careful
observation of what they are eating, is essential to success.
Many species are intolerant of pollution and are thus useful as water qual-
ity indicators. In many countries of the northern hemisphere the decrease in
water pH due to acid rain and the reduction in dissolved oxygen due to pollu-
tion of various kinds has led to a marked reduction in the abundance of mayfly
nymphs, which, in turn has caused a decline in the number of freshwater
fish.
Mayflies have an unusual sort of economic impact in the damage they oc-
casionally inflict on the paint work of new cars. Car manufacturers hold large
numbers of new cars together in floodlit pounds often located on low-value,
marginal land near wet areas or reservoirs. During hot summer months, the
surface temperature of car roofs can be very high indeed. Egg-laden females,
mistaking a large area of cars for the surface of water, alight to lay eggs and
are instantaneously fried. The stricken insects roll over on their backs and the
boiling contents of their guts spill out and seep under the outstretched wings,
etching a neat insect shape in the paint.
Mayflies have a rich fossil record, with unambiguous representatives dat-
ing to the Permian (300 mya). Some fossils from the Carboniferous may
represent early mayfly lineages, or close relatives that have since gone extinct.
Modern mayflies probably appeared sometime in the Late Triassic–Early
Jurassic (200 mya).
Key reading
Alba-Tercedor, J. and Sanchez-Ortega, A. (1991). Overview and strategies of Ephemeroptera
and Plecoptera. Sandhill Crane Press, Gainesville.
Barber-James, H. M., Gattolliat, J.-L., Sartori, M., and Hubbard, M. D. (2008). Global
diversity of mayflies (Ephemeroptera, Insecta) in freshwater. Hydrobiologia 595: 339–50.
Brittain, J. E. (1982) Biology of mayflies. Annual Review of Entomology 27: 119–47.

Campbell, I. C. (ed) (1990). Mayflies and stoneflies: life histories and biology. Series Ento-
mologica series 44. Kluwer Academic Publishers, Dordrecht.
Edmunds, G. F. Jr. and McCafferty, W. P. (1988). The mayfly subimago. Annual Review of
Entomology 33: 509–29.
Harker, J. (1992). Swarm behaviour and mate competition in Mayflies (Ephemeroptera).
Journal of Zoology 228(4): 571–87.
Hubbard, M. D. (1990). Mayflies of the world. A catalog of the family and genus group
taxa (Insecta: Ephemeroptera). Flora and fauna handbook series 8. Sandhill Crane Press,
Gainesville.
Liegeois, M., Sartori, M., and Schwander, T. (2021). Extremely widespread parthenogenesis
and a trade-off between alternative forms of reproduction in mayflies (Ephemeroptera).
Journal of Heredity 112: 45–57.
Macan, T. T. (1982). The study of stoneflies, mayflies and caddisflies. Amateur Entomologist’s
Society, London.
Misof, B., Liu, S., Meusemann, K. Peters, R. S., Donath, A., Mayer, C., Frandsen, P. B., et al.
(2014). Phylogenomics resolves the timing and pattern of insect evolution. Science 346:
7639–7767.
Ruffieux, L., Sartori, M., and L’Eplattenier, G. (1996). Palmen body: a reliable structure
to estimate the number of instars in Siphlonurus aestivalis (Eaton) (Ephemeroptera:
Siphlonuridae). International Journal of Insect Morphology and Embryology 25: 341–4.
Simpson, S. J. and McGavin, G. C. (1996). The right fly: An angler’s guide to identifying and
matching natural insects. Aurum Press, London.
Soluk, D. A. and Craig, D. A. (1988) Vortex feeding from pits in the sand: A unique method
of suspension feeding used by a stream invertebrate. Limnology and Oceanography 33(4,
pt 1): 638–45.
Soluk, D. A. and Craig, D. A. (1990). Digging with a vortex: Flow manipulation facilitates
prey capture by a predatory stream mayfly. Limnology and Oceanography 35(5): 1201–6.
51
Odonata
(dragonflies and damselflies)
Common name Dragonflies and Metamorphosis Incomplete (egg, nymph,

Damselflies adult)
Derivation Gk. Odon–tooth Distribution Worldwide
Size Wingspan 18–200 mm. Number of families 30
Body length up to 150 Known world species 5,952 (0.53%)
mm
Short antenna
Large compound eye
Mobile head
Strong pterothorax
Spiny legs
Net-like
arrangement
of veins
Elongate abdomen Pterostigmata

Hind wing with
anal lobe
Anal appendages
Libellula quadrimaculata — Odonata
Key features
• aquatic nymphs, with labium modified into a prey-catching ‘mask’
• adults with elongate bodies and two pairs of long, similarly sized wings
• often brightly coloured or metallic
• distinctive fast, hovering, or darting flight
• usually seen near or over water
© Zestin Soh
Dragonflies are among the

most efficient aerial
predators to have ever
evolved.
Many damselfly males

possess conspicuous wing
patterns which are thought
© Zestin Soh
to be attractive to females or
used to ward off competing
males. Euphaea fraseri, India.
ODONATA 53
Superlative flying skills with speeds over 50 kilometres per hour make these
colourful insects instantly recognizable and enduringly popular with artists
and poets. In many cultures, dragonflies have had mythological associations.
Devil’s darning needles, a common name for dragonflies, reflects people’s
fears, while in South America many believe that certain large, graceful dam-
selflies (Pseudostigmatidae) are the spirits of recently dead humans. In sev-
eral countries, dragonflies are considered a great delicacy, and are caught,
threaded on skewers, and lightly grilled.
Although closely tied to fresh water by virtue of their aquatic nymphs,
adults can fly many miles away in search of prey. Despite the fact that they
are easily recognizable and distinctive insects, it is estimated that at least a
quarter of species worldwide remains undiscovered. The order is split into two
major suborders: the dragonflies (suborder: Epiprocta) and the damselflies
(suborder: Zygoptera).
In general, the head, which is large and very mobile, has biting,
downwards-pointing mouthparts, short, hair-like antennae composed of
fewer than eight segments, very large compound eyes, and three ocelli. Drag-
onflies have round heads and very large eyes while damselflies have broader
heads with widely separated eyes. The large eyes give virtually all-round vi-
sion and (as expected of these highly active, aerial hunters) they are able to
resolve distant objects better than any other insect.
The enlarged thorax is packed with flight muscles to power the front and
hind wings, which beat out of phase with each other, producing the whirring
noise characteristic of larger species. The sound is produced by the back edge
of the front wings brushing past the front edge of the hind wings at mid-
stroke. Odonates can control the frequency and amplitude of their wing beat
as well as the phase and angle of attack of the front and hind wings separately,
and, as a result, can hover and fly upwards, sideways, and backwards at will.
The basalar and subalar muscles that produce the downstroke or power stroke
are attached directly to sclerites at the bases of the wings, just outside the hinge
point. Contraction of these two sets of muscles can produce a downstroke ei-
ther with the leading edge down (basalar muscle contracted) or a downstroke
with the leading edge up (subalar muscle contracted). The upstroke for each
wing is produced by two indirect dorso-ventral muscles (one anterior and one
posterior) running from the top of the thorax (the tergum) to the bottom (the
sternum) of the segment bearing the wing. Contraction of these muscles pull
the tergum down and, as a result, the wing pivots upwards. The same as for the
downstroke, the insect can control the angle of the wing on the upstroke by
varying the power produced by the dorso-ventral muscles. Contraction of the
anterior dorso-ventral muscle will raise the wing with the leading edge down,
whereas contraction of the posterior dorso-ventral muscle will raise the wing
with the leading edge up. Pairs of wings beat together (i.e., the two front or
hind wings are either both up or both down at the same time) but by control-
ling what each set of flight muscles does, odonates can do different things on
opposite sides enabling them to turn around within their own body length.
It’s a bit like rowing with one oar pulling forwards and the other pulling back-
wards at the same time to make a tight turn. Dragonflies and damselflies also
have unusually low wing loading (mass to wing area), which contributes con-
siderably to their aerobatic capabilities and explains why catching dragonflies
can be a tiring occupation. The odonate arrangement of the flight muscula-
ture is unique among insects, and it probably arose 300 mya, when their much
larger relatives, the Meganisoptera, with wingspans of a little under a metre,
were wheeling and gliding through the Coal Measure swamp forests. Species
such as aeshnids (hawkers), with long abdomens and their centre of mass well
below and behind the wing bases, tend to be more stable and less manoeu-
vrable than those with short squat abdomens such as libellulids (darters and
chasers), whose shape makes them less stable and thus more responsive in the
air. Think of the difference between a passenger plane and a fighter plane.
Most species are active during the day and need to warm up their flight
muscles to a certain temperature before taking off. Many species bask out-
stretched but, if they get too hot, they can adopt an obelisk position, pointing
their abdomen towards the sun to minimize surface area. Large dragonflies
are able to lower their thoracic temperatures when flying in very hot condi-
tions by gliding for longer periods and by increasing haemolymph circulation
to their abdomen, which acts as a radiator. While the larger species might only
have to worry about vertebrate predators fast enough to catch them, smaller
species can fall prey to other aerial predators, such as robber flies (Asilidae).
Apart from the size and shape of the head, adults of the two suborders
can be distinguished easily in the field. Resting dragonflies tend to hold their
wings sideways from the body at rest, and their hind wings are broader near
their bases than the front wings. In damselflies, the wings are held together
over the abdomen at rest, and both hind and front wings are of similar shape
and narrow at their bases. As a result of the backwards slant of the thoracic
segments, it may appear, especially in some of the damselflies, that the wings
have been folded back along the body; however, they are in fact held directly
above the thorax. Another effect of thoracic rotation has been to bring the
legs forward. The three pairs of spiny legs are therefore in an ideal position
to act as an in-flight prey capturing net or basket. The shorter, front pair al-
low in-flight feeding. Although adults have wide-ranging tastes, some species
of social wasp are avoided. Dragonflies are generally active hunters, cruis-
ing to snatch prey in the air, from vegetation, or the ground, while the more
delicate damselflies tend to sit and wait for suitably sized prey to appear before
attacking.
The bright colours of odonates, which are involved in mate and rival recog-
nition, very often fade quickly after death. Many species show pruinescent
(powdery) colouration due to the morphological and biochemical structure
of their cuticular waxes thought to be involved in intraspecific communica-
tion. The abdomens of males of some tropical species can change colour in
response to a drop in ambient temperature. At 30–35◦ C the abdomen may
be bright red or blue, but as the temperature falls and the insects become less
active, pigment granules in the epidermis become clumped near the surface,
causing a darkening to grey-black. It has been suggested that this might make
them less easily seen by enemies.
ODONATA 55
Courtship is very variable within the order, being at one extreme non-
existent—males simply seizing females as they pass. At the other end of the
spectrum, courtship might be quite elaborate, with males luring females into
a territory by means of special displays. Reproduction in these species can
be energetically costly. In many species the mass of adults rises and may
even double as they mature. In males this is due to additional flight muscles
needed to hold territories, and in females it is due to the growth of ovaries.
The size of territory a male holds depends on the species, the population den-
sity, and often on the size of the male himself—the larger ones being territory
holders—while the smaller ones are consigned to inferior areas. Even so, these
satellite males will try to sneak in to mate with a female, so territory holders
have to defend their patch vigorously by constant patrolling, hovering, and
the engaging in the occasional aerial skirmish. The whole exhausting business
is worthwhile, as holding a territory ensures more matings.
Dragonflies and damselflies are unique among modern insects in that the
male has secondary sexual organs located ventrally at the front end of the ab-
domen. Males transfer sperm from the primary genital organs on the ninth
abdominal segment to the secondary sperm storage organ located on the
underside of the second or third abdominal segment by curling their ab-
domen round. Once the secondary storage organ is loaded, the males are
ready to copulate. The end of the male abdomen is equipped with clasping
organs, which are used to hold the female by the head in dragonflies or, on
♂ the prothorax behind the head, in damselflies.
The pair, locked together in tandem, may mate straight away or after a nup-
tial flight. This male grip is maintained during mating, which usually takes
place on a perch in the middle of the male’s territory or in the air. Unrespon-
sive females try to repel males by raising their front legs over their thorax.
♀ The need for protection is witnessed by the fact that the heads of many fe-
males are physically damaged by the force exerted by the male’s abdominal
claspers. To copulate, the female bends the tip of her abdomen round to join
with the secondary genitalia of the male. In this posture, called the wheel po-
sition, sperm is transferred via the penis to the female’s internal storage organ,
the spermatheca. Females may mate several times, especially if they want to
lay eggs in prime areas controlled by resident males. As in many other in-
Wheel mating position in sects, male dragonflies and damselflies need to ensure that they will father
damselflies. The male the eggs laid, especially if they are making an investment in time and energy.
grasps the female behind
the head, and she curls her
Sperm competition is an effective means of ensuring paternity. In most dam-
abdomen forward to couple selflies the hard penis, located on the second abdominal segment, is first used
her terminal genitalia with to scrape out the sperm of other males from the female’s spermatheca. The
his, located at the base of
his abdomen. removal of sperm may take up most of the copulation time and even though
the male may not be able to remove all of the previous male’s sperm, it has
been shown that last male sperm precedence is high. In many cases, the last
male to mate before oviposition takes place will father more than 90% of the
offspring. In species of damselfly that mate only once, the males do not have
a sperm-removing apparatus, but instead copulate for periods as long as sev-
eral hours, effectively excluding rivals. Recent work has shown that sperm is
transferred either as free motile sperm in a featureless seminal fluid or ag-

gregated into special structures called spermatodesms. Spermatodesms are
found in species who do not defend a territory and do not guard the females,
which tend to move away after copulation to lay their eggs. Free sperm is
found in species that hold territories and guard their mates, who lay eggs im-
mediately after mating. In the latter situation males have predictable access
to females. Several ideas have been put forward to explain the function of
spermatodesms. The structures may protect the sperm in some way, increase
sperm life, or provide some benefit in terms of nutrients to the female.
Female dragonflies mostly lay their eggs directly into water or attach them
to plants underwater, but species belonging to the family Aeshnidae and all
damselflies lay eggs in slits they cut into marginal or submerged plant stems
using an ovipositor. Males may retain in tandem or guard the females closely
as they lay their eggs.
The aquatic nymphs (referred in old works as ‘naiads’) are highly preda-
cious and equipped with a specialized prehensile modification of the labium
for rapid prey capture. The labium, or facial mask, bears hooked or toothed
labial palps and is hinged and can be shot forward from under the head in a
matter of milliseconds to seize prey which, for larger nymphs, may include
frogs and even small fish.
Slender
shape
Robust
shape
Flap-like or
feathery
Gills inside
external gills
rectal chamber
Nymphs. Damselfly nymph Dragonfly nymph
Odonate larvae respond to a range of mechanical, chemical, and visual

cues that warn them of the presence of predators such as fish, water bugs,
and beetles. In experiments, damselfly larvae responded to water from tanks
that contained predatory fish that had been feeding on damselflies or small,
sympatric fish, but not to water from tanks where the fish had been feeding
on exotic prey such as mealworms.
ODONATA 57
Damselfly nymphs have a closed tracheal system with no spiracles. In-

stead they obtain their oxygen from the water by means of three, feathery
or flap-like external gills located at the end of the abdomen. Known as cau-
dal lamellae, they are used for swimming and evidence shows that damselfly
nymphs use them to escape from dragonfly nymphs and similar predators. In
dragonfly nymphs, the gills are internal and are located in the rectal cham-
ber. Water is drawn in and forced out to provide a respiratory current over
the gills. If required, water can be forced out of the rectal chamber at high
speed to provide a jet-propelled escape. Some damselfly nymphs also pump
water in an out of the rectum for a great deal of the time but for different rea-
sons. The rate and stroke volume increases in the nymphs of some species if
they are stressed or short of oxygen. It has also been shown that the pump-
ing motions are concerned with the exchange of ions in the epithelium of the
hindgut, in particular to maintain a good flow of water over the rectal pads,
where special chloride cells are responsible for the uptake of inorganic ions.
Constant movement of water around the caudal lamellae will also maximize
gaseous exchange.
The habitats favoured by nymphs of different families ranges from water-
filled tree holes and temporary pools of water to streams, rivers, and lakes.
Some make burrows in swamps and others are able to hunt for prey after dark
out of water, among marginal vegetation. The nymphs of several families have
left the aquatic realm, and instead live in moist terrestrial environments, such
as leaf litter, mosses, and logs.
Dragonfly nymphs will readily engage in cannibalism, particularly if there
is a shortage of food and nymphs of different instars are present. The removal
of younger instars promotes the synchronization of hatching and popula-
tion development. Development is very much dependent on temperature as
well as food supply, and in cool temperate regions nymphs may take sev-
eral years before they crawl out of the water onto stones or up vegetation,
to emerge as adults. They do not mate right away; instead, they spend some
time feeding and maturing, gaining strength and body mass for the rigours
of reproduction. Tropical species may have two or even three generations a
year.
Although dragonflies sometimes capture and eat beneficial insects such
as honey bees, they are not pests and may rather have a useful role in the
control of natural insect populations. In some tropical countries, dragonfly
nymphs have been kept in drinking water storage tanks to control the num-
bers of larval Aedes aegypti mosquitoes far more safely and economically than
pesticides.
Odonata are very well represented in the fossil record, due to their large
size, their dispersal capabilities, and their association with aquatic habitats,
which facilitate fossilization. Extinct groups related to Odonata, such as the
Meganisoptera and the Geroptera, represent the oldest known flying insects,
likely originating in the Late Carboniferous (300 mya). Like other very old
groups such as the mayflies, we are uncertain when the ancestors of modern
Odonata appeared, although the Late Permian or Early Triassic (around 250
mya) are likely.
Key reading
Bybee, S. M., Ogden, T. H., Branham, M. A., and Whiting, M. F. (2008), Molecules, mor-
phology and fossils: a comprehensive approach to odonate phylogeny and the evolution
of the odonate wing. Cladistics 24(4): 477–514.
Corbet, P. S. (1980). Biology of Odonata. Annual Review of Entomology 25: 189–217.
Corbet, P. S., Longfield, C., and Moore, N. W. (1985). Dragonflies. Collins, London.
Dijkstra, K.-D. B., Bechly, G., Bybee, Dow, R. A., Dumont, H. J., Fleck, G., Garrison, R. W.,
Hämäläinen, M., Kalkman, V. J., Karube, H., May, M. L., Orr, A. G., Paulson, D. R., Rehn,
A. C., Theischinger, G., Trueman, J. W. H., Van Tol, J., Von Ellenrieder, N., and Ware, J.
(2013). The classification and diversity of dragonflies and damselflies (Odonata). Zootaxa
3703(1): 36–45.
Fincke, O. M., Waage, J. K., and Koenig, W. D. (1997). Natural and sexual selection compo-
nents of odonate mating patterns. In: Choe, J. C. and Crespi, B. J. (eds), The evolution
of mating systems in insects and arachnids, pp. 58–74. Cambridge University Press,
Cambridge.
Cambridge.
Johnson, D. (1991). Behavioural ecology of larval dragonflies and damselflies. Trends in
Ecology and Evolution 6(1): 8–13.
Miller, P. L (1994). The responses of rectal pumping in some zygopteran larvae (Odonata)
to oxygen and ion availability. Journal of Insect Physiology 40(4): 333–9.
Polcyn, D. M. (1994). Thermoregulation during summer activity in Mojave Desert dragon-
flies (Odonata: Anisoptera). Functional Ecology 8: 441–9.
Rehn, A. C. (2003). Phylogenetic analysis of higher-level relationships of Odonata. System-
atic Entomology 28(2): 181–240.
Siva-Jothy, M. T. (1997). Odonate ejaculate structures. Odonatologica 26(4): 415–37.
Theischinger, G. and Hawking, J. (2006). The complete field guide to dragonflies of Australia.
CSIRO Publishing, Collingwood.
Waage, J. K. (1979). Dual function of the damselfly penis: sperm removal and transfer.
Science 203: 916–18.
Waage, J. K. (1984). Sperm competition and the evolution of odonate mating systems. In:
Sperm competition and the evolution of animal mating systems. Smith R. L. (ed), pp.
251–90. Academic Press, Cambridge, MA.
Waage, J. K. (1986). Evidence for widespread sperm displacement ability in Zygoptera
(Odonata) and the means for predicting its presence. Biological Journal of the Linnean
Society 28(3): 285–300.
Watson, J. A. L. (1982). A truly terrestrial dragonfly larva from Australia (Odonata:
Corduliidae). Australian Journal of Entomology 21: 309–11.
59
Division Neoptera
·····························································································
The arrangement of moveable sclerites at the base of the wings allows neopter-
ans to fold their wings out of the way back along the abdomen when not in
use. This ability has allowed species in these orders to colonize all kinds of spe-
cialized terrestrial microhabitats unavailable to palaeopterous adult insects.
The Neoptera is split into two unequal divisions: the Exopterygota, where
the immature stages are similar to the adults (incomplete metamorphosis, or
hemimetaboly), and the Endopterygota, where the immature stages are very
different from the adults and there is an intermediate pupal stage (complete
metamorphosis, or holometaboly). The evolution of a pupal stage took place
around 350 mya and was a major contributor to the success of insects, of
which more than 80% belong to the Endopterygota. Several theories attempt
to explain the advantages conferred by holometaboly, proposing factors such
as protection from climatic instability, the exploitation of different resources
between immatures and adults, the allocation of tissue growth to the lar-
val stage, and tissue differentiation to the pupal stage, among others. It is
likely that several of these factors acted in conjunction. We know that the
earliest insects did not undergo metamorphosis (ametaboly), much like sil-
verfish and bristletails today. The difference between hemimetabolous and
holometabolous development might, at first, seem striking. In fact, some the-
ories proposed that the holometabolan larva is a prolonged embryonic stage,
and that the pupa corresponds to the immature stages of hemimetabolan
insects. However, the latest developmental and endocrinological studies sug-
gest that the immatures of both Exopterygota and Endopterygota in fact
represent the same developmental stage, and that the holometabolan pupa
corresponds to the last instar(s) of hemimetabolan insects. Future studies
will undoubtedly further elucidate the origins and mechanisms of insect
developmental modes.
Key reading
Belles, X. (2020). The evolution of metamorphosis. In: Insect metamorphosis. From natural
history to regulation of development and evolution. Belles, X. (ed), pp. 251–72. Academic
Press, Cambridge, MA.
Jindra, M. (2019). Where did the pupa come from? The timing of juvenile hormone sig-
nalling supports homology between stages of hemimetabolous and holometabolous
insects. Philosophical Transactions of the Royal Society B 374: 20190064.
Rainford, J. L., Hofreiter, M., Nicholson, D. B., and Mayhew, P. J. (2014). Phylogenetic
distribution of extant richness suggests metamorphosis is a key innovation driving
diversification in insects. PLoS One 9(10): e109085.
Rédei, D. and Štys, P. (2016). Larva, nymph and naiad—for accuracy’s sake. Systematic
Rolff J., Johnston, P. R., and Reynolds, S. (2019). Complete metamorphosis of insects.
Philosophical Transactions of the Royal Society B 374: 20190063.
Superorder Exopterygota
(= Hemimetabola)
·····························································································
In these orders, the young stages are called nymphs and, in most cases,
they feed on the same foods as the adults. Their wings develop on the out-
side of the body (exopterygote) and metamorphosis is simple or incomplete
(hemimetabolous). An exception can be found in thrips (Thysanoptera) and
many scale insects (Hemiptera: Coccoidea), which undergo a non-feeding,
pupa-like stage during their development (neometabolous).
Polyneoptera
The Polyneoptera are one of the major lineages of Neoptera, and com-
prise earwigs (Dermaptera), angel insects (Zoraptera), grasshoppers, crickets,
and katydids (Orthoptera), stick and leaf insects (Phasmatodea), webspin-
ners (Embioptera), stoneflies (Plecoptera), cockroaches and termites (Blat-
todea), mantids (Mantodea), ice crawlers (Grylloblattodea), and the newly
discovered heel walkers (Mantophasmatodea). We describe these orders in
a sequence that reflects the taxonomic groupings proposed by the latest
phylogenetic studies.
Haplocercata
This name (meaning simple-cerci in Greek) refers to a group of two closely re-
lated orders: earwigs (Dermaptera) and the elusive angel insects (Zoraptera),
which are characterized by simple, one-segmented cerci (albeit of remark-
ably different morphology and function). The evolutionary relationships of
either order had remained enigmatic until recently, when morphological and
molecular studies established their place in the insect tree of life.
61
Dermaptera
(earwigs)
Common name Earwigs Distribution Worldwide,

Derivation Gk. Derma–skin, hide; predominantly tropical
pteron–a wing regions
Size Body length 3–85 mm Number of families 12
Metamorphosis Incomplete (egg, nymph, Known world species 1,978 (0.17%)
adult)
Compound eye
Broad flattened head
Squarish pronotum
Shortish Short front wings

legs (tegmina)
Smooth and Folded hind wings

veinless
Telescopic, mobile
abdomen
Forceps
Forficula auricularia — Dermaptera
Key features
• elongate and slightly flattened
• abdomen telescopic with distinctive terminal, forceps-like cerci
• first pair of wings short and hardened
• second pair of wings fan-shaped and foldable
• prefer confined, humid microhabitats such as soil, litter, or under bark
A pair of Forficula dentata
© Paul Brock
browsing for nectar. Note
that the male has larger
cerci than the female. UK.
Mostly drab, nocturnal and generally reluctant to fly, it is possible to distin-

guish the vast majority of earwigs from all other insects by their distinctive
abdominal forceps. The eyes may large, small, or absent, the antennae are
long, thread-like, and multi-segmented, and there are no ocelli.
The order is divided into one extant suborder, the Neodermaptera, and
two extinct suborders. The most peculiar members of Dermaptera belong to
the families Arixeniidae (five species) and Hemimeridae (11 species), which
are blind, wingless, and their slender, unsegmented cerci are much less well
developed.
Arixeniid earwigs are stout, fairly hairy species found in south Asia, where
they live epizoically (i.e., non-parasitically on the body surface of an animal
or plant) on the fur or roosts of two species of molossid bat, feeding on their
skin fragments and excreta. The Hemimeridae are flattened, cockroach-like
earwigs found only in Africa, where all but one species live epizoically on the
fur of giant rats belonging to the genus Cricetomys. A single species has only
been found in the nests of the pouched rat Beamys major. It is thought that
these epizoic species live mutualistically on their hosts, where they feed on
the tough outer layers of the skin and a species of fungus that, left unchecked,
would lead to hair loss in the host. The females of both families give birth to
live nymphs. The eggs do not have a food supply of their own and develop
inside the mother’s reproductive tract, where they obtain nutrients directly
from a special tissue similar to a mammalian placenta. The morphological
DERMAPTERA 63
differences of these two families from all other earwigs led some researchers
in the past to propose that they might even represent a different order. Molec-
ular studies, however, have shown that both Arixeniidae and Hemimeridae
definitely belong to Dermaptera, and that their modified morphology and
epizoic habits evolved independently from each other, from more general-
ized earwigs. It is possible that both arixeniids and hemimerids are in fact
modified members of other earwig families and will no longer be considered
families of their own in the near future.
A characteristic feature of the order (with the exception of the epi-
zoic species already mentioned), which enables these insects to enter small
crevices in soil, bark, and stones, is the small toughened or leathery, vein-
less, front wings, which cover and protect the much larger, semicircular hind
wings. The hind wings have a greatly expanded fanlike area, which possesses
a sequence of alternating long and short ‘ribs’. Each long rib has a bending
point that acts as a hinge, which allows the wings to be folded like a fan and
then folded again along two transverse folds to enable them to fit under the
front wings. In earwigs, the mechanism that folds the hind wings and packs
them under the fore wings in an origami-like manner is so effective that it
serves as an invaluable model in the design of biologically inspired technol-
ogy. This mechanism is very ancient, having been found in extinct earwig
relatives from the Permian (280 mya), and is thought to have contributed to
the success of this order, as it allows them to inhabit tight spaces but also
retain their flight capacity—something that few other polyneopteran insects
can do. Another peculiarity of earwigs are their terminal forceps, which are
usually straight in females and curved in males and are used in a variety of
ways, such as weapons for defence, prey handling, courtship displays, and to
fold the fan-like hind wings under the toughened front wings. The flexible
and telescopic abdominal segments allow earwigs to use their forceps in all
directions. In those cases studied, individuals dominant in male–male inter-
actions are able to gain exclusive access to females. Defence in the adults of
some species relies on the production of a repellent secretion from abdominal
glands. Earwigs like to be in confined spaces (rarely ears) and, when resting,
will ensure that as much of their body as possible is in contact with the sub-
strate. There is some disagreement about the origin of the common name. It
either refers to earwigs entering ears or possibly to the ear-shape of the hind
wing.
Earwigs have simple chewing mouthparts that face forwards. They are om-
nivorous, some species preferring predominantly a plant diet, while others
are more predacious. Some species can be serious pests of flowers and crops,
while others have been shown to be of benefit in eating small insect pests
in fruit trees. Earwigs can be parasitized by tachinid flies such as Triarthria
setipennis, which is an endoparasitoid of the European earwig Forficula auric-
ularia. The female flies lay their eggs near a suitable host and the first instar
larvae climb on to the earwig and enter the body by penetrating the inter-
segmental membranes. This fly was introduced to North America early in
the twenty-first century in an attempt to control the accidentally introduced
European earwig.
Copulation takes place end to end, and females may retain the sper-
matophore for months before the sperm it contains is used to fertilize the
eggs. Females typically lay their eggs in tunnels they dig in the soil and show
a high degree of maternal care. Sometime the males assist in tunnelling but
take no further part in raising the young. Females groom their eggs, licking
and turning them to remove fungal spores and they will carry their eggs up
and down the tunnel to keep them in an even ambient temperature. Females
also guard their eggs from predators. Maternal care continues for some time
after the eggs hatch and the mother will feed the young nymphs by bringing
food into the nest or by regurgitation of part of her own meal. Eventually the
nymphs have to disperse, for as they grow older, their mother stops guard-
ing them and starts regarding them as a potential meal. Earwigs moult up
to five times and, apart from getting bigger and gaining additional antennal
segments at each moult, they look similar to their parents.
The earliest earwig relatives appear in the Late Permian–Early Triassic
(about 250 mya) and differed in some features from modern Dermaptera.
Their cerci were long and segmented, as is the general condition in most
other hemimetabolans, and their forewings were veined. The ancestors of
modern earwigs started making their appearance sometime between the Late
Jurassic–Early Cretaceous (150–100 mya), perhaps earlier.
Key reading
Deiters, J., Kowalczyk, W., and Seidl, T. (2016). Simultaneous optimisation of earwig
hindwings for flight and folding. Biology Open 5(5): 638–44.
Engel, M. S. and Haas, F. (2007). Family-group names for earwigs (Dermaptera). American
Museum Novitates 3567(1): 1–20.
Cambridge.
Jarvis, K. J., Haas, F., and Whiting, M. F. (2004). Phylogeny of earwigs (Insecta: Dermaptera)
based on molecular and morphological evidence: reconsidering the classification of
Dermaptera. Systematic Entomology 30(3): 442–53.
Sato, K., Pérez-de la Fuente, R., Arimoto, K., Seong, Y., Aonuma, H., Niiyama, R., and You,
Z. (2020). Earwig fan designing: biomimetic and evolutionary biology applications. Pro-
ceedings of the National Academy of Sciences of the United States of America 117(30):
17622–6.
Kuhlmann, U. (1995). Biology of Triarthria setipennis (Fallen) (Diptera: Tachinidae), a
native parasitoid of the European earwig, Forficula auricularia L. (Dermaptera: Forfi-
culidae), in Europe. Canadian Entomologist 127(4): 507–17.
Moore, A. J. and Wilson, P. (1993). The evolution of sexually dimorphic earwig forceps: so-
cial interactions among adults of the toothed earwig, Vostox apicedentatus. Behavioural
Ecology 4: 40–8.
Naegle, M. A., Mugleston, J. D., Bybee, S. M., and Whiting, M. F. (2016). Reassess-
ing the phylogenetic position of the epizoic earwigs (Insecta: Dermaptera). Molecular
Phylogenetics and Evolution 100: 382–90.
Nakata, S. and Maa, T. C. (1974). A review of the parasitic earwigs. Pacific Insects 16:
307–74.
Popham, E. J. (1965). A key to the Dermaptera subfamilies. Entomologist 98: 126–36.
Simmons, L. W. and Tomkins, J. L. (1996). Sexual selection and the allometry of earwig
forceps. Evolutionary Ecology 10(1): 97–104.
Terry, M. D. and Whiting, M. F. (2005). Mantophasmatodea and phylogeny of the lower
neopterous insects. Cladistics 21(3): 240–57.
65
Zoraptera
(angel insects)
Common name Angel insects, ground lice Distribution Primarily tropical, some
Derivation Gk. zoros–pure; species in subtropical and
a+pteron–wingless temperate regions
Size Body length 1.5–3 mm Number of families 2 (Spiralizoridae,
Metamorphosis Incomplete (egg, nymph, Zorotypidae)
adult) Known world species 44 (0.004%)
9 segmented antenna
Squarish pronotum
This is a wingless morph
Short cercus
Zorotypus hubbardi — Zoraptera
Key features
• very small, dimorphic (coming into two forms)
• wingless form is eyeless, with a termite-like appearance
• winged form with compound eyes, ocelli, and two-segmented tarsi
• short, one-segmented cerci
• extremely complex male genitalia
• rarely encountered, mostly associated with rotting wood
Zorapterans, such as this
© Piotr Naskrecki
Latinozoros sp. from Costa
Rica, are among the most
elusive of all insects, with
only a handful of known
species.
These small, delicate insects, whose wingless forms resemble minuscule ter-
mites, are among the least known insect orders. Their elusive habits in rotting
wood and leaf litter, combined with their small size, make them difficult to
collect, and as a result they have been poorly studied. Furthermore, their
extreme morphological uniformity has puzzled scientists regarding their clas-
sification for more than a century. Zoraptera were initially placed close to the
Paraneoptera (thrips, true bugs, bark lice, and lice), but subsequent studies
showed that they are in fact most closely related to earwigs (Dermaptera)
in the Polyneoptera. The possession of short, one-segmented abdominal
cerci and the generalized type of downward-pointing mouthparts betray their
polyneopteran affinities. Zoraptera is an ancient group, with a few fossils
known from Burmese amber (about 100 mya). Molecular clock estimates
suggest an origin in the Jurassic (200 mya), before the breakup of the super-
continent Pangaea, which explains the worldwide distribution of Zoraptera
(although they have not yet been discovered in Australia).
Zorapterans are sub-social, forming aggregations of up to 120 individ-
uals under bark or in piles of wood dust and leaf litter, where they eat
fungal threads, spores, springtails, and occasionally engage in cannibalism.
Zoraptera appear to be dependent on these aggregations, as individuals
removed from the group soon perish. It is possible that the grooming be-
haviour that these insects exhibit spreads essential antimicrobial substances
or perhaps endosymbiotic organisms, which are crucial for their survival.
ZORAPTERA 67
Adults are dimorphic, being either blind, pale, and wingless (resembling the
nymphs), or darkly pigmented with eyes, ocelli, and two pairs of pale, sparsely
veined wings. Winged morphs disperse to new locations and the wings are
then shed. This order was first discovered in the early twentieth century, and
all known species were initially contained in the genus Zorotypus in the sin-
gle family Zorotypidae. Subsequent morphological and molecular analyses
split Zorotypus into nine genera belonging to two families—Spiralizoridae
and Zorotypidae—while many more species are likely to be discovered in the
near future.
Although species from both families are morphologically extremely simi-
lar to each other, each family shows striking differences in male genitalia and
in the choice of reproductive strategy. In Spiralizoridae, male genitalia are
symmetrical, containing a very long, coiled intromittent organ. During mat-
ing, the intromittent organ uncoils inside the female reproductive tract, where
it serves to remove or destroy sperm deposited by previous males. Insemina-
tion is internal in this family. The family Zorotypidae, whose male genitalia
are asymmetrical and lack a coiled intromittent organ, follows an entirely
different reproductive strategy, which is unique not only among insects, but
Hexapoda as a whole. In some zorotypid species, the male deposits several
spermatophores externally on the abdominal tip of the female, each contain-
ing a single giant sperm, which is as long as the male’s entire body length.
The spermatophore and large sperm likely function as physical obstacles that
prevent the females from copulating with more than one male. Therefore,
males from both families go at great lengths at ensuring the paternity of their
offspring, but they achieve this in remarkably different ways.
Both families of Zoraptera also exhibit elaborate pre-copulatory courtship
behaviours, and males of many species give females gifts of secretions from
their cephalic gland to induce females to accept them as mates. It seems that
females can judge the quality of the gift and will terminate a copulation before
sperm is transferred if the male is inferior. Females mate two or three times
in succession with suitable males and evidence shows that by adopting this
strategy, they can lay more eggs and get more cephalic secretions than if they
mated with different males. In some species, males are larger than females
and wrestle, often using hind-leg kicking and head butting to gain matings.
Mating success appears to be determined by age and not necessarily by size,
and dominant males may get three quarters of all matings.
Zorapterans have been increasingly studied in the last few years, and there
is much to learn from these elusive yet fascinating insects.
Key reading
Choe, J. C. (1995). Courtship feeding and repeated mating in Zorotypus barberi (Insecta:
Zoraptera). Animal Behaviour 49: 1511–20.
Choe, J. C. (1994). Sexual selection and mating in system in Zorotypus gurneyi Choe (In-
secta: Zoraptera): II determinants and dynamics of dominance. Behavioural Ecology and
Sociobiology 34: 233–7.
Choe, J. C. (1994). Sexual selection and mating system in Zorotypus gurneyi (Insecta: Zo-
raptera): I dominance hierarchy and mating success. Behavioural Ecology and Sociology
34: 87–93.
Choe, J. C. (1997). The evolution of mating systems in Zoraptera: mating variations and
sexual conflicts. In: The evolution of mating systems in insects and arachnids. Choe, J. C.
and Crespi, B. J. (eds), pp 130–45. Cambridge University Press, Cambridge.
Dallai, R., Mercati, D., Gottardo, M., Machida, R., Mashimo, Y., Matsumura, Y., and Beu-
tel, R. G. (2014). Giant spermatozoa and a huge spermatheca: A case of coevolution of
male and female reproductive organs in the ground louse Zorotypus impolitus (Insecta,
Zoraptera). Arthropod Structure & Development 43(2): 135–51.
Dallai, R., Mercati, D., Gottardo, M., Machida, R., Mashimo, Y., Matsumura, Y., and Beutel,
R. G. (2013) Divergent mating patterns and a unique mode of external sperm transfer in
Zoraptera: an enigmatic group of pterygote insects. Naturwissenschaften 100: 581–94.
Dallai, R., Mercati, D., Gottardo, M., Machida, R., Mashimo, Y., and Beutel, R. G. (2011).
The male reproductive system of Zorotypus caudelli Karny (Zoraptera): Sperm structure
and spermiogenesis. Arthropod Structure & Development 40(6): 531–47.
Engel, M. S. (2007). The Zorotypidae of Fiji (Zoraptera). Bishop Museum Occasional Papers
91: 33–8.
Kočárek, P., Horká, I., and Kundrata, R. (2020). Molecular phylogeny and infraordinal
classification of Zoraptera (Insecta). Insects 11: 51.
Matsumura, Y., Beutel, R. G., Rafael, J. A., Yao, I., Câmara, J. T., Lima, S. P., and Yoshizawa,
K. (2020). The evolution of Zoraptera. Systematic Entomology 45: 349–64.
Valentine, B. D. (1986). Grooming behavior in Embioptera and Zoraptera. Ohio Journal of
Science 86(4): 150–2.
Wipfler, B., Letsch, H., Frandsen, P. B., Kapli, P., Mayer, C., Bartel, D., Buckley, T. R.,
Donath, A., Edgerly-Rooks, J. S., Fujita, M., Liu, S., Machida, R., Mashimo, Y., Misof,
B., Niehuis, O., Peters, R. S., Petersen, M., Podsiadlowski, L., Schütte, K., Shimizu, S.,
Uchifune, T., Wilbrandt, J., Yan, E., Zhou, X., and Simon, S. (2019). Evolutionary his-
tory of Polyneoptera and its implications for our understanding of early winged insects.
Proceedings of the National Academy of Sciences of the USA 116: 3024–9.
69
Plecoptera
(stoneflies)
Common name Stoneflies Metamorphosis Incomplete (egg, nymph,

Derivation Gk. plectos–plaited, adult)
woven; pteron–a wing Distribution Worldwide, but mainly
Size Body length 3–48 mm. cool temperate regions
Maximum wing span Number of families 16
about 100 mm Known world species 3,743 (0.33%)
Thread-like
antenna
Bulging eye
Squarish prothorax
Meso- and
metathorax
similar
Hind wing broader

than front wing
Slightly flattened body

Cercus
Perlodes microcephala — Plecoptera
Key features
• aquatic nymphs
• weak flying insects near water
• adults typically rest with wings held flat or rolled around body
• important in aquatic food chains
© Anne Riley
Stoneflies use vibrational
signals to attract each other
and mate. Nemoura sp., UK.
Stoneflies are soft or leathery looking, generally slender insects with soft,
slightly flattened bodies. Although they have two pairs of membranous wings,
they are not good fliers, seldom travel far from water, and spend a great deal
of time resting on marginal rocks and vegetation. The front wings are much
narrower and slightly longer than the hind wings, which are folded in pleats
beneath. Both pairs of wings may have complex patterns of cross veins. The
head has bulging eyes, two or three ocelli, and thread-like, multi-segmented
antennae. The mouthparts are weakly developed or non-functional and many
species, which live less than a fortnight as adults, do not feed at all. In other
species, adults feed by scraping algae, lichen, and similar material from rocks
and other substrates. The legs are quite long, robust, and appear widely sep-
arated. The elongate abdomen may be cylindrical or slightly flattened and
has a pair of single- or multi-segmented cerci at its posterior end. The or-
der is divided into two suborders: the Arctoperlaria (12 families) and the
Antarctoperlaria (four families). With the exception of one family, the No-
tonemouridae, all Arctoperlaria are found in the northern hemisphere. All
families in the Antarctoperlaria, (Austroperlidae, Eustheniidae, Diamphip-
noidae, and Gripopterygidae) are found in the southern hemisphere.
Males attract females by rubbing, tapping, or drumming a species-specific
signal on the substrate to which the females respond with acoustic signals
of their own. The noises are produced by a hardened finger or hammer-like
structure on the underside of the abdomen being struck or rubbed against the
ground, although in a few species belonging to the family Chloroperlidae, no
contact is made, and the sound is produced by tremulation—body vibrations
transmitted to the substrate via the legs and tarsi. Theories suggest that this
method of signalling may have evolved as it is less likely to be intercepted
PLECOPTERA 71
by acoustically searching predators. In most cases, male and female duetting

occurs and in some, parts of the female’s call may overlap with that of the
male. Duetting has been shown to reduce the time spent by the male search-
ing for the stationary responding female. Male stoneflies typically follow a
pattern of drumming and directed searching until the source of the vibrations
(the female) is found. Females will move to a different location if the males
take too long to find them, which may have the double advantage of reduc-
ing the risk of predation and selecting a fit mate. Once the male has located
her, mating takes place almost immediately on the ground or on plants. Male
stonefly calls vary from series of single beats to quite complex songs with dis-
tinct phases varying in beat interval, beat number, and other characteristics,
whereas female responses are generally simple.
Females, who can lay many hundreds or thousands of eggs, deposit slimy
egg masses into the water. The eggs may be spherical and adhesive, flattened,
spindle-shaped, with filamentous projections or other devices to ensure that
they stick to objects or lodge in crevices. The aquatic nymphs are quite broad
and flattened with noticeable developing wing pads and a pair of long ter-
minal, abdominal tails. They have a closed tracheal system and obtain their
oxygen by straightforward diffusion through the surface of the body or by
means of a variety of branched or feathery gill tufts located on different parts
of the body. Some species have gill tufts at the end of their abdomens, while
others have gills on the neck and thorax. Remnants of thoracic and abdominal
gill tufts may still be visible in the adult stages. Cold, fast-flowing, stony- or
gravel-bottomed streams are preferred, and many species are very intolerant
of pollution. Stonefly nymphs are mostly herbivorous but some families (Per-
lidae, Chloroperlidae, and Perlodidae) contain omnivorous and predacious
species. Nymphs pass through anything up to thirty moults and may take
from one up to four years before moving away from the water to emerge as
adults, often during the hours of darkness. The nymphs of predatory stone-
flies (Perlodidae) have been shown to be able to respond to the swimming
patterns of their prey and can distinguish between prey and non-prey species
of mayfly nymphs. Females may mate with several males and mate guarding
to prevent access by rival males has been recorded for some species. Males
may guard a newly moulted female until she is ready to mate, often before
her cuticle has fully hardened.
Plecopterans are generally adapted to cold conditions and in the northern
hemisphere, many species, such as those belonging to the Capniidae, may
become adults, mate, and die during the cold months of Winter.
Stonefly nymphs are not defenceless. It has been shown that species such
as Pteronarcys dorsata (Pteronarcyidae) have complex defensive repertoires.
They are able recognize different types of predator and respond either by
feigning death if the predators are fish, or by backing away and using re-
flex bleeding if the predators are tactile hunters such as crayfish. Some adult
pteronarcyids also show reflex-bleeding if attacked and, in some species, the
haemolymph is forced out a distance of many centimetres, accompanied by
a distinct popping noise. Nymphs are able to learn paths to shelters like
particular groups of stones that provide a refuge from predatory fish.
Plecopterans are a very important component in aquatic food chains and,

like other aquatic insect groups such as mayflies and caddisflies, are imitated
in patterns of fishing fly used by anglers. As stone flies are usually very intol-
erant of low water oxygen, they are useful as aquatic bio-indicators. The large
nymphs of the California salmonfly (Pteronarcys californica) may have been
eaten by native Americans of the Achomawi, Modoc, and Wintun nations of
northern California.
The origins of Plecoptera are lost in the abyss of time. Possible plecopteran
relatives are known from the Permian (300–250 mya), although the ancestors
of the modern representatives likely appeared much later, around the Jurassic
(200 mya), based on phylogenomic and biogeographical data.
Key reading
Alba-Tercedor, J. and Sanchez-Ortega, A. (1991). Overview and strategies of Ephemeroptera
and Plecoptera. Sandhill Crane Press, Gainesville.
Campbell, I. C. (ed) (1990). Mayflies and stoneflies: life histories and biology. Entomologica
series 44. Kluwer Academic Publishers, Dordrecht.
Du Bois, C. A. (1935). Wintu ethnography, 36. University of California Press, Berkeley.
Illies, J. (1965). Phylogeny and zoogeography of the Plecoptera. Annual Review of Entomol-
ogy 10: 117–40.
Landolt, P. and Sartor M. (eds) (1997). Ephemeroptera & Plecoptera: biology, ecology,
systematics. MTL—Mauron, Tingley & Lachat SA, Fribourg.
Marden, J. H. and Kramer, M. G. (1994). Surface-skimming stoneflies: a possible interme-
diate in insect flight evolution. Science 266(5184): 427–30.
Society, London.
Moore, K. A. and Williams, D. D. (1990). Novel strategies in the complex defensive
repertoire of a stonefly (Pteronarcys dorsata) nymph. Oikos 57(1): 49–56.
Pekarsky, B. L. and Wilcox, R. (1989). Stonefly nymphs use hydrodynamic cues to discrim-
inate between prey. Oecologia 79(2): 265–70.
Szczytko, S. W. and Stewart, K. W. (1979). Drumming behaviour of four western Nearctic
Isoperla (Plecoptera) species. Annals of the Entomological Society of America 72: 771–86.
Stewart, K. W. and Maketon, M. (1991). Structures used by Nearctic stoneflies (Plecoptera)
for drumming, and their relationship to behavioral pattern diversity. Aquatic Insects
13(1): 33–53.
Stewart, K. W. (1994). Theoretical considerations of mate finding and other adult behaviors
of Plecoptera. Aquatic Insects 16: 95–104.
Stewart, K. W. (1997). Vibrational communication in insects: epitome in the language of
stoneflies. American Entomologist 43(2): 81–91.
Sutton, M. Q. (1985). The California salmon fly as a food source in northeastern California.
Journal of California and Great Basin Anthropology 7(2): 176–82.
Ward, J. V. (1994). Ecology of Alpine streams. Freshwater Biology 32(2): 277–94.
Zwick, P. (2000). Phylogenetic system and zoogeography of the Plecoptera. Annual Review
of Entomology 45: 709–46
73
Orthoptera
(grasshoppers and crickets)
Common name Grasshoppers, crickets, Metamorphosis Incomplete (egg, nymph,

and their relatives adult)
Derivation Gk. orthos–straight; Distribution Worldwide but mostly in
pteron–a wing warm regions
Known world species 23,434 (2.07%)
Hind legs enlarged for jumping

Front wings toughened
Saddle-shaped pronotum (tegmina)
Large compound
eye
Downwards-facing mouthparts
Schistocerca gregaria — Orthoptera
Key features
• hind legs usually much larger and longer than other legs and used for
jumping.
• many species make sounds using hind legs and/or front wings
• characteristic ‘grasshopper’ shape with pronotum extended down at each
side
• several species can be serious crop pests
A spermatophore
transferred by the male to a
© Nicky Bay
female katyidid, serves as a
nutritious meal. Sabah,
Malaysia. © Nicky Bay
A female katydid ovipositing

inside a branch. Onomarchus
sp., Singapore.
ORTHOPTERA 75
Also known as dune crickets,
© George McGavin
schizodactylids often
construct deep tunnels in the
sandy riverine habitats.
Schizodactylus sp., Northern
India.
Springing and singing might describe the species in this, the largest order
of hemimetabolous insects that have chewing mouthparts. Of course, a few
cannot jump and quite a few do not sing but very many of the ones com-
monly encountered do both things quite well. Orthopterans can be found in
just about every conceivable terrestrial habitat, even in caves or burrowing in
soil, and some live permanently in association with ant nests. The range of
the order is divided into two suborders: the Ensifera and the Caelifera. These
taxa share several common characteristics. Both have conspicuous eyes and
may have ocelli. They all possess downward-pointing chewing mouthparts, an
enlarged, saddle or shield-shaped pronotum, toughened, narrow front wings
(or tegmina) with larger fan-folded hind wings (when present), and large hind
legs modified for jumping. The abdomen has a pair of short, terminal cerci.
The Ensifera, comprising the crickets, bush-crickets, and katydids, have
long or very long, multi-segmented antennae, and are mainly nocturnal
and solitary (Table 2.1). Most species are cryptically coloured brown or
Table 2.1 Principal ensiferan families
Family Habits Examples of genera

Tettigoniidae (katydids) Mostly in foliage. Her- Acanthodis, Anabrus,
(~ 7,100 spp.) bivorous, saprophagous, Ephippiger, Orchelimum,
predacious. Tettigonia
Some pests
·········································································································································································
Gryllidae and related families Mainly ground-living. Acheta, Gryllodes, Gryllus,
(true crickets) (~ 4,900 spp.) Omnivorous Oecanthus
·········································································································································································
Rhaphidophoridae (cave Many in caves, burrows. Ceuthophilus,
crickets) (~ 660 spp.) Saprophagous, predacious Dolichopoda, Tachycines,
Troglophilus
·········································································································································································
Gryllotalpidae (mole Burrowing. Gryllotalpa, Scapteriscus
crickets) (~ 110 spp.) Omnivorous.
Some are pests
green to mimic dead or living leaves. For example, the new world katydid,
Markia hystrix, is a remarkable mimic of the lichens on which it lives. In
female ensiferans, the ovipositor is always prominent and sword-, sickle-,
or stiletto-shaped. Many species are herbivorous, eating mostly plants other
than grasses, but some are partly or wholly predacious. The Caelifera, com-
prising grasshoppers and locusts, have short antennae and the females never
have prominent ovipositors. They are generally ground-living, diurnal, grass-
feeders, and many are cryptically coloured to blend in with the ground, or
they are warningly coloured to advertise their ability to produce unpalatable
or repellent secretions from special glands. Some species regurgitate their gut
contents as a defensive strategy. The brightest colours and striking patterns
of black, red, yellow, and orange are seen in the chemically defended species
belonging to the small families Pyrgomorphidae and Romaleidae. Cryptic
species may have bright colour patches on their hindwings to distract preda-
tors as they jump up, fly, and then drop to the ground out of sight. Members
of several families (e.g., Chorotypidae, Trigonopterygidae) take crypsis to
the extreme, with their entire body shape and colouration imitating that of
a leaf.
Nymphal development goes through 4–11 instars. As in the Odonata, wing
pad orientation changes in development. In young nymphs the front edge of
the wing pads is lateral and ventrally directed. In that last instar the orienta-
tion changes so that the front edges of the wing pads are directed dorsally, and
the hind wings overlap the front wings. The normal orientation is resumed in
the adult stage (front edges of the wings directed ventrally).
The ability to jump, seen in the majority of species, explains an older name
for the order—Saltatoria. Jumping can be part of an escape from potential
danger or to enable the insect to take to the air. The strong hind legs are
also used to kick out at attackers or rivals. Some groups of orthopterans, who
have evolved habits such as burrowing or digging, cannot jump very well.
Although orthopteran legs vary greatly in size, the mechanically ingenious
jumping mechanism of the hind leg is essentially the same. The tibia is con-
trolled by a large extensor muscle, which occupies most of the volume of the
femur, and a much smaller flexor muscle. Energy, produced by contraction
of the large extensor muscle, is stored prior to the jump. The energy is stored
in two cuticular springs at the knee joint (semilunar processes) and in the
slightly elastic extensor apodeme (= tendon). The catch, which keeps the tib-
iae from extending too soon, is formed by a pocket in the flexor apodeme,
which sits over a lump at the end of the femur. When the grasshopper has
contracted its extensor muscle fully and has stored as much energy as it can,
it releases the tension on the flexor muscle, which causes the catch to dis-
engage and the full force of the jump to be developed. The system allows
the power of the muscles to be amplified. In the case of the adult Desert
Locust, the amplification is about ten times, allowing a long jump of up to
80 cm.
Singing, usually by males, is another well-known behavioural character-
istic of the order. The songs, and the manner in which they are produced,
ORTHOPTERA 77
are diverse. The songs are generally designed to attract females but can also
be involved in territoriality or signalling aggression or alarm. As a very
rough guide grasshoppers (Acrididae) generally sing during the day, katydids
(Tettigoniidae) sing at night, and crickets (Gryllidae) sing during the day or
night.
Male ensiferans typically produce songs by stridulating, that is, rubbing
parts of their front wings against each other. In gryllids, for example, the un-
derside of both wings has a vein near the base which is modified as a file with
File
between 35 and 300 teeth. The stiffened edge of the other wing forms a scraper
or plectrum. As the right wing overlaps the left wing, the file on the right wing
and the scraper on the left wing rasp against each other and set up vibrations
that pass through the wings and excite a special clear harp or mirror region
on each wing. The resonant frequency of these harp regions closely matches
the song frequency. When singing, crickets raise their wings at an angle to
allow the sound to be radiated more effectively. A pulse of sound is made
on the closing stroke of the front wings and the songs vary in the number
and character of the pulses produced (4–200 per second) and the way the
The underside of the front
pulses are grouped. Although the stridulatory structures on the front wings
wing of a gryllid showing of crickets, bush-crickets, and katydids are superficially very similar, remark-
the stridulatory file. ably, they have evolved independently at least four times—suggesting that
sound communication also evolved several times. Both winged and wingless
crickets can make vibrations that pass through the substrate (e.g., bark, soil,
but not air) by tremulating (shaking their body) or by drumming their ab-
domen or feet against a solid surface. Substrate vibrations are considered to be
the original mode of signalling in Ensifera, with sound appearing later in their
evolution.
Cricket songs can be very musical, and frequencies vary from 1500 Hz–10
kHz. Katydids tend to be noisier and can employ frequencies from 20–100
kHz, but the details of sound production differ in the two groups. Several
species of tree crickets (Oecanthus spp.) increase the volume of their song
by positioning their wings across a hole they chew in a large leaf, which acts
as a baffle. An even more incredible example of acoustic engineering is seen
in mole crickets (Gryllotalpidae). Mole crickets are remarkable for their in-
credible morphological resemblance to mammalian moles. The body is very
The unique acoustic burrow robust, cylindrical, and covered with very short, velvety hairs. The legs are
of the more cricket. short and strong, and the front legs are particularly modified for digging, be-
ing much broadened and armed with stout teeth. The eyes are small, and the
ovipositor is very short or vestigial. But they are also famous for their ability
to construct elaborate singing burrows to make their songs more audible. The
burrow has a bulb region tuned to resonate at the song’s carrier frequency (3.4
kHz) and twin, exponentially flared horns to amplify and carry the song to
the surface. The burrow is very effective and at a metre above the burrow the
peak sound level can be 92 decibels, which is enough to carry nearly a mile
in quiet conditions.
© George McGavin
Tropidacris cristata is a large
grasshopper from Central
and South America with a
wingspan of up to 18 cm.
Costa Rica.
Once a female has been attracted, the two mate above ground. Eggs are laid
in an underground chamber and full development through the ten or so
nymphal stages may take up to two years.
Caeliferan species make sounds in a variety of ways, ranging from hind-
wing snapping and palp and/or mandible rubbing to the more familiar
rubbing of the inner face of the hind femur against the edge of the tegmina.
The inside of each femur has a row of tiny teeth or pegs that set up vibrations
in the front wings. As in ensiferan singers, the tone of the song is dependent
on the spacing of the teeth and the speed at which the leg is moved (Table 2.2).
As for the mechanisms of sound production, the tympanal or hearing or-
gans in Orthoptera take two distinct forms. In the Caelifera, a tympanum is
situated on either side of the first abdominal segment. Each organ consists of
a thin tympanic membrane located over an air sac. Attached to the underside
of the membrane is a chordotonal organ of varying complexity. A more com-
plicated system is found in the Ensifera, where a single or a pair of tympanal
Table 2.2 Principal caeliferan families
Family Habits Examples of genera

Acrididae (grasshoppers, On ground and low plants. Acrida, Chortoicetes, Locusta,
locusts) (~ 6,700 spp.) Herbivorous. Melanoplus, Nomadacris,
Many serious pests Schistocerca
·········································································································································································
Tetrigidae (pygmy locusts) Ground-living, woods and Discotettix, Paratettix, Tetrix,
(~ 1,823 spp.) wet areas. Xerophyllum
Saprophagous, herbivorous
·········································································································································································
Pyrgomorphidae (Bush Ground-living. Atractomorpha, Petasida,
hoppers, Bush locusts) Bushes, shrubs, some in Phymateus, Pyrgomorpha
(~ 500 spp.) trees and on grasses.
Some local pests
ORTHOPTERA 79
organs are situated on the front tibiae close to the knee joint. The leg around
the tympana is slightly swollen and typically has two longitudinal slits that
each open to a tympanic chamber. Where there is a pair of tympanic mem-
branes, they are unequal in size and the trachea behind them is divided by
a rigid membrane. Vibration of the tympanic membranes is picked up by a
number of chordotonal organs, the most important of which is the called the
crista acustica. The crista acustica comprises a row of sensory cells attached to
the anterior tympanal membrane. In gryllids the trachea associated with the
tympana are part of the normal tracheal system but the most complex form
of orthopteran hearing, which is found in tettigoniids, relies on a specialized
acoustic trachea that runs up the middle of each front leg and is not part of
the normal tracheal system. The acoustic tracheae on each side are connected
and joined to specialized acoustic spiracles in the prothorax. If hearing is to be
of use in locating a distant sound source, it must be directional. The problem
common to insects is that their small size makes differences in intensity or
time of arrival of sound at two different tympanal organs very small indeed.
The problem is overcome by the introduction, mechanically or acoustically,
of a time delay between the two ears. In tettigoniids the main input of sound
to the system comes from the acoustic spiracles on the thorax; it then trav-
els down and is amplified by the tapering acoustic tracheae before it reaches
the tympanal organs. The effects of these sounds are modified by sound vi-
brations picked up directly through the slits’ opening to the outside from the
tympanic chambers. The precise mechanism of cricket hearing is, however,
exceedingly complex and not yet fully understood.
Courtship in the Orthoptera varies enormously from very simple affairs
where males do not engage in any sort of foreplay, to long and very complex
‘wooing’ involving precise movements and mutual contact of many parts of
the body accompanied, at the right time, by different types of songs. Humans
may find the singing of crickets attractive and in several countries, they are
kept as pets for their songs. But the intended audience are just as choosy. It has
been shown that the quality of sound a male cricket makes is very important
to his chances of a successful mating. Smaller males with asymmetrical harps
produce less pure tones or a higher frequency than do larger males with larger
symmetrical harps. Not surprisingly, females prefer males who sing better,
louder, and longer.
Being the last male to mate with a female is very advantageous. The way
in which sperm is used—a last-in first-out basis—makes it likely that the last
mate will father most of the eggs. In the gryllid, Truljalia hibinonis, males are
able to remove more than 87% of the sperm of a previous mating after they
have copulated. The rival’s sperm is not wasted but eaten. Simply hanging on
tightly to the female’s back until she lays her eggs and kicking out at any other
male who comes near is a commoner strategy. Where females are in short
supply, lone males will make strenuous attempts to take over. Fighting be-
tween male crickets is common and has even been popular in some countries
as a gambling sport.
Ensiferans typically mate with the smaller male beneath or below the fe-
male and the resultant fertilized eggs are often laid singly or in small batches
in plant tissues. Males transfer sperm in spermatophore consisting of an am-

pulla containing the sperm and a nuptial gift in the form of a spermatophylax,
which the female eats after mating. The consumption of the spermatophylax
enables the female to produce more offspring. Males who have the chance of
lots of matings provide smaller individual gifts than those males who mate
with few females. This suggests that the gift might have to do with guarantee-
ing paternity when the father’s investment is high. But some other research
has shown that spermatophylax size simply correlates with sperm number
and ejaculate volume, suggesting a simple sperm protective function. A larger
spermatophylax will take the female longer to eat so, by the time she has fin-
ished, the sperm will be transferred. There are probably two types of gift,
depending on species: one, a low-value, easy-to-produce spermatophylax,
which is protective; the other, a large, highly nutrient-rich spermatophylax,
which also increases the fecundity of the females. In some gryllids the sper-
matophylax provided by the male is a bit of a confidence trick. True, it does
contain a significant amount of amino acids—enough to persuade the female
to eat it but not so much that the male is depleted. She might actually use up
more energy eating it than she gets back.
The mating of behaviour of North American sagebrush crickets belonging
to the genus Cyphoderris is remarkable for it involves males offering portions
of their body tissue as nuptial gifts to the female. In Cyphoderris strepitans,
once the female has mounted the male’s back, she begins to eat his hind wings.
These are especially fleshy, and the female also drinks the haemolymph that
issues from the wounds she makes. Males, rather than wishing to terminate
the cannibalism, make it easy for the female by raising their wings into a
convenient position and make sure that she stays put by grasping the end of
her abdomen very tightly with a special hooked clamping device while they
transfer their spermatophore. It has been suggested that this device may allow
males to force females to copulate either because the male has an insufficient
amount of hind wing to keep her interest or, more likely, it has evolved to
prevent females from taking advantage of free meals. In species of another
genus, Bradyporus, the female simply bites their partner’s back and drinks
haemolymph.
An interesting reversal of the normal sexual roles in seen in the Mormon
cricket, Anabrus simplex, and other similar situations are known. In poorer
habitats it is only well-nourished males who choose a mate. These males have
a large, nutritious nuptial gift, which can be one-third of their total body
weight and their love song will attract several females who compete for ac-
cess. The female who succeeds will mount the male’s back but might still not
be chosen. The male is able to tell how heavy she is (and therefore how many
eggs she might be carrying), and if not satisfied, he may choose another mate.
The reversal of normal sexual role is seen when paternal investment is high
and can be induced experimentally in some bush cricket species.
In the Caelifera, mating takes place with the male on top of the female and
the eggs (up to 200) are normally laid in the soil, protected by an egg pod made
of a special protective foam produced by the female. Caeliferan ovipositors are
short and squat, composed of pairs of valvulae that expand and grip the side
ORTHOPTERA 81
of the egg-laying burrow to pull the abdomen down, in some cases, to more
than double its length.
Many acridid species are agricultural pests, capable of causing immense
crop losses in tropical, subtropical, and even in temperate regions. In North
America, several species in the spur-throated grasshopper genus Melanoplus
can be serious pests. The most notorious acridid species, however, are com-
monly referred to as ‘locusts’. Locust is the generic common name given
to fewer than ten species in the genera Schistocera, Locusta, Locustana, No-
madacris, Anacridium, and Chortoicetes. The defining feature of these species
is that they can exist in solitary or gregarious forms. As solitary individuals,
even large populations do little lasting damage, but as gregarious swarms they
can devastate vast swathes of vegetation. The Desert locust, Schistocerca gre-
garia, probably one of the most damaging insects in the world, has been a pest
since agriculture first developed and is mentioned as the eighth of the ten bib-
lical plagues of Egypt. When conditions are right, and rain brings about the
growth of lush vegetation, the locusts are able to feed and lay more eggs than
usual. More nymphs hatch but now, rather than shy away from each other,
they mass together in growing numbers. They have become gregarized. These
gregarious black, yellow, and orange-striped hoppers (as they are called) look
very different from the pale green solitary nymphs. Once locust nymphs be-
come aggregated in the habitat and reach a critical local population density,
they suddenly shift from uncoordinated movement to collective mass march-
ing. This striking behavioural transition has been explained by computer
modelling and is a classic case of an emergent phenomenon, arising because
of local interactions among individuals. The hoppers feed and march until
they become adult. The result can be massive swarms of adults, covering as
much as 1000 km2. Although they are good fliers, they tend to move in the
direction of the prevailing winds. When a food source is located, the swarm
will descend, strip it bare, and move on. As each locust in a single swarm,
consisting of up to 50 billion individuals (5 × 101⁰), can eat its own weight
of food in a day (1.5–2.0 grams), it takes but a few moments to calculate the
amount of food consumed; somewhere between 75,000–100,000 tons.
Today, locust swarms can be tracked by satellite and the use of insecti-
cides and biopesticides at the right time and in the right place can do much
to control outbreaks of certain species. Several studies using modelling and
biochemical analysis have revealed the mechanisms of gregarization. If food
is scarce, a female will do better to lay eggs destined to become solitary in-
dividuals, which are well camouflaged (green or brown in colour) and avoid
one another. However, as population density rises when food is plentiful, it
is more advantageous for locusts to band together and become gregarious. In
the Desert locust, this transition in behaviour occurs rapidly upon crowding
and is induced by touch-sensitive hairs on the hind legs, which, when stim-
ulated by contacting another locust, elicit the release of serotonin within the
CNS, which sets in train behavioural gregarization and ultimately leads to
other changes in colour and shape.
Female locusts use their own experience of being crowded to set the de-
velopmental trajectory of their unhatched embryos. The more recently the
mother was crowded before laying her eggs in the soil, the higher the propor-
tion of gregarious hatchlings that emerge. This ‘epigenetic’ effect is mediated
by a chemical added by the female to the foam surrounding her eggs.
As large and easily cultured insects, locusts have also proved them-
selves to be incredibly useful as model systems for the investigation of
neurophysiology. For instance, they were once thought to be gluttons but
now turn out to be gourmets. Studies on locust feeding behaviour have shown
them to be capable of regulating their food intake to a high degree. Further-
more, their selection of macronutrients in response to their own changing
nutritional needs and a variable nutritional environment has led to the de-
velopment of a geometric framework for considering feeding and nutrition
in insects and other animals, including vertebrates. Indeed, insights gained
from studying locust appetites has revolutionized our understanding of hu-
man obesity and its causes—in particular, the role of a powerful appetite for
protein.
Orthoptera are remarkably well-represented in the fossil record, with the
origins of the order going back to the Late Permian (around 250 mya). Extinct
relatives of Orthoptera appeared even earlier, and some occupied ecological
niches that are now filled by other orders. For example, the Chresmodidae
possessed very long legs and may have walked on water, much like modern
water striders (Hemiptera) do.
Key reading
Anstey, M. L., Rogers, S. M., Ott, S. R., Burrows, M., and Simpson, S. J. (2009). Serotonin
mediates behavioral gregarization underlying swarm formation in desert locusts. Science
323: 627–30.
Bailey, W. J. and Rentz, D. C. F. (eds) (1990). The Tettigoniidae: biology, systematics and
evolution. Crawford House Press, Bathurst.
Bennet-Clark, H. C. (1975). The energetics of the jump of the locust, Schistocerca gregaria.
Journal of Experimental Biology 63: 53–83.
Bennet-Clark, H. C. (1987). The tuned singing burrow of mole crickets. Journal of Experi-
mental Biology 128: 383–409.
Bennet-Clark, H. C. (1989). Songs and the physics of sound production. In: Cricket be-
haviour and neurobiology. Huber, F., Moore, T.E. and Loher, W. (eds), pp. 227–61.
Cornell University Press, Ithaca.
Bennet-Clark, H. C. (1990). Jumping in Orthoptera. In: Biology of grasshoppers. Chapman,
R. F. and Joern, A. (eds), pp. 173–203. John Wiley and Sons, New York.
Bennet-Clark, H. C. (1998). Size and scale effects as constraints in insect sound communi-
cation. Philosophical Transactions of the Royal Society of London B 353: 407–19.
Bidau, C. J. (2014). Patterns in Orthoptera biodiversity. I. adaptations in ecological and
evolutionary contexts. Journal of Insect Biodiversity 2(20): 1–39.
Buhl, J., Sumpter, D. J. T., Couzin, I. D., Hale, J. J., Despland, E., Miller, E. R., and Simpson,
S.J. (2006). From disorder to order in marching locusts. Science 312: 1402–6.
Chapman, R. F. and Joern, A. (eds) (1990). Biology of grasshoppers. John Wiley and Sons,
New York.
Desutter-Grandcolas, L., Jacquelin, L., Hugel, S., Boistel, R., Garrouste, R., Henrotay, M.,
Warren, B. H., Chintauan-Marquier, I. C., Nel, P., Grancolas, P., and Nel, A. (2017). 3-D
imaging reveals four extraordinary cases of convergent evolution of acoustic communi-
cation in crickets and allies (Insecta). Scientific Reports 7: 7099.
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ORTHOPTERA 83
Gage, A. R. and Barnard, C. J. (1996). Male crickets increase sperm number in relation to
competition and female size. Behavioural Ecology and Sociobiology 38(5): 349–53.
Gwynne, D. T. (1984). Sexual selection and sexual differences in Mormon crickets (Or-
thoptera: Tettigoniidae), Anabrus simplex. Evolution 38: 1011–22.
Gwynne, D. T. and Simmons, L. W. (1990). Experimental reversal of courtship roles in an
insect. Nature 346(6280): 172–4.
Gwynne, D. T. (1997). The evolution of edible ‘sperm sacs’ and other forms of courtship
feeding in crickets, katydids and their kin (Orthoptera: Ensifera). In: The evolution of
mating systems in insects and arachnids. Choe, J. C. and Crespi, B. J. (eds), pp. 110–29.
Cambridge University Press, Cambridge.
Huber, F., Moore, T. E., and Loher, W. (1990). Cricket behaviour and neurobiology. Cornell
University Press, Ithaca.
McCaffery, A. R., Simpson, S. J., Islam, M. S., and Roessingh, P. (1998). A gregarizing fac-
tor present in the egg pod foam of the Desert Locust Schistocerca gregaria. Journal of
Experimental Biology 201(3): 347–63.
Miller, G. A., Islam, M. S., Claridge, D. W., Dodgson, T., and Simpson, S. J. (2008). Swarm
formation in the desert locust (Schistocerca gregaria): isolation and NMR analysis of the
primary maternal gregarizing agent. Journal of Experimental Biology 211: 370–6.
Preston-Mafham, K. (1990). Grasshoppers and mantids of the world. Blandford Press,
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Raubenheimer, D. and Simpson, S. J. (2020). Eat like the animals: what nature teaches us
about the science of healthy eating. Houghton Mifflin Harcourt: New York.
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as a device facilitating coercive mating in sagebrush crickets. Proceedings of the Royal
Society of London B 261(1360): 65–71.
Simmons, L. W. (1995). Male bushcrickets tailor spermatophores in relation to their
remating intervals. Functional Ecology 9(6): 881–6.
Simmons, L. W. and Ritchie, M. G. (1996). Symmetry in the songs of crickets. Proceedings
of the Royal Society of London B 263(1375): 1305–11.
Simpson, S. J., McCaffery, A. R., and Hägele, B. (1999). A behavioural analysis of phase
change in the desert locust. Biological Reviews 74: 461–80
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Stritih, N. and Čokl, A. (2013). Mating behaviour and vibratory signalling in non-hearing
cave crickets reflect primitive communication of Ensifera. PLoS One 7(10): e47646.
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Zuk, M. and Simmons, L. W. (1997). Reproductive strategies of the crickets (Orthoptera:
Gryllidae). In: The evolution of mating systems in insects and arachnids. Choe, J. C. and
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84
Dictyoptera
Cockroaches, mantids, and their extinct relatives are united under the super-
order called the Dictyoptera because they share morphological features to do
with wing venation, internal musculature, the structure of the mouthparts,
the male genitalia, and the production of an ootheca. Termites, who were
(until recently) assigned to their own distinct order, have now been shown
to be highly modified cockroaches, based on an overwhelming amount of
molecular and morphological evidence.
85
Blattodea
(cockroaches and termites—alternative name: Blattaria)
Common name Cockroaches and termites Distribution Worldwide

Derivation L. blatta–cockroach Number of families 19
Size Body length 3–100 mm Known world species 7,314 (0.65%)
Metamorphosis Incomplete (egg, nymph,
adult)
Maxillary palp Head directed

downwards Large compound eye
Long
multi-segmented
antenna
Large pronotum
Toughened, leathery
front wings
Strong,
spiny legs
Distinctive
longitudinal
venation
Short cercus
Periplaneta americana — Dictyoptera
Key features
• flattened, broadly oval and leathery-bodied insects (cockroaches)
• front wings toughened as protective ‘tegmina’ to cover the membranous
hind wings
• eggs typically laid in a toughened case
• median ocellus lost
• obligate mutualism with endosymbiont bacteria involved in nitrogen re-
cycling
• soft-bodied, social insects with different castes (termites)
• in soil, litter, wood, or in association with habitations
A female Epilamprinae
cockroach guarding her
© Nicky Bay
nymphs, which are hiding
under her body and wings.
Singapore.
Female Perisphaerus
(Blaberidae) cockroaches
have evolved a body type
similar to that of a pill
millipede and will roll into a
ball when disturbed. Their
© Zestin Soh
nymphs feed on milk-like

secretions from specialized
pouches under their
mother’s thorax. Singapore.
BLATTODEA 87
Cockroaches and termites are of immense ecological importance in tropi-

cal and subtropical ecosystems across the globe, while their behaviour and
physiology have revolutionized our understanding on the evolution of so-
ciality and mutualism in animals. Cockroaches and termites will be described
here separately, due to their highly divergent morphology and habits.
Cockroaches
Cockroaches are fairly uniform in general appearance, being leathery-bodied,
and mostly sombre-coloured insects with an oval outline. Some tropical
day-active species are brightly coloured to advertise distastefulness to poten-
tial predators. The body, when viewed from the side, is flattened, especially
in species living under bark and stones, and this enables them to squeeze
through very tight spaces. The downward-directed head, bearing generalized
chewing and biting jaws, is shielded and largely concealed from above by the
large pronotum. The compound eyes are well developed, there are two ocelli-
like spots, and the long, thread-like antennae are multi-segmented. There are
two pairs of wings, although many species are wingless. The front pair of
wings, called tegmina, are toughened and overlap at rest to cover the larger
fan-shaped, membranous hind wings. Wing venation is mainly longitudinal
in orientation with numerous pale cross veins in the hind wing. Cockroaches
are generally fast runners, and many species fly infrequently. The abdomen,
which is normally completely concealed by the folded tegmina, bears a pair
of sensitive, single- or multi-segmented cerci at the posterior end. A unique
species that departs from this general body plan is the leaproach (genus Salto-
blattella), which has enormous hind legs that allow it to move around by
jumping, much like a grasshopper. Although fossils of extinct cockroach-like
‘roachoids’ of ambiguous affinities were among the most numerous neopteran
insects in the hot, humid coal forests of the upper Carboniferous around 300
mya, the origins of modern cockroaches only go back to the Cretaceous (139–
66 mya), which is considerably younger compared to other insect groups (e.g.,
beetles and wasps).
Cockroaches play an important role in the nitrogen cycle of soil and lit-
ter communities in tropical and subtropical regions, as the vast majority of
species are free-living scavengers and omnivores that eat a wide range of dead
or decaying organic materials, including bird droppings and bat guano, and a
few are able to digest wood. Some species damage growing plants, and canni-
balism and predation also occur. Many cockroaches live in association with
other species and can be found in wasp, ant, or termite colonies, or in the nests
of birds, caves with bats, or the lairs and burrows of mammals, while several
species are semi-aquatic. Cockroaches are an important source of food for
many species, both invertebrate and vertebrate, and are hosts for numerous
parasitic species. Some species of the family Blatellidae are the main pollina-
tors of certain woody climbing plants in the lowland rain forests of Sarawak.
The flowers produce odours of rotting wood or fungi to attract the cock-
roaches, which feed on a secretion produced by the stigma and on the pollen
itself.
In spite of their ecological diversity, cockroaches have been inadequately

studied, as most of the extensive literature concerning these ubiquitous in-
sects deals with effects of pest species and their control. The bad press
regarding the unpleasant habits of certain species is misguided, as in fact,
only 0.005% (or about 40 cockroach species) are considered pests because
of their close association with humans, of which 20 have a significant impact.
Synanthropic species have become widely distributed around the world due
to extensive commercial trade and are associated with warm conditions and
poor hygiene and sanitation. The main problem is that these species are at-
tracted to both filth and foodstuffs, and, as a result, may carry a huge diversity
of pathogenic organisms on their feet and other body parts. The list of diseases
caused by mechanically transmitted bacterial species includes dysentery, di-
arrhoea, and gastroenteritis. They also act as intermediate hosts for a number
of helminths that are pathogenic to vertebrates and are able to harbour and
excrete dangerous viruses and protozoans. When they feed, they regurgitate
partly digested food and leave behind their faeces and a characteristic offen-
sive odour. The exposure, especially of children, to high levels of cockroach
allergens in house dust can produce serious health problems such as allergies,
dermatitis and eczema, and asthma. Insecticides of many types are routinely
used to control pest species, but the development of resistance has gener-
ated interest in integrated pest management using biological control agents,
such as hymenopteran parasitoids and fungal pathogens. The most common
method of control remains the use of toxic sugar baits. However, in a fasci-
nating example of natural selection, populations of the German cockroach,
Blattela germanica, have evolved an aversion for sugar baits, greatly reducing
the traps’ effectiveness. The cockroaches have achieved this by having muta-
tions that modified their taste buds, which make them taste sugary foods as
highly bitter, therefore avoiding them altogether.
Principal cockroach families
Family Distribution Examples of genera

Blaberidae Tropical Blaberus, Gromphadorhina,
Leucophaea, Nauphoeta
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Blattellidae Worldwide Blattella, Parcoblatta
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Pseudophyllodromiidae Mostly Tropical Saltoblattella, Supella
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Blattidae Tropical Periplaneta
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Cryptocercidae N. America and East Asia Cryptocercus
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Nocticolidae Africa, Asia, Australia Nocticola, Spelaeoblatta
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Corydiidae Worldwide Arenivaga, Polyphaga, Tivia
In terms of their general habits, cockroaches are very sensitive to air- and
substrate-borne vibration, and their normal defence is to run and hide, but
some species have warning colouration to signal non-palatability to preda-
tors, and several tropical species mimic the colour patterns of distasteful
beetles, such as lycids, ladybirds, and blister beetles. Many cockroaches, espe-
cially those in the blattid subfamily Polyzosterinae, produce noxious-smelling
BLATTODEA 89
repellent chemicals from ventrally located intersegmental or other glands.

These compounds can produce skin rashes and temporary blindness, and a
number of species, such as the Florida cockroach, Eurycotis floridana, are able
to spray secretions over a distance of many centimetres.
Sexual behaviour ranges from relatively simple to quite complex reper-
toires of behaviour involving sequences of distinctive body movements. In
general, females produce sexual pheromones to which males respond to over
relatively long distances. Males sometimes also produce sexual pheromones
and, when they meet, there is usually much antennal contact and a variety
of body-rocking or trembling movements. The males of many species have
special dorsal abdominal glands that produce aphrodisiac secretions to en-
courage the female. By raising their wings and adopting a particular posture,
the male entices the female to mount his back partially or completely. In this
position the female can lick the tergal gland secretions and the male can en-
gage his genitalia with hers. The female then dismounts and turns to face
backwards to complete the coupling.
In some species, such as members of the blaberid subfamily, Oxyhaloinae,
courtship involves the production of sounds. Males of Nauphoeta cinerea
stridulate by rubbing a ridged portion of the costal veins of the front wing
against similar fine ridges on the hind corners of the pronotum. The Madagas-
can Hissing cockroach, Gromphadorhina portentosa, a large, wingless species
that lives under forest litter, has an unusual mating behaviour. The males,
which are larger than the females and can be up to 10 cm long, have en-
larged thoracic humps with which they batter rivals in head-to-head contests.
The fights are accompanied by a loud hissing produced when air is forcibly
squeezed out of the second pair of abdominal spiracles, and are won by the
biggest, noisiest males. The hissing noises also act as an aphrodisiac to the
females, who will refuse to couple with silent partners.
Sperm is transferred in a spermatophore, and, after mating, female cock-
roaches lay up to forty eggs in two rows surrounded by a tough, protective egg
case or ootheca. Some cockroaches are oviparous and stick their egg cases
to the ground and cover them or carry them about partly projecting from
the end of the abdomen. In all species of the family Blaberidae and a few
in the Blattellidae, females are ovoviviparous. The egg case, which is much
softer, thinner, and more flexible than in oviparous species, is fully extruded
from the end of the abdomen and then rotated by 90◦ so that the long axes
of the eggs within are horizontal. The ootheca is then drawn back inside and
brooded within the body of the female until the eggs hatch.
Ovoviviparity differs from viviparity in that the young do not receive any
nutrients from the mother while retained inside her body. Viviparity, the
most derived reproductive strategy, is seen in the blaberid species Diploptera
punctata, where females produce only half a dozen eggs, which are stored in-
side their abdomen in a ‘uterus’. The developing embryos are supplied with a
unique form of cockroach ‘milk’, which flows through the walls of the uterine
wall. This milk is rich in fats and proteins and has even been considered as
a potential superfood for humans. Female cockroaches in many species are
known to provide varying degrees of maternal care after their young hatch.
In the simplest cases, young nymphs may crowd under their mother’s body
until their cuticle becomes sclerotized. In some species, the young may crawl
into the space beneath the tegmina of the female to find refuge. Females of
the species Thorax porcellana take this approach one step further. Once her
young are born, they hide underneath her wings, on her abdomen. She then
starts making compressing motions with her abdomen, which squeeze a nu-
tritious fluid out of microscopical pores from her abdominal membranes, on
which the nymphs feed. The first instar nymphs are entirely dependent on
their mother for nutrition and protection and perish rapidly if they are re-
moved from her. Perhaps the most peculiar type of maternal care is provided
by the females of Perisphaerus, also known as pill cockroaches. Males have
a typical cockroach appearance, but females are wingless and look remark-
ably like pill millipedes, and they can roll themselves into a ball in a similar
manner. Their nymphs are born blind and depend on their mother of sur-
vival. They cling on the underside of her abdomen, where they insert their
elongated head in a lock and key manner in small holes at the base of their
mother’s legs, which are thought to provide the nymphs with nourishment,
much like suckling mammals. In the presence of predators, the mother is also
able to roll herself into a defensive ball with her nymphs still attached, in order
to ensure their protection.
One of the best-known species, the so-called American cockroach, Peri-
planeta americana (Blattidae), originally spread from Africa on ships and is
now cosmopolitan. Due to its relatively large size and ease of culturing, it has
been used extensively as a laboratory animal for studies ranging from insec-
ticide testing to research on the function of the nervous system. The German
cockroach, Blattella germanica, is probably the most significant single pest
species found in domestic and commercial buildings in many parts of the
world, and considerable efforts are directed towards its control. A single fe-
male, which can live for six months, is capable of producing around 20,000
offspring.
The family Cryptocercidae contains a single genus Cryptocercus with a
handful of unusual species found in North America and East Asia. Crypto-
cercids are sub-social species that live in family groups inside tunnels eaten
out of dead trees. A pair of adults and their developing nymphs stay together
in a family group, and it may take the parents several years to raise a com-
plete brood, feeding the young nymphs and protecting them from enemies.
They are able to eat wood because they have symbiotic protozoans in their
guts to digest the cellulose. At each moult the nymphs have to re-acquire
their symbionts by eating the excrement of their parents or other nymphs. The
protozoans belong to the same genus as those found in early termites (Mas-
totermes: Mastotermitidae); these termites also share with all cockroaches
(except the family Nocticolidae) intracellular bacterial endosymbionts, which
are passed from mother to offspring and are essential for their survival. Both
types of microbial symbionts provided the first indication that they might
have been inherited in cockroaches and termites from a common ancestor—
termites were until recently placed in their own order (Isoptera) and were
thought to be related to cockroaches. The latest molecular studies have now
shown that termites are in fact a lineage of highly modified cockroaches, and
BLATTODEA 91
their closest living relatives are cryptocercids, suggesting that wood feeding
and complex sociality evolved in their common ancestor.
Termites
Termites are undoubtedly the most successful cockroach lineage, and one
of the dominant organisms on the planet. There can be no more impressive
a sight in the African savannahs than the spectacularly large mounds built
by termites in the genus Macrotermes (Termitidae). These towering, multi-
vented chimneys, which can be more than six metres high, provide essential
air conditioning for the nests below ground level, without which the ter-
mites might overheat. An intricate system of tunnels and vents throughout
the structure of the nest maintains optimum temperature and gas levels in
the brood chambers and gardens, where they cultivate a special fungus to eat.
Other equally impressive termite mounds can be seen in northern Australia,
where wedge-shaped nests more than three metres tall are made by the mag-
netic or compass termite, Amitermes meridionalis (Termitidae). The long axis
of the nest always runs roughly north–south, so that the flat surface of the
nest is orientated to act as a solar collector that warms the nest in the morn-
ing and evening. When the sun is at its highest during the heat of the day, the
small surface presented by the top of the nest prevents overheating. Termite
mounds are so effective in keeping cool in spite of the scorching heat that their
structure inspired architects to design buildings in Africa and Australia that
use the same passive cooling mechanism, instead of using air-conditioning.
Confined to tropical and subtropical regions, between 45–50◦ north and
south of the Equator, termites have an immense impact in many habitats.
They may consume up to one-third of the annual production of dead wood,
leaves, and grass, and can be present in huge numbers, sometimes more than
8,000/m2 (800/ft2). Although individually small, their total biomass per unit
area can vary from 1–5 g/m2 to 22 g/m2, which is double that of the biggest
herds of African hoofed grazing mammals. In Amazonian terra firma forests,
termites constitute up to 10% of the biomass of soil fauna, but more than 80%
of the biomass of dead wood fauna.
Termites are sometimes called ‘white ants’ because their workers are pale-
bodied, and because of similarities in their social organization. Termites live
polymorphic in permanent social colonies and are polymorphic, that is, they have a number
having more than two of distinct castes. Termites evolved their social structures much earlier than
forms
ants, but they are, nevertheless, truly social, as they feed, groom, and protect
caste each other, and offspring of one generation help their parents raise the next
any group of individuals in a
generation.
colony of social insects that is
structurally or behaviourally In general, termites are pale, soft-bodied, and wingless with shortish an-
different from individuals in tennae composed of fewer than 32 segments. The rear end of the abdomen
other groups
has a pair of short cerci, just like other cockroaches. A colony, which may
vary in size from a small nest with a couple of dozen individuals, to a mas-
sive, architecturally complex, air-conditioned mound containing millions of
termites, contains four different castes: primary reproductives, supplemen-
tary reproductives, soldiers, and workers. Primary reproductives (the kings
and queens) are large, darkly hardened individuals whose sole job is to repro-
duce. Primary reproductives were once winged termites that left to found the
new colony, and which shed their two pairs of wings after a short dispersal or
nuptial flight. Colonies normally have a single queen and a few reproductive
males. Supplementary reproductives are less hardened than kings and queens
and these will become reproductive if the king or the queen unexpectedly die.
Soldiers are sterile males and females with heavily sclerotized and modified
pear-shaped heads.
© George McGavin
Termites are among the most
abundant and ecologically
important organisms on the
planet. Northern India.
Soldiers have the primary function of defending the colony against predators,
which usually come in the form of termite-hunting ants. Although soldier
termites are usually blind, they are variously armed with large jaws that slice
ants, or they possess the ability to eject sticky secretions from a tube-shaped
head gland that entraps and kills ant invaders. The ongoing arms race be-
tween ants and termites have led some species of the genus Pericapritermes to
develop remarkable ways to defend themselves. The soldiers have peculiarly
shaped, twisted asymmetric mandibles. When the soldiers are attacked by an
ant, their mandibles strike against each other at a speed of 132.4 m/s2, in what
is the fastest movement in the animal kingdom. When the mandibles touch
the ant, the force generated by this ultra-fast impact is so large (4,600 times
the termite’s body weight), that it can dismember ants at a single blow, with
nearly 100% success.
Considering this combative strength, worker termites are rather unim-
pressive in their appearance, and they resemble nymphs. However, their
role is essential in maintaining the colony, and they are the most numer-
A soldier termite ous caste, outnumbering soldiers by 50 to 1. Workers, like soldiers, can be
(Rhinotermitidae).
of both sexes (in contrast to the social Hymenoptera, where workers are all
females). Normally blind with simple, chewing mouthparts, workers build
and repair the nest, forage, feed nymphs, and groom other colony members.
BLATTODEA 93
In the Kalotermitidae, there is no true worker caste, and the jobs are done
instead by older nymphs.
The founding of new colonies happens at certain times of the year with
the production of large numbers of winged reproductives that usually emerge
around sunset. After a short nuptial flight, they settle on the ground, shed
their wings, and females adopt a characteristic stance with their abdomen
pointed up in the air. In this pose they release sexual pheromones to attract
males. A pair will then move off to find a good place nearby in which to bur-
row. They make a small chamber for themselves in soil or wood, seal it, and
mate. The job of rearing the first brood falls to the reproductives, thereafter all
tasks are taken over by workers or nymphs. Soldiers may not be produced un-
til the colony grows larger, and new reproductive individuals may not appear
for many years.
Although some termites eat a range of plant matter, including seeds,
grasses, and leaves, the majority eat dead or decaying wood.
All termites digest cellulose, an ability dependent on symbiotic interac-
A worker termite tions with bacteria, protozoa, and fungi. Six families make up the so-called
(Rhinotermitidae).
lower termites. In these taxa, the gut contains symbiotic flagellate protozoans
trophallaxis to enable cellulose digestion, and they are passed on by trophallaxis via oral
the exchange of food between or anal feeding, just like in their relatives, the cryptocercid cockroaches.
colony members in social
insects In addition to cellulose-digesting symbionts, all termites (except the
fungus-growing Macrotermitinae) have nitrogen-fixing gut bacteria.
Members of the Termitidae, comprising some 70% of all species, digest
cellulose, that is pre-digested either by fungus or by the action of anaerobic
bacteria. The family is made up of four distinct subfamilies. The Apicoter-
mitinae, or soldier-less termites, are humus-feeding species with cultures of
bacteria in their hindgut to ferment plant material. In species belonging to
the genus Globitermes, workers act as soldiers and are able to self-destruct
by bursting their abdomens and covering their enemies (usually ants) with
slimy, sticky gut contents. The Termitinae, or subterranean termites, also
have bacterial cultures in their hindgut to ferment plant material, as do
the Nasutitermitinae (snouted termites). This latter subfamily has strange
snouted soldiers with reduced jaws. Their heads contain well-developed
frontal glands, which produce sticky and noxious chemicals that are ejected at
predators. The Macrotermitinae (fungus-growing termites) construct a per-
manent mud-covered network of tunnels to reach their food. The fungus
gardens are spongy, greyish brown combs made of their own faeces. The fun-
gus, belonging to the genus Termitomyces, is only found inside termite nests,
and it breaks down the faeces and termites that eat it. Fungal hyphae are mixed
with regurgitated food and fed to nymphs. Fungus-growing species are very
important in decomposition processes in seasonal habitats in Asia and Africa,
because, by cultivating the fungus, the termites are able to survive through the
dry season.
Termite families
Family Distribution Habits

Mastotermitidae* 1 species in northern Australia Pest of wooden structures
(Darwin termite) (Mastotermes darwiniensis)* and crops
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Kalotermitidae > 460 species, widespread Prefer dead wood, timber
(dry wood termites) pests
·········································································································································································
Hodotermitidae 21 species in Africa and Asia Grass-harvesters, pasture
(harvester termites) pests
·········································································································································································
Archotermopsidae > 6 Species in North America Prefer decaying wood,
(dampwood termites) and Asia timber pests
·········································································································································································
Serritermitidae 3 species in South America Live in mound walls of
(saw-toothed termites) another termite, food
unknown
·········································································································································································
Stolotermitidae 10 species in Australia, New Prefer decaying wood,
Zealand and South America occasional timber pests
·········································································································································································
Stylotermitidae > 50 species in Indian sub- Inhabit living trees, on which
continent and Southeast they feed
Asia
·········································································································································································
Rhinotermitidae > 300 species, widespread Prefer decaying wood, crop
(subterranean termites) and timber pests
·········································································································································································
Termitidae > 2,070 species, widespread Food variable (grass, wood,
(higher termites) humus, etc.), many pests
* The Darwin termite shares many characteristics with woodroaches (Cryptocercidae), especially in
that they possess similar protozoan symbionts that they transfer in faecal food to the young.
Lower termites have flexible development pathways and there is no true

pseudergate worker caste; this role is taken by a pseudergate caste made up of nymphs.
a caste in lower termites made Moulting in lower termites is complicated and, depending on colony structure
up from immatures who take
the role of workers and the presence of certain pheromones, can produce no change (a station-
ary moult), can produce an individual belonging to a different caste (i.e., a
pseudergate moults to become a reproductive), or can produce a reversion to
an earlier caste. To make it worse, instars can also be added or missed out
entirely.
Termitids have a more elaborate and rigid caste system than other ‘lower’
termites and, in some species, the physogastric queens can grow to 1,000%
of their original size. The abdomen, which is grossly swollen with fat body
and ovaries, can produce several thousand eggs each day. Generally, after
eggs hatch there are two developmental pathways. One leads, through a se-
ries of nymphal stages, to individuals that develop wing buds, and eventually
become reproductives. The other path, known as the sterile pathway, leads
through a series of moults to the production of sterile workers and soldiers.
This pathway can be complicated in some species with the production of two
separate subpathways, one leading to the production of small workers and
small soldiers and the other to the production of large soldiers and workers.
Another way that termites are essential components in ecosystems is that
they are food for a great many other species. Many ant species specialize on
termites as their main prey. Another, less-aggressive approach is followed by
BLATTODEA 95
species of ants that live inside termitaria and defend the termites from attack
by other ants. The relationship is almost certainly mutualistic as the ants re-
ceive shelter (and perhaps food) in return for their services as bodyguards.
Representing such an abundant source of food as they do, it is not at all sur-
prising that termites have many specialized vertebrate predators around the
world. Many bird species eat termites, but also many mammals, such as the
aardvark, aardwolf, pangolins, armadillos, anteaters, sloth bears, and echid-
nas. Chimpanzees spend more than one-quarter of their foraging time all year
round using sticks and branches from particular tree species as tools to extract
termites from their mounds. Two types of tool are used: a sharp-ended stick
the width of a thick pencil and up to 50 cm long to make holes in the walls,
and a thinner, flexible probe to act as a lure. The end of the probe is made
brush-like, and it is pushed into the hole where the soldier termites seize it.
The chimpanzee withdraws the probe and quickly eats the insects. Humans,
very close relatives of chimpanzees, harvest termites as well, especially when
huge numbers of reproductives leave the nest to mate and disperse. They are
a rich food source, providing more than twice as much energy, twice as much
iron and phosphorus, and four times as much calcium per 100 g as lean beef.
Unsurprisingly, in parts of Africa, Australia, Asia, and the Americas, where
termites are abundant, all stages are readily consumed by locals and indige-
nous communities, while their mounds can form important family-owned
resources.
Despite the fact that they are an essential component in tropical terres-
trial food chains and primary nutrient recyclers, and they are considered
the functional equivalent of earthworms in temperate regions, the termites
are (somewhat unfairly) notorious, as many species are destructive pests
and attack structural timbers, wooden buildings, furniture, wooden artefacts,
books, and crops in many areas of the world. In parts of Africa, there may be
800 termite mounds per hectare, which can tie up a huge amount of soil, and
can encourage erosion and subsidence. Recent studies using carbon-14 dat-
ing techniques show that some South African termite nests or termitaria may
have been in existence for 4,000 years. Termites cause huge commercial dam-
age to pasture grasses and crops both above and underground (sugarcane,
potatoes, yams). Termite damage is a major problem in tropical forestry and
explosives and persistent insecticides are routinely used to destroy and poi-
son colonies before areas are planted. The use of resistant tree varieties will
probably be most cost-effective in the long run. Biological control has not
been widely successful so far, but some products containing bacteria that are
genetically engineered to express the Bacillus thuringiensis endotoxin are use-
ful against subterranean termites (Rhinotermitidae), and a number of fungal
pathogens have been used to control termites that damage crops. Another
unusual feature of termites is that, as result of bacterial fermentation in their
hindguts, vast quantities of methane (a greenhouse gas) are released by the
termite’s rear ends to the atmosphere (similarly to ruminants such as farmed
cattle), which represent 1–3% of the world’s methane emissions. However,
unlike cattle and rice paddies, which release harmful amounts of methane,
before they reach the atmosphere more than half of the termites’ methane
emissions are broken down and recycled into the soil by methane-consuming
bacteria in the mound’s walls. Other termite gut symbionts instead release
hydrogen in the atmosphere, as they can convert 1 g of wood into 10 l of
hydrogen, which is harmless for the planet. This process is so effective that
scientists are trying to harness the powers of these symbionts in order to
massively produce hydrogen as an ecologically friendly alternative source of
energy. It is evident that cockroaches and termites are much more than the
annoying pests that most of us think they are, and that, without them, major
ecosystems on the planet would collapse.
Extinct cockroach relatives are among the most abundant insect groups in
the fossil record, especially during the Carboniferous (350–300 mya). These
‘roachoids’ had a similar body plan with modern cockroaches, but many had
important differences, such as a long ovipositor, which is absent in their ex-
tant relatives. Some roachoids even had beak-like mouthparts to perforate
plant tissue, while certain basal cockroaches looked like beetles or praying
mantises. The ancestors of modern cockroaches evolved much later, around
the Jurassic (200 mya). Termites evolved from a Cryptocercus-like ancestor in
the Late Jurassic (about 150 mya) and are unique among insects in having no
extinct families, which attests to the endurance of this group.
Key reading
Barth, R. H. (1968). The mating behaviour of Gromphadorhina portentosa (Schaum) (Blat-
taria, Blaberoidea, Blaberidae, Oxyhaloinae): an anomalous pattern for a cockroach.
Psyche 75(2): 124–31.
Bell, J. W., Roth, L. M., and Nalepa, C. A. (2007). Cockroaches: ecology, behaviour and natural
history. Johns Hopkins University Press, Baltimore.
Bhoopathy, S. (1998). Incidence of parental care in the cockroach Thorax porcellana
(Saravas) (Blaberidae: Blattaria). Current Science 74(3): 248–51.
Bohn, H., Picker, M., Klass, K.-D., and Colville, J. F. (2010). A jumping cockroach from
South Africa, Saltoblattella montistabularis, gen. nov., spec. nov. (Blattodea: Blattellidae).
Arthropod Systematics and Phylogeny 68(1): 53–69.
Cambridge.
Evangelista, D. A., Wipfler, B., Béthoux, O., Donath, A., Fujita, M., Kohli., M. K., Legendre,
F., Liu, S., Machida, R., Misof, B., Peters, R. S., Podsiadlowski, L., Rust, J., Schuette, K.,
Tollenaar, W., Ware, J. L., Wappler, T., Zhou, X., Meusemann, K., and Simon, S. (2019). An
integrative phylogenomic approach illuminates the evolutionary history of cockroaches
and termites (Blattodea). Proceedings of the Royal Society B 286(1895): 20182076.
Grandcolas, P. (1994). Phylogenetic systematics of the subfamily Polyphaginae, with the
assignment of Cryptocercus Scudder to this taxon (Blattaria, Blaberoidea, Polyphagidae).
Systematic Entomology 19: 145–58.
Howse, P. E. (1970). Termites: a study in social behaviour. Hutchinson University Library,
London.
Jacklyn, P. M. (1992). ‘Magnetic’ termite mound surfaces are orientated to suit wind and
shade conditions. Oecologia 91: 385–95.
Krishna, K., Grimaldi, D. A., Krishna, V., and Engel, M.S. (2013). Treatise on the Isoptera of
the world: introduction. Bulletin of the American Museum of Natural History 2013(377):
1–200.
Krishna, K. and Weesner, F. M. (eds) (1970). Biology of termites (2 Vols). Academic Press,
London.
Kuan, K.-C., Chiu, C.-I., Shih, M.-S., Chi, K.-J., and Li, H.-F. (2020). Termite’s twisted
mandible presents fast, powerful, and precise strikes. Scientific Reports 10: 9462.
BLATTODEA 97
Lo, N., Bandi, C., Watanabe, H., Nalepa, C., and Beninati, T. (2003). Evidence for coclado-
genesis between diverse dictyopteran lineages and their intracellular endosymbionts.
Molecular Biology and Evolution 20(6): 907–13.
Lo, N., Tokuda, G., Watanabe, H., Rose, H., Slaytor, M., Maekawa, K., Bandi, C., and Noda,
H. (2000). Evidence from multiple gene sequences indicates that termites evolved from
wood-feeding cockroaches. Current Biology 10(13): 801–4.
Martius, C. (1994). Diversity and ecology of termites in Amazonian forests. Pedobiologia
38: 407–28.
Nalepa, C. A. (1984). Colony composition, protozoan transfer and some life history
characteristics of the woodroach, Cryptocercus punctulatus Scudder (Dictyoptera: Cryp-
tocercidae). Behavioural Ecology and Sociobiology 14: 273–9.
Nalepa, C. A. (1988). Reproduction in the woodroach Cryptocercus punctulatus Scudder
(Dictyoptera: Cryptocercidae): mating, oviposition, and hatch. Annals of the Entomolog-
ical Society of America 81(4): 637–41.
Nauer, P. A., Hutley, L. B., and Arndt, S. K. (2018). Termite mounds mitigate half of termite
methane emissions. Proceedings of the National Academy of Sciences 115(52): 13306–11.
Pearce, M. J. (1997). Termites: biology and pest management. CAB International, Walling-
ford.
Roth, L. M. (1970). Evolution and taxonomic significance of reproduction in Blattaria.
Roth, L. M. (1981). The mother-offspring relationship of some blaberid cockroaches (Dic-
tyoptera: Blattaria: Blaberidae). Proceedings of the Entomological Society of Washington
83: 390–8.
Roth, L. M. and Alsop, D. W. (1978). Toxins of Blattaria. In: Arthropod venoms. Handbook
of experimental pharmacology series 48. Bettini, S. (ed), pp. 465–87. Springer-Verlag,
Berlin/Heidelberg.
Roth, L. M. and Willis, E. R. (1960). The biotic associations of cockroaches. Smithsonian
miscellaneous collections series 141. Smithsonian Institution, Washington, DC.
Schal, C., Gautier, J. Y., and Bell, W. J. (1984). Behavioural ecology of cockroaches. Biological
Reviews 59(2): 209–54.
Schal, C. and Hamilton, R. L. (1990). Integrated suppression of synanthropic cockroaches.
Sreng, L. (1993). Cockroach mating behaviors, sex pheromones and abdominal glands
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Wada-Katsumata, A., Silverman, J., and Schal, C. (2013). Changes in taste neurons support
the emergence of an adaptive behavior in cockroaches. Science 340(6135): 972–5.
Watson, J. A. L. and Abbey H. M. (1985). Seasonal cycles in Nasutitermes exitiosus (Hill)
Isoptera: Termitidae. Sociobiology 10: 73–92.
Wendelken, P. W. and Barth, R. H. (1987). The evolution of courtship phenomena in
neotropical cockroaches of the genus Blaberus and related genera. Ethology 27(Suppl):
98.
Wilde, de, J. and Beetsma, J. (1982). Caste development in social insects. Advances in Insect
Physiology 16:167–246.
Williford, A., Stay, B., and Bhattacharya, D. (2004). Evolution of a novel function: nutritive
milk in the viviparous cockroach, Diploptera punctata. Evolution & Development 6(2):
67–77.
98
Mantodea
(praying mantids)
Common name Mantids Distribution Predominantly tropical

Derivation Gk. mantis–diviner, and subtropical regions
prophet Number of families 15
Metamorphosis Incomplete (egg, nymph,
adult)
Large forward-facing
Thread-like compound eye
antenna
Elongate prothorax
Elongate front coxa
Narrow toughened front

Specialized wings (tegmina)
tibia folds
back on femur
Trochanter
Sharp hook
Front femur with spines
Sphodromantis centralis — Mantodea
MANTODEA 99
Key features
• distinctive highly mobile, triangular head with large, forward-facing com-
pound eyes
• front pair of legs characteristically modified for prey capture
• asymmetric male genitalia
• eggs laid in papery, foam-like, or cellophane-like egg cases
© Anne Riley
A Liturgusa sp. mantis from
Ecuador, feeding on a
cricket.
As their principal
© Anne Riley
prey-capturing appendages,
the mantis’s sickle-shaped
forelegs require regular
cleaning. Brazil.
© Paul Brock
An exquisitely camouflaged
female Empusa pennata
from France.
The common name ‘praying’ mantid comes from the way in which these in-
sects rest with their front legs held up and together as if in prayer. An equally
appropriate homonym would be ‘preying’ mantid, because the features that
separate these insects from their close relatives the cockroaches are mainly
concerned with their highly specialized predatory lifestyle. Mantids are mas-
ters of ambush and rely on being difficult to see. They keep very still or sway
gently to mimic the movement of the foliage, and their colour patterns are
usually sombre greens and browns to blend in perfectly with the background.
Some tropical species may be very brightly coloured and ornamented with
strange cuticular outgrowths and texturing to mimic the flowers upon which
they rest. Certain species take this to the extreme by having a body shape,
colouration, and texture that mimics that of dead or live leaves, sticks, and
grasses. Others, such as Vespamantoidea wherlyi, instead want to be seen—
they mimic the form and colouration of potentially dangerous wasps, so that
predators confuse them with the latter and leave them alone.
Mantids have distinctive triangular and very mobile heads with a pair of
large compound eyes placed laterally and facing forward to give true binocu-
lar vision. Binocular triangulation to calculate the distance of prey objects is
known in many vertebrates, but has also been proved conclusively in mantids,
showing that even the smallest brains can undertake complex sensory tasks.
Other predacious insects, such as tiger beetles and dragonfly larvae, might
also use triangulation when hunting. In addition to the eyes, there may be
MANTODEA 101
three ocelli. The first segment of the thorax is very long indeed and bears the
specialized raptorial, front legs.
The coxa of the front legs is also very elongate, and the femur is enlarged
and variously equipped with rows of sharp spines and teeth. The tibia, which
is also spined or toothed, folds back on the inner face of the femur like a
jackknife.
The middle and hind pair of legs are of normal construction and ap-
pear to arise much further back down the body. The front wings are narrow
and toughened while the fan-folded, hind wings are much larger, membra-
nous, and have a more obvious net-like arrangement of veins, although some
species have short wings or may be wingless. Some mantids can be incredi-
The mobile head of the
bly stick insect-like, but the shape of the head and the raptorial front legs are
mantid (Mantis religiosa) characteristic.
supported by a very Mantids, which are active during the day and some by night, are preda-
elongate prothorax, which
also bears the raptorial cious on a wide range of insects, spiders, and other arthropods, which they
front legs. ambush or stalk. Larger species have even been recorded catching and eat-
ing vertebrates such as salamanders, mice, lizards, and even hummingbirds.
Using triangulation, mantids are able to calculate the exact distance, speed,
and direction of their victim, and can snatch insects out of the air as they fly
past. The speed and elegance of their movements makes mantids popular in-
sect pets and has even inspired a style of karate. The strike, which takes place
Mantid front legs are well
in two distinct phases, lasts less than 100 ms. In the initial phase, the tibiae
equipped with spines and are fully extended in readiness for the second phase, which takes the form of
teeth to impale and hold a rapid sweeping action. The femora are quickly extended, and, at the same
prey.
time, the tibiae are flexed around the prey. It has been shown that some of the
spines projecting from the ventral surface of the femora are moveable and,
when stimulated, elicit a tibial flexion reflex, which helps the insect maintain
a strong grip on struggling prey. The size of prey that mantids can cope with
depends on the species, but, generally, if prey can kick or jump, the mantid
will not attack anything much larger than half its own body weight. Portions
of the prey containing poisonous secretions are left uneaten. However, man-
tises of the genus Carrikerella follow a different hunting approach. The spines
of their raptorial fore tibiae are forwardly oriented and barbed—much like
a spear or a harpoon. While most species use the spines of their raptorial
forelegs to simply restrain their prey, Carrikerella use their foretibial spines to
impale microscopic arthropods that they fish out of small crevices that most
mantises would not be able to access. This allows Carrikerella to hunt insects
of multiple size categories, which makes them more effective predators.
Many species have been shown to be cannibalistic, and mating for the
smaller male can be a dangerous business if he attempts to mate with a non-
receptive female. Females will signal their receptivity by emitting pheromones
and by changes in their behaviour. Nevertheless, males are very cautious to
keep out of the way of the front legs of females, and normally approach and
jump on their prospective mate from behind, keeping low and out of the way
of the female’s front legs. Males do get eaten, even while copulating, and al-
though this happens often in captivity, the extent to which it occurs in the
wild is unclear. Female-biased sex ratios seen in nature might be due to sex-
ual cannibalism but could also be due to higher predation rates on males,
the fact that females live longer, or simply that more female offspring are pro-
duced. However, it matters little to the female if her suitor loses his head. Due
to reflexes in the CNS, the half-devoured male will continue to copulate for
some time ensuring the transfer of his sperm. Eggs are laid within a protective
papery or foam-like ootheca made from abdominal gland secretions. This egg
case contains anything from a dozen to a few hundred eggs, depending on the
species.
In some cases, a balloon of a cellophane-like secretion surrounds an inner
ootheca. The ootheca is fixed to twigs or rock surfaces and, in some species,
is guarded against predators by the female. Several species of parasitic wasps
with appropriately long ovipositors seek out mantid egg cases and will para-
sitize every egg within. In guarding species, maternal care may even continue
A typical mantid egg case,
which may contain a few
for a few days after the young have hatched. On hatching, the very small
hundred developing eggs. nymphs moult again before they start feeding; they become adult after ten
or twelve moults.
Mantids not only have superb eyesight for hunting, but also they have an
additional remarkable sense to avoid being hunted by bats. An ultrasonic-
detecting ‘cyclopean’ ear, responsive to frequencies of between 25 and 45
kHz, is located in a deep groove on the ventral midline of their bodies be-
tween their metathoracic legs. Pressed against the insides of the groove walls
are three pairs of tracheal air sacs, the largest pair of which have a neural
connection. Ultrasound triggers immediate evasive behaviour. Rapid exten-
sion of the front legs and a flick up of the abdomen causes a rapid stall and
forces the insect into a roll or a steep dive. They can recover control from such
an extreme manoeuvre, but if the pursuing bat is too close, they will crash-
land without injury. Predation pressure by bats to flying mantises must be
considerable, as cyclopean ears are present in many mantises and may have
evolved independently multiple times. They are frequently best developed in
only the sex that flies the most (usually the male) while short-winged flightless
species and females usually have reduced or no auditory capability. However,
cyclopean ears are maintained in certain flightless species and may be used
in communication, prey detection, or the avoidance of different predators, al-
though we currently have no concrete proof. Recently, a species of mantid has
been shown to have additional, equally sensitive hearing at lower frequencies
(2–4 kHz), but the reason for this is not yet known. The sensitivity of the man-
tid system is similar to bat-detecting devices found in crickets and lacewings,
but less sensitive than the tympanal hearing organs of moths.
Mantids also need to defend themselves against birds, reptiles, and mam-
mals. They can run, fly, and hide, but also use threat postures, holding their
front legs out sideways to look bigger and displaying bright markings on the
inside of the femora. No idle threat, a strike from a mantid can easily draw
blood or damage an eye, and a predator might be persuaded to seek an easier
autotomy
the casting off of a part of the
meal elsewhere. Like stick insects, mantids will shed a leg if trapped, but auto-
body, for example, when tomy rarely involves the front legs, without which the insect would eventually
under threat die of hunger.
MANTODEA 103
As to the origins of mantids, their earliest fossils date back to the Creta-
ceous, although they started diversifying considerably only from the Tertiary
onwards. The Alienopteridae, a remarkable family discovered in Cretaceous
Myanmar amber, is thought to represent a lineage that is either ances-
tral to Mantodea, or its sister group. Alienopterids are unique in that they
possess characteristics present in both Mantodea and cockroaches, further
reaffirming the sister-group relationship between the two orders.
Key reading
Bai, M., Beutel, R.G., Klass, K.-D., Zhang, W., Yang, X., and Wipfler, B. (2016).
Alienoptera—A new insect order in the roach—mantodean twilight zone. Gondwana
Research 39: 317–26.
Grimaldi, D. (2003). A revision of Cretaceous mantises and their relationships, including
new taxa (Insecta: Dictyoptera: Mantodea). American Museum Novitates 3412: 1–47.
Johns, P. M. and Maxwell, M. R. (1997). Sexual cannibalism: who benefits? Trends in Ecology
and Evolution 12: 127–8.
Reitze, M. and Nentwig, W. (1991). Comparative investigations into the feeding ecology of
six Mantodea species. Oecologia 86: 568–74.
Rivera, J. and Callohuari, Y. (2020). A new species of praying mantis from Peru reveals
impaling as a novel hunting strategy in Mantodea (Thespidae: Thespini). Neotropical
Rosner, R., von Hadeln, J., Tarawneh, G., and Read, J. C. A. (2019). A neuronal correlate of
insect stereopsis. Nature Communications 10: 2845.
Roussel, S. (1983). Binocular stereopsis in an insect. Nature 302: 821–2.
Yager, D. D. (1990). Sexual dimorphism of auditory function and structure in praying
mantises (Mantodea: Dictyoptera). Journal of Zoology 221: 517–38.
Yager, D. D. (1996). Serially homologous ears perform frequency fractionation in the pray-
ing mantis, Creobroter (Mantodea, Hymenopodidae). Journal of Comparative Physiology
A, Sensory, Neural and Behavioural Physiology 178: 463–75.
Yager, D. D. and Hoy, R. R. (1986). The cyclopean ear: a new sense for the praying mantis.
Science 231: 727–9.
Wieland, F. (2013). The phylogenetic system of Mantodea (Insect: Dictyoptera): species,
phylogeny and evolution. Universitätsverlag Göttingen: Göttingen.
Wieland, F. & Svenson, G. J. (2018). Biodiversity of Mantodea. In: Insect biodiversity: science
and society, II. Foottit, R. G. and Adler, P. H. (eds), pp. 389–416. John Wiley and Sons,
Hoboken.
104
Xenonomia
This is one of the most obscure clades of insects, which comprises two
wingless orders—the ice crawlers (Grylloblattodea), and the heel walkers
(Mantophasmatodea), which are the most recently described order of insects.
Some classifications unite both groups under the single order Notoptera, al-
though the latest molecular and morphological studies validate their status as
separate, deeply divergent orders.
105
Grylloblattodea
(ice crawlers—alternative name: Notoptera)
Common name Ice crawlers or rock Metamorphosis Incomplete (egg, nymph,

crawlers adult)
Derivation Gk. combination of cricket Distribution Cooler regions of North
(gryllus) and cockroach America and north-east
(blatta) Asia
Size Body length 14–43 mm Number of families 1 (Grylloblattidae)
Known world species 33 (0.003%)
Thread-like antenna
Small compound
eyes
Squarish
pronotum
Wingless
Elongated
abdomen
Slender cercus
Ovipositor
Grylloblatta campodeformis —
Grylloblattodea
Key features
• eyes reduced or absent, ocelli absent
• wingless
• eversible vesicles on first abdominal segment
• confined to certain cold, high-altitude regions
Grylloblattodea, one of the

smallest insect orders, can
only survive in very damp
© Piotr Naskrecki
and cold environments. This
species, Galloisiana
nipponensis lives
underground, in the wet and
cool forests of Japan.
These slender, wingless, slightly hairy, grasshopper-like insects were first dis-
covered in the Canadian Rockies in 1913. They were originally thought of as
being a small ‘primitive’ family of the Orthoptera, but a number of earwig-,
stonefly-, cockroach-, and even silverfish-like features made it inevitable that
they should have an order all of their own. For a long time, there was much
disagreement on exactly where these ‘living fossils’ fit in relation to the other
polyneopteran orders.
The head has small, compound eyes, although these are sometimes absent;
slender, thread-like antennae; and simple chewing, forward-facing mouth-
parts. There are no ocelli. The pronotum is square and the abdomen is
cylindrical with a pair of slender, multi-segmented cerci.
Species in only five genera are currently known: Galloisiana recorded from
China, Japan, and South Korea; Grylloblattina from South Korea; Grylloblatta
from western parts of North America and Canada; Grylloblatella from China
and southern Siberia; and Namkungia from South Korea.
Grylloblattodea can be found in a range of different habitats and eleva-
tions, although all are characterized by a cool climate with high humidity.
South Korean species of Galloisiana have been found in limestone caves,
whereas, in Japan, the commonest species Galloisiana nipponensis is a moun-
tain dweller and lives among rocks, under stones, and in decaying wood
in subalpine, deciduous forests up to 2000 m. The dependence of Gryl-
loblattodea on cool and glacial habitats means that most populations are
fragmented and could be severely affected by climate change.
GRYLLOBLATTODEA 107
Not much is known about the behaviours of Grylloblattodea. Mat-

ing, which has been observed in some Galloisiana species, involves some
preliminary tactile courtship and antennal fencing. The male then grips the
female’s prothorax with his mandibles and mounts her back. He then twists
his abdomen round to engage his genitalia and partially slides off sideways
(invariably to his mate’s right-hand side). The two may remain in this posi-
tion for up to four hours. The female may not lay her eggs for another ten
weeks or more. It is thought that there may be eight nymphal instars, and
complete nymphal development may take up to five or six years. As nymphs
age they become darker coloured and add segments to their antennae at each
moult. The adults typically live for fewer than two years.
With a few exceptions, most species of Grylloblatta are extremophiles
adapted to life in the cold conditions of mountainous regions, where they can
be found moving about on the surface of snow and ice, particularly after dark.
During the day they can be found in rock crevices or under snow. Nymphs
and adults are primarily scavengers, their preferred food being dead or torpid
insects and other small arthropods that they find. Wind-borne insects, par-
ticularly aphids, that end up as fallout on the surface of snow in mountainous
regions are the most important source of food for rock crawlers. There is some
evidence that they feed on a range of organic matter, including moss and, per-
haps, some plant material, and, in laboratory cultures, they are cannibalistic.
In the summer months, they retreat underground to avoid the desiccating
conditions. Females use a short ovipositor to lay relatively large, dark eggs in
moss or soil and these may enter diapause for a year. Species are distinguished
on the structure of the male genitalia. Life history details of most species are
not well known. Due to their reliance for low temperatures, climate change
may pose a severe challenge for the survival of ice crawlers across the globe.
The origins of Grylloblattodea are very obscure, as they are very scarce in
the fossil record. Currently available fossils from the Permian and the Jurassic
may represent groups that are only distantly related to modern ice crawlers
and may not be that informative on their evolution. However, morphological
data and phylogenomics overwhelmingly support a sister-group relationship
to the Mantophasmatodea, placing the origin of both orders to the Jurassic
(around 170 mya).
Key reading
Jarvis, K. J. and Whiting, M. F. (2006). Phylogeny and biogeography of ice crawlers (In-
secta: Grylloblattodea) based on six molecular loci: designating conservation status for
Grylloblattodea species. Molecular Phylogenetics and Evolution 41(1): 222–37.
Kamp, J. W. (1979). Taxonomy, distribution and zoogeographic evolution of Grylloblatta in
Canada (Insecta: Notoptera.) Canadian Entomologist 111: 27–38.
Kim, B.-W. and Lee, W. (2007). A new species of the genus Galloisiana (Grylloblattodea,
Grylloblattidae) from Korea. Zoological Science 24: 733–45.
Schoville, S. D., Simon, S., Bai, M., Beethem, Z., Dudko, R. Y., Eberhard, M. J. B., Frandsen,
P. B., Küpper, S. C., Machida, R., Verheij, M., Willadsen, P. C., Zhou, X., and Wipfler,
B. (2021). Comparative transcriptomics of ice-crawlers demonstrates cold specialization
constrains niche evolution in a relict lineage. Evolutionary Applications 14(2): 360–82.
Wipfler, B., Bai, M., Schoville, S., Dallai, R., Uchifune, T., Machida, R., Cui, Y., and Beutel,
R. G. (2014). Ice crawlers (Grylloblattodea)—the history of the investigation of a highly
unusual group of insects. Journal of Insect Biodiversity 2(2): 1–25.
108
Mantophasmatodea
(heel walkers—alternative name: Notoptera)
Common name Heel walkers or gladiators Distribution Southern and east Africa
Derivation Gk. combination of (Namibia, South Africa,
praying mantis and stick Tanzania)
insect (phasma) Number of families 3 (Austrophasmatidae,
Size Body length up to 35 mm Mantophasmatidae,
Metamorphosis Incomplete (egg, nymph, Tanzaniophasmatidae)
adult) Known world species 20 (0.0018%)
Cylindrical body
Tarsi five-segmented
Short cerci
Tyrannophasma gladiator — Mantophasmatodea
MANTOPHASMATODEA 109
Key features
• wingless
• mantis-like predatory head, ocelli absent
• tarsal pads distinctly enlarged and held upwards
• drumming organ on male abdomen
• confined to arid and semi-arid regions of Southern and East Africa
Mantophasmatodea are the

most recently described
extant insect order. They
resemble stick insects and
mantids, hence the name.
They are also called
© Mike Picker
heelwalkers, as their tarsal
arolia are always held
upwards. Sclerophasma sp.,
South Africa.
Heel walkers represent the most recently discovered insect order, only found
in 2002. Since their initial discovery, a comprehensive body of literature has
described their behaviour, morphology, physiology and evolution, rendering
them one of the best-known orders of insects.
The details of their discovery are fascinating. The first known specimen
was described from Eocene Baltic amber (48–34 mya) collected in 1992, as
belonging to an undetermined order. It was only in 2002, when a series of
unidentified museum specimens collected in the 1950s allowed the exami-
nation of additional characters that made it clear that these insects are part
of a previously unrecognized group. Mantophasmatodea are the sister group
to Grylloblattodea, with which they share a common ancestor in the Jurassic
(about 170 mya).
Heel walkers bear a superficial resemblance to stick insects and praying
mantises. Mantophasmatodeans walk in a characteristic manner, where their
enlarged tarsal pads (arolia) are held upwards to protect them, with their
‘heels’ being the only part touching the ground—hence their common name,
heel walkers. This is a feature shared with many stick insects (Phasmatodea).
All heel walkers are medium sized with a slender body. Adults are wingless,
which made some researchers to initially believe that they are nymphs of
winged insects. Their head is triangular, mantis-like, and facing vertically, and
it is equipped with powerful mandibles that are used to chew small insects.
The antennae are long. All of their legs are fairly long and slender, with the
fore and middle tibiae bearing small spines that assist heel walkers in catch-
ing and restraining small insects. The abdomen bears one-segmented cerci,
and the male genitalia are asymmetrical.
Much is known about the habits of Mantophasmatodea. They generally
inhabit shrubland and grassland areas in the arid and semi-arid regions in
southern and eastern Africa. All life stages are found on grasses, bushes and
short trees. They have not been found in forested areas, and seem to occupy
habitats at varying altitudes, with particular diversification in the Fynbos
biome. All species are well-camouflaged, being green, brown, yellow, and
grey, which allows them to blend with their surroundings. Their colour can
gradually change if they are placed on a differently coloured environment.
All mantophasmatodeans are generalist predators of small and medium-
sized insects, which they typically hunt during the evening and at night-time.
They use their armed fore- and mid-legs to grasp their prey, which they
quickly devour using sharp mandibles. Their enlarged tarsal pads enable
them to hold on to surfaces while their other legs are busy restraining their
struggling prey.
When it comes to courtship, heel walkers likely use pheromones to attract
mates from long distances. Once in closer range, both sexes communicate
and locate each other with low-frequency vibrational signals, which they de-
tect using highly sensitive scolopidial organs on their legs. The vibrations are
produced by a distinct drumming behaviour, where the abdomen is rhyth-
mically tapped against the substrate. In males, the tip of their abdomen has
a lip-like spine which they use to tap the substrate—much like a drumstick.
Like praying mantises, the male may run a risk of being eaten by the female
and tries to mount her cautiously and rapidly.
Oviposition takes place on small holes in sandy soil, where the eggs are
deposited inside a foam that hardens and incorporates sand particles to form
a protective pod. Females lay multiple egg pods, each containing between 10–
30 eggs, depending on the species. The egg pods hatch after approximately
eight months, with the first rains that mark the end of the dry season. Due
to the highly seasonal environments they inhabit, heel walkers are generally
short-lived. Once hatched, the nymphs mature in five months, and the adults
survive for only a couple more months.
Key reading
Eberhard, M. J. B., Lang, D., Metscher, B., Pass, G., Picker, M. B., and Wolf, H. (2010). Struc-
ture and sensory physiology of the leg scolopidial organs in Mantophasmatodea and their
role in vibrational communication. Arthropod Structure & Development 39(4): 230–41.
Eberhard, M. J. B. and Picker, M. D. (2008). Vibrational communication in two sympatric
species of Mantophasmatodea (Heel walkers). Journal of Insect Behaviour 21: 240.
Huang, D., Nel, A., Zompro, O., and Waller, A. (2008). Mantophasmatodea now in the
Jurassic. Naturwissenschaften 95: 947–52.
Klass, K.-D., Zompro, O., Kristensen, N. P., and Adis, J. (2002). Mantophasmatodea: a new
insect order with extant members in the Afrotropics. Science 296: 1456–9.
Roth, S., Molina, J., and Predel, R. (2014). Biodiversity, ecology, and behavior of the recently
discovered insect order Mantophasmatodea. Frontiers in Zoology 11: 70.
Terry, M. D. and Whiting, M. F. (2005). Mantophasmatodea and phylogeny of the lower
neopterous insects. Cladistics 21(3): 240–57.
MANTOPHASMATODEA 111
Wipfler, B., Letsch, H., Frandsen, P. B., Kapli, P., Mayer, C., Bartel, D., Buckley, T. R.,
Donath, A., Edgerly-Rooks, J. S., Fujita, M., Liu, S., Machida, R., Mashimo, Y., Misof,
B, Niehuis, O., Peters, R. S., Petersen, M., Podsiadlowski, L., Schütte, K., Shimizu, S.,
Uchifune, T., Wilbrandt, J., Yan, E., Zhou, X., and Simon, S. (2019). Evolutionary his-
tory of Polyneoptera and its implications for our understanding of early winged insects.
Proceedings of the National Academy of Sciences of the United States of America 116:
3024–9.
Wipfler, B., Theska, T., and Predel, R. (2018). Mantophasmatodea from the Richtersveld in
South Africa with description of two new genera and species. ZooKeys 746: 137–60.
112
Eukinolabia
This group includes two morphologically divergent orders: the stick insects
(Phasmatodea) and the webspinners (Embioptera). Their close relationship
was first proposed based on structures of their mouthparts and their eggs,
which was then confirmed by a suite of morphological and phylogenomic
studies.
113
Phasmatodea
(stick insects and leaf insects)
Common name Stick and leaf insects, Metamorphosis Incomplete (egg, nymph,
walking-sticks adult)
Derivation Gk. phasma–apparition, Distribution Mainly tropical and
spectre subtropical
Size Body length up to 360 Number of families Uncertain
mm. Mostly 10–100 mm Known world species Uncertain
Pronotum
Hollow in femora to
allow front leg to be
extended forwards
Widely spaced legs

Mesonotum
Metanotum Elongate stick-like body
Lonchodes sp. — Phasmatodea
Key features
• slender, stick-like, or broad, leaf-like body with widely spaced legs
• slow moving and herbivorous
• confined to vegetation, which many mimic
Stick and leaf insects are
© Rupert Soskin
masters of camouflage, such
as this Phyllium sp., which
perfectly imitates the shape,
colour, and veins of a leaf.
Phasmatodea may exhibit

marked sexual dimorphism
in size, colouration, and
© Anne Riley
structure, as can be seen

from these mating
Spinopeplus sp. from
Sumaco, Ecuador.
Stick insects are mostly nocturnal, slow moving and herbivorous. Their char-
acteristic stick-like or leaf-like bodies, the cryptic colouring, and the way they
move make them very difficult to see among foliage and afford them a high
degree of protection from predators.
PHASMATODEA 115
The relationships within stick and leaf insects still remain poorly under-
stood, hence the uncertainty in the number of families—some classification
systems propose as few as three, while others up to 30. The evolutionary ori-
gins of Phasmatodea are also hotly debated, with some studies suggesting they
are among the youngest insect orders dating back to the Late Jurassic–Early
Cretaceous (about 150–120 mya), and others proposing a significantly older
origin in the Permian–Triassic (more than 250 mya). What is more certain, is
that visual predators have driven the remarkable evolution of camouflage in
Phasmatodea, and there is strong evidence that most modern stick and leaf
insects have diversified their morphology as a result of bird predation.
Some species freeze motionless when disturbed, holding the middle and
hind legs tightly along the body and stretching the front legs out. Others
sway gently, imitating the movements of the vegetation. The leaf insects are
contained in one group, the Phylliinae, of which there are about 50 species
confined to southeast Asia, New Guinea, and Australia. Leaf insects are broad
and flattened, brown or green, and with fantastic leaf-like expansions on all
leg and body segments.
The disguise is further improved by surface texturing, and blotches of
colour or mottling, resembling leaf damage and veining on the short, front
wings.
The body of stick insects can be very variable from short and smooth to
large and very spiny. Many species have thorny or leafy outgrowths on vari-
ous body parts to break up the outline of the insect and to help it blend in with
the background. The head is often characteristically domed and carries quite
long, thread-like antennae, simple chewing mouthparts, a pair of small, lat-
erally placed compound eyes, and, in winged species, ocelli. The front wings
(tegmina) are short, toughened, and, at rest, slightly overlapped to protect the
much larger fan-shaped, membranous hind wings. Many species have very
short wings or none at all. The ovipositor of the females is very short and
The characteristic leaf-like
bodies of leaf insects make often difficult to see. The abdomen carries a pair of small, terminal cerci.
them very difficult to see Males are generally smaller than females, sometimes differently shaped,
among foliage and afford
them a high degree of and have one less moult. Mating in phasmids is, generally, rather a dull af-
protection from predators. fair. The male is presumably attracted by a sexual pheromone emitted by the
female, and then simply mounts his mate, engages his genitalia by twisting
his abdomen round underneath, and hangs on. Males may remain mounted
for many days; in the genus Timema (Timematidae), males may spend their
entire adult lives being carried about by the much larger female. A bit of
excitement is seen in the North American species Diapheromera veliei (Phas-
matidae), where competing males fight each other, kicking out with spined
legs as they try to gain exclusive access to a mature female. Parthenogenesis
is common in stick insects, and, in some species, males are very rare or even
unknown.
The eggs of stick insects look remarkably like seeds in size, surface tex-
ture, colour, and shape, and may be dropped, scattered, stuck to vegetation,
Stick insect eggs look very or lightly covered with sand or soil. In species where the eggs are dropped
seed-like and have a
protuberance called a
to the ground, rather than being glued to vegetation or buried, the eggs have
capitulum. protuberance called a capitulum. The resemblance between these eggs and
seeds of higher plants, whose seeds have elaiosomes, is striking and for good
reason. Both capitula-bearing eggs and elaiosome-bearing seeds attract ants
that take them back to their nests. Here, seeds and eggs are safe from herbi-
vores and parasitic wasps, respectively. Nymphs, which look very much like
the adults (especially in wingless species), may go through as few as two or as
many as eight instars before becoming mature.
Although the main defensive strategy in stick insects is crypsis and death
feigning, some will jump and fall from the foliage if disturbed by predators,
such as birds. Others flash bright colours on their hind wings to startle ene-
mies, adopt threatening scorpion-like postures by raising their abdomen, or
they produce hissing sounds. A few species possess thoracic glands, which
can produce noxious substances to repel ants, and even vertebrate predators,
and they may be brightly coloured to advertise this ability. The hind femora
of Eurycantha males from New Guinea have a very large inner spine and a re-
flex closing action, which can trap and crush. The spine is so hard and sharp
that it is used as a fishing hook. An interesting feature of many stick insects is
that, if attacked, they can shed their legs and partially regenerate them during
subsequent moults. Autotomy of limbs takes place at a fracture line at the base
of the legs, between the femur and the trochanter, and a special diaphragm
seals off the wound rapidly to prevent loss of body fluids. Many species, such
as Carausius morosus, can change colour. Epidermal cells contain granules
of pigment, which are able to come together or disperse. When it is hot and
sunny, the pigment clumps making the animals pale. When it is cool and
overcast, the granules disperse, making the surface appear dark and allowing
the insect to absorb more heat. Becoming darker in colour at night is also a
good antipredator strategy.
Most species are of no economic significance, but a few species of stick
insect can be pests. For example, the Australian spur-legged phasmid Didy-
muria violescens is responsible for defoliation in highland Eucalyptus forests.
Many species, such as Macleay’s Spectre (Extatosoma triaratum), Thailand
stick insects (Bacillus spp.), the Pink-winged stick insect (Sipyloidea sipylus),
the Jungle Nymph (Heteropteryx dilatata), and the Giant Spiny stick insect
(Eurycantha calcarata), are popular as pets, and all can be easily reared on a
diet of fresh bramble leaves.
Key reading
Bässler, U. (1983). Neural basis of elementary behavior in stick insects. Studies of brain
function series 10. Springer-Verlag, Berlin.
Bradler, S. (2009). Die Phylogenie der Stab- und Gespenstschrecken (Insecta: Phasmatodea).
Species, phylogeny, and evolution series 2. University of Göttingen Press, Göttingen.
Brock, P. D. (1999). The amazing world of stick and leaf insects. The amateur entomologist
series 26. A.E.S. Publications.
Brock, P. D. (1991). Stick-insects of Britain, Europe and the Mediterranean. Fitzgerald
Publishing, London.
Bedford, G. O. (1978). Biology and ecology of the Phasmatodea. Annual Review of Ento-
mology 23: 125–30.
Hughes, L. and Westoby, M. (1992). Capitula on stick insect eggs and elaiosomes on seeds:
convergent adaptations for burial by ants. Functional Ecology 6: 642–8.
PHASMATODEA 117
Robertson, J. A., Bradler, S., and Whiting, M. F. (2018). Evolution of oviposition techniques
in stick and leaf insects (Phasmatodea). Frontiers in Ecology and Evolution 6: 216.
Schmitz, J. (1993). Load compensating reactions in the proximal leg joints of stick insects
during standing and walking. Journal of Experimental Biology 183: 15–33.
Simon, S., Letsch, H., Bank, S., Buckley, T. R., Donath, A., Liu, S., Machida, R., Meusemann,
K., Misof, B., Podsiadlowski, L., Zhou, X., Wipfler, B. and Bradler, S. (2019). Old World
and New World Phasmatodea: phylogenomics resolve the evolutionary history of stick
and leaf insects. Frontiers in Ecology and Evolution 7: 345.
Sivinski, J. (1978). Intrasexual aggression in the stick insect Diapheromera veliei and D.
covilleae and sexual dimorphism in the Phasmatodea. Psyche 85: 395–403.
Strauß, J., von Bredow, C.-R., von Bredow, Y. M., Stolz, K., Trenczek, T. E., and Lakes-Harlan,
R. (2017). Multiple identified neurons and peripheral nerves innervating the prothoracic
defense glands in stick insects reveal evolutionary conserved and novel elements of a
chemical defense system. Frontiers in Ecology and Evolution 5: 151.
Tihelka, E., Cai, C., Giacomelli, M., Pisani, D., and Donoghue P. C. J. (2020). Inte-
grated phylogenomic and fossil evidence of stick and leaf insects (Phasmatodea) reveal
a Permian–Triassic co-origination with insectivores. Royal Society Open Science 7(11):
201689.
118
Embioptera
(webspinners—alternative name: Embiidina)
Common name Webspinners Metamorphosis Incomplete (egg, nymph,

Derivation Gk. embios–having life, adult)
pteron–a wing Distribution Mainly tropical; some in
Size Body length 3–25 mm. warm temperate regions
Mostly under 12 mm Number of families 13
Known world species 400 (0.035%)
Mouthparts
facing forwards
Small compound eye
Swollen basal
Meso- and metathorax tarsal segment
broad (males only)
Narrow wing
(males only)
Short, stout hind legs
Elongate, cylindrical body
Cercus
Antipaluria caribbeana — Embioptera
EMBIOPTERA 119
Key features
• gregarious in silk galleries
• swollen first tarsal segment contains silk glands
• hind femur distinctly swollen
Embioptera are unique
© Piotr Naskrecki
among insects in that they
produce silk from their
enlarged front tarsi, as can
be seen in this female Embia
sp. from Mozambique.
Web spinners are pale or dark-coloured, gregarious or sub-social insects who

get their common name from their ability to make elaborate (and some-
times extensive) silk tunnels and galleries in soil, litter, and on or beneath
bark. The tunnels protect the web spinners and their young from enemies,
desiccation, and rain. They are narrow-bodied, elongate, and cylindrical or
slightly flattened, with small, kidney-shaped eyes, no ocelli, antennae with
10–35 segments, and simple, forward-pointing, biting mouthparts. Both sexes
are equipped with swollen, muscular hind legs, which, when threatened, allow
them to rapidly move backwards inside their tunnel without having to turn.
The remaining legs are short, and the front legs of both sexes and nymphs
are characterized by having swollen basal tarsal segments, which contain
numerous silk glands.
The liquid silk produced by each secretory gland passes through a minute
tubule to one of about a hundred hollow, hair-like or bristle-like silk ejec-
tors on the ventral surface. The mid-tarsal segments also have silk glands and
ejectors but are not swollen. As the front feet are rubbed to and fro against
the substrate, silk strands are pulled from the ends of the ejectors to build up
a mat of silk. As the colony grows, galleries and tunnels are extended to take
in new food sources such as dead plant material, litter, lichens, and mosses.
Only the adult females and the nymphs, which they resemble, feed. Males do
not feed as adults, and their forward-facing jaws are used to grasp the female
during copulation. The only individuals to leave the confines of the colony
are mature males who disperse to find mates elsewhere. Females are wing-
less, but the males usually have two equally sized pairs of long, narrow wings,
which are very unusual in that they have certain hollow veins which can be
inflated with haemolymph to make them stiff for flight. When the veins are
not inflated, the wings can fold forwards to enable the male to run backwards
through the galleries (if necessary) without damaging them.
When males locate females, they typically court each other with low fre-
quency substrate vibrations. After mating, eggs are laid in the galleries and
the female builds a protective nest over them composed of silk mixed with
chewed food and debris. The female will remain nearby and guard the eggs
against parasites and other females competing for oviposition sites. Some
species have a rich repertoire of territorial vibrational signals that they pro-
duce against female competitors and unwanted males, which allows them to
communicate over long distances without having to leave their oviposition
site. Once the eggs hatch, the female must break open the nest to release her
young. In some species, maternal care extends to providing the first instar
nymphs with a paste of masticated food. Some species are parthenogenetic.
Despite the protection of their silk tunnels and cryptic habits, web spinners
are attacked by ants and parasitic wasps. The very few known species in the
bug genus Embiophila (Plokiophilidae) live in association with web spinners
and suck out the eggs, and dead or dying larvae and adults. The web spinners
do not seem to mind their presence; therefore, it is likely that they get some-
A typical adult female web thing in return. Ten species of wasp in the family Sclerogibbidae are specialist
spinner with a swollen ectoparasitoids of web spinner nymphs. The larvae feed on their hosts through
tarsal segments containing
silk glands.
the intersegmental membranes and then pupate in the web spinners’ tunnels.
The systematics of Embioptera are still poorly understood, and most of the
current families have been shown to be rather artificial. Furthermore, their
cryptic habits make them difficult to collect, and they are generally scarce in
most entomological collections. It is likely that at least 1,500 species remain
undescribed, which outnumber described species by more than 3 times. This
highly specialized insect order is in dire need of further study. Fossils of web
spinners are also scarce, with the oldest representatives of the order having
been found in Jurassic deposits.
Key reading
Büsse, S., Hörnschemeyer, T., Hohu, K., McMillan, D., and Edgerly, J. S. (2015). The spin-
ning apparatus of webspinners—functional-morphology, morphometrics and spinning
behaviour. Scientific Reports 5: 9986.
Choe, J. C. (1994). Communal nesting and subsociality in a webspinner, Anisembia texana
(Insecta: Embiidina: Anisembiidae). Animal Behaviour 47: 971–3.
Dejan, K. A., Fresquez, J. M., Meyer, A. M., and Edgerly, J. S. (2013). Maternal territori-
ality achieved through shaking and lunging: an investigation of patterns in associated
behaviors and substrate vibrations in a colonial embiopteran, Antipaluria urichi. Journal
of Insect Science 13: 82.
Edgerly, J. S. (1988). Maternal behaviour of a web-spinner (Order: Embiidina): mother-
nymph associations. Ecological Entomology 13: 263–72.
Edgerly, J. S. (1994). Is group living an antipredator defense in a facultatively communal
webspinnner (Embiidina: Clothodidae)? Journal of Insect Behaviour 7: 135–47.
Edgerly, J. S., Büsse, S., and Hörnschemeyer, T. (2012). Spinning behaviour and morphology
of the spinning glands in male and female Aposthonia ceylonica Enderlein, 1912 (Em-
bioptera: Oligotomidae). Zoologischer Anzeiger—A Journal of Comparative Zoology 251:
297–306.
EMBIOPTERA 121
Hodson, A. M., Cook, S. E., Edgerly, J. S., and Miller, K. B. (2014). Parthenogenetic and sex-
ual species within the Haploembia solieri species complex (Embioptera: Oligotomidae)
found in California. Insect Systematics and Evolution 45: 93–113.
Huang, D.-Y. and Nel, A. (2009). Oldest webspinners from the Middle Jurassic of Inner
Mongolia, China (Insecta: Embiodea). Zoological Journal of the Linnean Society 156:
889–95.
Miller, K. B., Hayashi, C., Whiting, M. F., Svenson, G. J., and Edgerly, J. S. (2012). The
phylogeny and classification of Embioptera (Insecta). Systematic Entomology 37: 550–70.
Ross, E. S. (1970). Biosystematics of the Embioptera. Annual Review of Entomology 15:
157–72.
Ross, E. S. (1987). Studies on the insect order Embiidina: a revision of the family Clothodi-
dae. Proceedings of the California Academy of Sciences 45: 9–34.
Ross, E. S. (2001). EMBIA: contributions to the biosystematics of the insect order Embiid-
ina. Part 3: The Embiidae of the Americas (Order Embiidina). Occasional Papers of the
California Academy of Sciences 150: 1–86.
Valentine, B. D. (1986). Grooming behavior in Embioptera and Zoraptera. Ohio Journal of
Science 86: 150–2.
122
Paraneoptera (= Acercaria)
This assemblage of orders comprises barklice and true lice (Psocodea), true
bugs (Hemiptera), and thrips (Thysanoptera). The latter two orders are fur-
ther classified under the category of Condylognatha to highlight the special-
ization of their mouthparts and their sister-group relationship. The origins of
Paraneoptera go back to before the Carboniferous, placing them among the
oldest insect groups. Hemiptera are the most diverse hemimetabolan lineage
and, together with the remaining paraneopteran orders, constitute 10% of all
known insect species.
123
Psocodea
(barklice, booklice, and true lice)
Common name Barklice, booklice, and Metamorphosis Incomplete (egg, nymph,

true lice adult)
Derivation Gk. psokos–gnawed Distribution Worldwide
Size Body length 1–10 mm. Number of families 61
Mostly under 6 mm Known world species > 11,270 (0.99%)
Long thread-like
antenna
Bulging head
Ocelli
Prominent compound eye
Simple
venation
Slender leg
Soft-bodied
Graphopsocus cruciatus — Psocodea
Key features
Bark lice:
• small and generally cryptic
• clypeus distinctly bulging outwards, wings macropterous, brachypterous,
or absent
• very common on trees, vegetation, and in litter
• some species are pests of stored products
True lice:
• small, wingless, flattened ectoparasites living permanently on vertebrate
hosts
• eyes with two ommatidia, ocelli absent, legs modified for clinging to fur or
feathers
• feed on skin debris, secretions, feathers, or blood
• several species are significant vectors of human and animal diseases
An aggregation of strikingly
© Zestin Soh
coloured barklice (Psocidae:
Clematoscenea sp.) from
Singapore.
Barklice and booklice are very common, small, squat, and soft-bodied in-
sects, which, on account of their size, cryptic colouration, and habits, are often
overlooked and certainly understudied. True lice, on the other hand, are ec-
toparasitic, wingless insects found on mammal or bird hosts, and spend all
their lives in the microhabitat provided by the skin, fur, hair, or feathers. True
lice are better studied as they infest humans and are of significant veterinary
and financial importance. Psocodea can be found in a very wide range of ter-
restrial habitats including caves and the nests of birds, bees, and wasps, but
are particularly abundant in litter, soil, and on the bark and foliage of trees
and shrubs. Despite the fact that they can make up a significant proportion
of the total biomass in tree canopies and are preyed on by ants, spiders, and
birds, their ecological role and impact has not been well studied and is little
understood. Although true lice bear little resemblance to bark- and booklice
PSOCODEA 125
© Gilles San Martin

A male head louse, Pediculus
humanus capitis, on its
natural habitat – a human
scalp. Lice use their claw-like
tarsi to hold onto hairs.
© John Gausas, The Academy of Natural Sciences of Drexel University

A pair of mating Ibidoecus sp.
lice on a feather of the
Australian white ibis
(Threskiornis molucca).
and were previously classified in a distinct order known as the Phthiraptera,

overwhelming morphological and molecular evidence suggest that their sys-
tematic position lies deeply within Psocodea. Due to their radically different
morphology and life history, we describe true lice separately from the rest of
Psocodea.
The Psocodea are subdivided into three main subgroups: the Trogiomor-
pha, the Troctomorpha (which also includes true lice), and the Psocomorpha
(containing the vast majority of the known species). The relationships within
and between different psocodean families are poorly understood, and current
classification systems are likely to change in the future.
The head, which is large and appears very bulging at the front, carries a pair
of prominent compound eyes, simple chewing mouthparts, long thread-like
antennae, and, in winged psocodea, three ocelli. The antennae may have any-
thing between 10 and 50 segments and are generally half as long as the body
or longer. Most species can be classified as epiphytic grazers. The chewing
mandibles are asymmetrical, and parts of the maxillae are rod-shaped and
used to brace the head against the substrate as the mandibles scrape away at
the microflora, which may consist of algae, lichens, and moulds and fungal
spores. Some species eat pollen and other plant tissues. The genus Psilopsocus
Typical front wing venation (Psilopsocidae) contains some very unusual species, which have wood-boring
in the Psocidae. habits.
When wings are present, they are held roof-like over the body at rest and
arise from a thorax that appears slightly humped in side view. The front wings
are slightly larger than the hind wings, and both pairs have simple venation.
A few species have scales on their wings, and bodies that can make them
look superficially like very small moths. There are many species in which
short-winged and wingless forms occur in either sex.
Sexual dimorphism in wing development is typical of certain genera and
families, and species living in stable habitats that do not need to disperse as
much as those in other habitats tend to have shortened wings. The three pairs
of legs are of a normal, walking design, and although some species have ex-
panded hind femora and can jump weakly, most species are simply quick
runners. Many species are solitary scavengers, living on vegetation, bark,
and leaf litter, and under stones or anywhere that has adequate moisture.
Some species exhibit varying degrees of gregariousness; groups of adults and
nymphs living and feeding. Members of the tropical family Archipsocidae are
very gregarious, and remarkable in that the dense silken sheets produced by
their labial silk glands can be so extensive that they cover whole bushes and
trees.
Courtship in some species may involve the production of sound from spe-
cialized organs at the base of the hind legs. These structures, called Pearman’s
A typical wingless organs, take the form of a ridged area and tympanum on the inner face of
psocodean.
the coxa. In general, the males, following courtship, which may involve wing
fluttering and other vibrational signals, transfer sperm to the females in a
sperm packet known as a spermatophore. The courtship behaviour of many
Psocodea is quite unusual. In the trogiid species Lepinotus patruelis, males
and females compete for mates and will try to struggle with already copulat-
ing couples. Females do not appear to be very fussy about their mates. Males,
on the other hand, exhibit a great deal of mate choice, and, interestingly,
males on a high-quality diet are choosier in picking mates than males on a
low-quality diet. This may have something to do with the investment that the
males put into their special sclerotized spermatophore, coupled with the fact
that sperm may serve as a source of nutrients for the females. An extreme of
this condition has been found in two members of the family Prionoglarididae
(Afrotrogla and Neotrogla spp.), where the sexual roles have been completely
reversed. The males possess vagina-like genitalia, while the females have a
penis-like organ known as the gynosome. The females compete for access to
males, and mate with several individuals. Mating lasts for a remarkably long
PSOCODEA 127
period (40–70 hours), during which the males transfer to the females volumi-
nous and nutritious spermatophores. It is thought that the nutrient-poor cave
environments in which these species live have forced females to mate with as
many males as possible (up to 11 spermatophores have been found in some
females), as the spermatophores provide them with necessary food resources
that would be hard to obtain only by scavenging.
Eggs may be laid singly or in clusters, on vegetation or under bark, where
they may be exposed or covered with silk, detritus, or adult faecal material.
Females of many species are attracted to fresh conspecific eggs and may add
their own eggs to the batch or near to it. Guarding of egg batches by males
and females has been observed in some species, but in most species the eggs
are not attended. The nymphs of many species stay together in groups and
normally pass through six instars before becoming adults, although in some
species the number is reduced to five or four.
The females of a single genus (Archipsocopsis spp.) give birth to live young,
and the reproductive tract may contain more than ten embryos, which ob-
thelytoky tain nutrition from the walls of the ovarian tubules down which they pass.
parthenogenesis where only Thelytoky is common, and species in which this occurs tend to have wider
female offspring are produced distributions than bisexual species.
Many psocodean species, such as those belonging to the genera Lepinotis,
Liposcelis, and Trogium, are associated with humans and have become very
widespread. Pest species are associated with houses and stored products, such
as dried fruit, grains, and flour, and some can be pests in museum collec-
tions. Outbreaks are often due to poor house-keeping, and a combination of
warm and damp conditions. However, non-parasitic Psocodea overall are of
negligible economic importance.
Psocodea are a remarkably old group, with phylogenomic estimates sug-
gesting their first radiations started in the Carboniferous (330 mya), if not
earlier. Modern families start appearing in the Cretaceous.
One of the most peculiar branches of the Psocodea are undoubtedly the
true lice, which are among the most prevalent ectoparasites in birds and many
mammals (but not bats). It seems odd that there are no lice on bats, one of
the most diverse groups of mammals on earth today, and there is no clear ex-
planation available. A reasonable hypothesis suggests that although animals
can, of course, ‘pick up’ lice from other species, the aerial lifestyle of bats does
not bring them into intimate contact with other animals from which they
could acquire lice secondarily. Furthermore, the habitats of true lice ances-
tors may not have favoured an association with bats: true lice evolved from
barklice-like ancestors who had probably become adapted to eating the de-
tritus in nests, lairs, and burrows. Indeed, true lice are most closely related
to the barklice family Liposcelididae (suborder Troctomorpha), whose mem-
bers are still frequently associated with vertebrate nests, and some have even
been found on the fur or plumage of the animals that constructed them. A
close association of true lice ancestors with animal nests, coupled with the
loss of wings, could easily have led to a more intimate and permanent re-
lationship with the animal itself. Early mammals and birds arose from the
end of the Cretaceous and, by the end of first part of the Tertiary Period (the
Palaeocene and Eocene 65–53 mya), the forerunners of modern mammals

and most of the main groups of birds had evolved. The earliest recognizable
parasitic louse comes from an Eocene fossil (44 mya) with chewed up feath-
ers still in its foregut. Molecular divergence estimates push the origins of true
lice back to the Cretaceous (up to 140 mya) when mammals and birds started
radiating, showing a clear co-evolution between host and parasite. Older es-
timates going back to the Permian (> 200 mya) have been suggested but need
to be further re-evaluated.
Suborders of the Phthiraptera
Suborder Feeding Food Hosts

method
Amblycera Biting Skin, secretions, Birds and mammals
feathers, (blood*)
·········································································································································································
Ischnocera Biting Skin, secretions, Birds and mammals
feathers, (blood*)
·········································································································································································
Anoplura Sucking Blood Mammals, including humans
·········································································································································································
Rhynchophthirina Biting Skin, debris, etc. Warthogs and elephants
* Some species have mandibles modified to pierce the skin of their hosts.
The order has been split into four suborders within the Phthiraptera (see
Box), although their validity remains to be confirmed by molecular data. We
mention these suborders and the term Phthiraptera primarily for historical
and practical reasons, as most of the published literature prior to the 2000s
employs this classification system.
Amblyceran and ischnoceran lice use birds and mammals as hosts, while
the only hosts of Rhyncophthirina and anoplurans are placental mammals.
All lice are wingless and have tough, flexible, dorso-ventrally flattened bod-
ies (unlike fleas which are laterally flattened). Additionally, the eyes are very
small or totally absent, there are no ocelli, and the antennae, which are short
and stout, have between three and five segments. In some species, the anten-
nae are used for holding onto the host. The three pairs of legs are short and
robust with the tarsi and claws usually highly modified for grasping hair or
feathers. The surface of the body is well endowed with sensory hairs of various
kinds.
Lice eggs are typically stuck to the hair or feather with a waterproof, fast-
acting glue. The nymphs pass through three nymphal stages, taking anything
from two weeks to a few months to reach adulthood. Many lice have symbiotic
relationships with bacteria, which live in special mycetocytes associated with
the digestive system. These bacteria allow the lice to digest feather protein
(keratin) and blood. Unlike fleas (Siphonaptera), which are holometabolous
and have larvae that are not attached to the host, the entire life cycle of lice is
tied to the host animals, and they will die if separated from their host for too
long.
The mouthparts of species in the suborders Amblycera and Ischnocera are
very similar in design to those of non-parasitic Psocodea in that they have
PSOCODEA 129
short biting mandibles, and the head is braced against the substrate by a stiff,
rod-like component of the maxillae. The head is generally broader than the
thorax, and the antennae are less than half the length of the head. These lice
are mostly host specific on a variety of bird and mammal species. It is esti-
mated that all bird species and a quarter of all mammal species are used as
hosts by ischnoceran and amblyceran lice. Some species feed on the blood
of the host, but the great majority eat hair, feather, and skin fragments, the
products of sebaceous glands, and other dermatological detritus.
Species of the family Menoponidae are typical amblyceran lice. They are
confined to birds and most feed on feathers or live in the feather shafts, but
some suck blood. The head is large, broadly triangular, and expanded behind
the eyes, and the mandibles bite horizontally. The short antennae are slightly
clubbed and can be concealed in grooves on the underside of the head. The ab-
domen is broadly oval, and the legs are short and stout, each with two specially
adapted claws for holding on to their host’s feathers. Some species are quite
serious pests of poultry. The Large Hen or Chicken Body Louse (Menacanthus
stramineus) and the Shaft Louse (Menopon gallinae) are two well-known pest
species. Infestations of these poultry lice lead to feather loss and reduced host
health. Species in the genus Piagetiella actually live inside the throat pouches
of cormorants and pelicans where they suck blood. Oddly, the females leave
the pouch to lay their eggs on neck feathers. Studies of the lice on nestling
swifts have shown that higher than normal parasite loads do not seem to af-
fect their growth rate or success, confirming the prediction that a vertically
transmitted parasite (transmitted between generations) would not be success-
A typical biting louse of the ful if it adversely affected the reproductive success of its hosts. Another study
suborder Amblycera. on cliff swallows showed that fumigation of birds to rid them of a suite of
ectoparasites, including blood-sucking cimicid bugs, fleas, and chewing lice,
increased adult survivorship by 12% over non-treated birds.
Mammal-chewing lice of the family Trichodectidae are typical ischno-
cerans. The head is not triangular, but rather squarish in appearance. The
antennae are prominent and have three segments, and the mandibles bite
vertically. The legs are short and stout, and each bears a single tarsal claw. Tri-
chodectids are parasitic on Carnivora as well as rodents, hyraxes, lorises, and
some monkeys. They live in the fur of their hosts and feed on skin fragments,
hair, secretions, and blood. If populations of these small insects become large,
the hair or fur of the host can become very damaged. Some species in this fam-
ily are pests of domesticated animals. The Cattle Biting Louse (Bovicola bovis),
the Horse Biting Louse (Bovicola equi), and the Dog Louse (Trichodectes ca-
nis) are typical examples and all can cause severe irritation to their hosts, as
well as loss of hair due to prolonged scratching. The Dog Louse is known to
transmit tapeworms from dog to dog, but no other species in this family is a
known disease vector.
The mouthparts of species in the suborder Anoplura (sucking lice) are
modified to form a piercing structure composed of three stylets, which can
be retracted into a pouch in the head when not in use. In these lice, the head
is generally narrower than the thorax, and the antennae are about the same
length as the head. Anoplurans are all attached to mammals (including hu-
mans) where they suck blood, and some can be serious vectors of several
human and animal diseases. Body lice belong to the family Pediculidae and
have pear-shaped, flattened bodies. Like all lice, they are well adapted to their
hosts and have short, strong, inwardly curved legs, each armed with a large
claw for grasping and climbing through hair.
The family contains only one genus Pediculus, which comprises two
species. Pediculus humanus has two subspecies that live on humans (Pedicu-
lus humanus humanus (the Body Louse) and Pediculus humanus capitis (the
Head Louse)), which also occurs on monkeys from the New World. The other
Modifications to the leg of
the Head Louse to enable it
species, Pediculus schaeffi, is found on gibbons and apes. Human head lice and
to grip human hair. body lice are sometimes referred to by trinomens (as mentioned), or they are
treated as sibling species. They are very similar in appearance but have very
different habits. The Head Louse is confined to head hair and rarely acts as a
vector of disease. The female attaches its eggs (nits) to hair shafts with a strong
glue, and a complete cycle from egg to adult takes about 3–4 weeks.
The Head Louse is transmitted from host to host by the exchange of head
wear, combs, brushes, direct contact, and fallen hair carrying unhatched eggs.
Bites can be very irritating and can become infected or cause allergic reac-
tions. Outbreaks of human head lice are an increasingly common occurrence
in schools, especially among younger children, who play and work together
in very close contact. The use of insecticidal shampoos has inevitably led to
resistance developing in the lice, but regular washing and combing with a fine
nit or ‘dust’ comb is very effective.
The Body Louse attaches its eggs to clothing, especially along the seams,
and lives mainly in the fibres of the clothing. It leaves the clothing to feed on
blood and then returns to its hiding place. Each meal may take a few minutes
and they feed at frequent intervals. The whole life cycle from egg to adult may
take anywhere between eight days or as long as five weeks. The Body Louse is
transmitted by changing or sharing infested clothes or bedding, can survive
The eggs or ‘nits’ of the longer off the host than the Head Louse, and is capable of producing many
Head Louse are glued firmly more offspring. This species has had an immense impact on the course of hu-
to the shaft of human hair.
man history, and its effects in determining the final result of many conflicts
has been documented since the fifteenth century. Until the invention of DDT
during the Second World War, many more people died of louse-borne dis-
eases, such as epidemic typhus and relapsing fever, than were killed by their
opponents. The best-known example is the defeat of Napoleon’s army, not
by the Russians but by the loss of hundreds of thousands of men to typhus,
spread by the infestations of Human Body Louse. Only a few thousand of
more than half a million men that started the campaign, survived. Lice carry
The Pubic Louse has a diseases caused by bacteria-like organisms called rickettsiae (singular rick-
characteristic broad ettsia). Epidemic typhus is caused by Rickettsia prowazeki. These organisms
appearance.
develop inside gut cells of the louse and are passed out in the excrement. They
can then be scratched into louse bites, or can be inhaled as a component of
dust, or swallowed.
PSOCODEA 131
The Pubic Louse (Phthirus pubis) belongs to the anopluran family, the
Pthiridae, and is superbly adapted for clinging to the thick pubic hair of
humans.
The tarsal claws of the middle and hind legs are large and fold back to
grip the hair shaft firmly. Pubic lice do not move around a great deal and
are certainly incapable of the prodigious feats of athleticism alluded to in the
work of toilet-wall poets. The lice can only move between humans who are
engaged in sexual activity. The imaginative sexual behaviours of humans have
led pubic lice to occasionally infest other parts of the body, such as beards and
eyelashes. The family also contains another species, Phthirus gorillae, which,
as the name implies, is confined to gorillas.
Short antenna
Small compound eye

Single tarsal
claw
Tibia
Short,
Thoracic robust legs
segments
fused
Dorsoventrally
flattened body
Pediculus humanus
capitis—the Human Head
louse.
Other interesting anopluran lice belong to the family Echinophthiriidae,

whose member species are blood-sucking ectoparasites on sea mammals. For
instance, Lepidophthirus macrorhini is found on the hind flippers of elephant
seals, and the hosts of Arctocephalus ogmorhini are Weddell seals. Louse re-
production takes place when the hosts are breeding on land, and egg laying
is stimulated by temperature rise. Mammals like dolphins, porpoises, and

whales that are permanently at sea do not have parasitic lice.
The fourth and smallest phthirapteran suborder is the Rhyncophtherina,
which contains just two species in the family Haematomyzidae. One of these
lice, whose mouthparts, carried at the end of an extended proboscis, is specific
to elephants, the other to the African warthog.
The degree of host specificity in lice is high, and many species are restricted
to particular areas of the body, especially where the host finds it difficult
to groom. In this way, more than one louse species can inhabit the host si-
multaneously. Fahrenholtz’s Rule states that ectoparasites, such as lice, have
co-evolved with their hosts so closely that when the host speciates the lice
do as well. This has been termed phyletic tracking. If Fahrenholtz’s Rule al-
ways held true, then you would expect there to be exactly the same number of
parasites as there are hosts, and that there would be a one-to-one match be-
tween parasites and hosts. In other words, the phylogenies of the two groups
would be pretty much the same. However, when comparing actual phyloge-
nies, many discrepancies appear. There are various reasons to explain this. A
species of louse may have speciated on a single host, or a host species may
have speciated independently of its louse. Lice may have switched to an ad-
ditional, unrelated host species, either because it was geographically close or
in some way readily accessible because the hosts interacted, perhaps socially.
These phenomena have been called resource tracking. Further complications
arise when a host becomes extinct, and its louse survives on a secondary host.
The lousiness of an animal might have an effect, not only on its own sur-
vival, but also on its inclusive fitness. Hamilton and Zuk (1982) put forward
the idea that certain showy, secondary sexual characteristics might enable the
selection of mates that were less infested with parasites. In birds, high loads of
parasitic lice might lead to the birds having damaged or less bright plumage,
and they might spend more time in preening activities. A study of pigeons
did indeed show that females picked mates that had no lice in preference to
ones that had artificially high numbers of lice. Although the clean males did
not seem to groom themselves any less than the lousy males, they did spend
significantly more time displaying to females.
Louse-infested animals can remove lice by biting them through preening
or grooming. Birds may get some relief from their parasites by dust bathing,
but ‘anting’ is certainly more effective. Here, birds spread their wings out and
flap them over the nest of a formicine ant species. The workers swarm over
the bird ejecting formic acid, which is an effective louse deterrent. However,
as fast as hosts evolve parasite avoidance behaviours, it is sure that the lice
co-evolve strategies to minimize their effects.
Key reading
Barker, S. C. (1994). Phylogeny and classification, origins and evolution of host associations
of lice. International Journal of Parasitology 24: 1285–91.
Broadhead, E. and Richards, A. M. (1982). The Psocoptera of East Africa—a taxonomic and
ecological survey. Biological Journal of the Linnean Society 17: 137–216.
Brown, C. R., Brown, M. B., and Rannala, B. (1995). Ectoparasites reduce long-term survival
of their avian host. Proceedings of the Royal Society of London B 262: 313–19.
PSOCODEA 133
Clayton, D. H. (1990). Mate choice in experimentally parasitized rock doves: lousy males
lose. American Zoologist 30: 251–62.
Greenwood, S. R. (1988). Habitat stability and wing length in two species of arboreal
Psocoptera. Oikos 52: 235–8.
Grimaldi, D. and Engel, M. S. (2006). Fossil Lipsocelididae and the lice ages (Insecta:
Psocodea). Proceedings of the Royal Society B: Biological Sciences 273: 625–33.
Hamilton, W. D. and Zuk, M. (1982). Heritable true fitness and bright birds: a role for
parasites. Science 218: 384–7.
Hellenthal, R. A. and Price, R. D. (1991). Biosystematics of the chewing lice of pocket
gophers. Annual Review of Entomology 36: 185–203.
Johnson, K. P., Dietrich, C. H., Friedrich, F., et al. (2018). Phylogenomics and the evolu-
tion of hemipteroid insects. Proceedings of the National Academy of Sciences of the United
States of America 115: 12775–80.
Lyal, C. H. C. (1986). Coevolutionary relationships of lice and their hosts: a test of Fahren-
holtz’s Rule. In: Coevolution and systematics. Stone, A. R. and Hawksworth, D. L. (eds),
pp. 77–91. Systematics Association, Oxford.
Lyal, C. H. C. (1987). Co-evolution of trichodectid lice (Insecta: Phthiraptera) and their
mammalian hosts. Journal of Natural History 21: 1–28.
Loye, J. E. and Zuk, M. (eds) (1991). Bird-parasite interactions. Ornithology series 2. Oxford
University Press, Oxford.
Murray, M. D. (1976). Insect parasites of marine birds and mammals. In: Marine insects.
Cheng, L. (ed), pp. 79–96. North Holland Publishing Co., Amsterdam.
Smith, V. S., Ford, D., Johnson, K. P., Johnson, P. C. D., Yoshizawa, K., and Light, J. E. (2011).
Multiple lineages of lice pass through the K–Pg boundary. Biology Letters 7: 782–5.
Thornton, I. W. B. (1985). The geographical and ecological distribution of arboreal Pso-
coptera. Annual Review of Entomology 30: 175–96.
Yoshizawa, K., Ferreira, R. L., Yao, I., Lienhard, C., and Kamimura, Y. (2018). Independent
origins of female penis and its coevolution with male vagina in cave insects (Psocodea:
Prionoglarididae). Biology Letters 14: 20180533.
Yoshizawa, K., Ferreira, R. L., Kamimura, Y., and Lienhard, C. (2014). Female penis, male
vagina, and their correlated evolution in a cave insect. Current Biology 24: 1006–10.
Yoshizawa, K. and Lienhard, C. (2010). In search of the sister group of the true lice: a sys-
tematic review of booklice and their relatives, with an updated checklist of Liposcelididae
(Insecta: Psocodea). Arthropod Systematics and Phylogeny 68: 181–95.
Yoshizawa, K. and Johnson, K. P. (2010). How stable is the ‘Polyphyly of Lice’ hypothesis
(Insecta: Psocodea)?: a comparison of phylogenetic signal in multiple genes. Molecular
Phylogenetics and Evolution 55: 939–51.
Wappler, T., Smith, W. S., and Dalgleish, R. C. (2004). Scratching an ancient itch: an Eocene
bird louse fossil. Proceedings of the Royal Society B: Biological Sciences 271: 255–8.
Wearing-Wilde, J. (1996). Mate choice and competition in the barklouse Lepinotus patruelis
(Psocoptera: Trogiidae): the effect of diet quality and sex ratio. Journal of Insect Behaviour
9: 599–612.
134
Hemiptera
(true bugs)
Common name Bugs, aphids, hoppers, etc. Metamorphosis Incomplete (egg, nymph,
Derivation Gk. hemi–half; pteron–a adult)
wing Distribution Worldwide
Size Body length 1–120 mm. Number of families About 170
Antenna
Compound eye
Pronotum
Scutellum
Front wing
thickened basally
Slender legs
Membranous
part of front wing
Rhabdomiris striatellus — Hemiptera
HEMIPTERA 135
Key features
• mouthparts forming a piercing/sucking beak or rostrum for liquid feeding
• stink glands and sound producing organs sometimes present
• many are significant crop pests, and some transmit human and animal
diseases
Spittle bug nymphs hide in a
© Zestin Soh
mass of their liquid waste
colloquially known as
‘cuckoo spit’.
Ants frequently tend

hemipteran insects for their
honeydew. Here, an ant is
tapping a treehopper
© Zestin Soh
(Membracidae), requesting a
drop of sugary liquid.
Sri Lanka.
The nymphs of many

assassin bugs (Reduviidae)
camouflage themselves by
attaching debris and
discarded prey on their
body. In this way, they are
© Zestin Soh
both invisible to their
unsuspecting prey, as well as
larger predators. Singapore.
Bugs, which comprise about 9.5% of all insect species, are the largest and most
successful of the exopterygote orders. They range from minute, wingless scale
insects, hardly resembling insects at all to giant water bugs with raptorial front
legs capable of catching fish and frogs. Virtually every type of terrestrial and
freshwater habitat has a particular and characteristic bug fauna, and ocean
striders of the gerrid genus Halobates can be found on the sea, hundreds of
miles from land.
The order is split into four suborders: the Auchenorrhyncha (cicadas,
froghoppers, lantern bugs, leafhoppers, planthoppers, and treehoppers), the
Coleorrhyncha (moss bugs), the Heteroptera (the true land and water bugs),
and the Sternorrhyncha (jumping plant lice, whiteflies, conifer woolly aphids,
phylloxerans, aphids, scale insects, and mealy bugs). Older classifications
Pond skaters are predators

who locate their prey by
detecting the ripples they
produce on the water
surface when drowning.
© Anne Riley
Here, a group of Gerris

lacustris are feeding on a
dead Bombus pascuorum
bumble bee. UK.
HEMIPTERA 137
© Anne Riley
Female aphid bearing live
young. Megoura viciae, UK.
lumped the Auchenorrhyncha, Coleorrhyncha, and Sternorrhyncha into a

single suborder known as the Homoptera, which has now been shown by
molecular and morphological studies to be paraphyletic and is therefore ob-
solete. Two pairs of wings are usually present and in heteropterans, the front
part of the front wings is toughened, leaving a membranous region at the rear.
In Auchenorrhyncha, the front wings and hind wings may be membra-
nous, or the front wings may be entirely toughened.
The earliest bugs were probably plant feeders and lived on land, sim-
ilar to many species today. All coleorrhynchans, auchenorrhynchans, and
sternorrhynchans are herbivorous, feeding on the sap or cell contents of
vascular plants. While the Heteroptera contains many predacious species,
mixed feeders, and a few specialist blood feeders, the majority of species are
A typical heteropteran bug
herbivorous.
(wings held flat over body). Bugs are sometimes called the ‘perfect suckers’ because the single unique
character linking them all is the possession of piercing, sucking mouthparts
in the form of a long, typically four-segmented beak (known as the labium
or rostrum in entomological terminology). In heteropteran bugs, the labium
arises from the front part of the head and can be hinged forward to point
down or forwards, in front of the head.
Suborders of Hemiptera
Suborder Number of Feeding habits

families
Coleorrhyncha 1 (Peloridiidae) Herbivorous on mosses and liverworts
·········································································································································································
Heteroptera About 88 Herbivorous, predacious, parasitic
·········································································································································································
A typical auchenorrhynchan Auchenorrhyncha 33 Herbivorous
bug (wings held roof-like ·········································································································································································
over body). Sternorrhyncha About 49 Herbivorous
(i) (iii)
Front leg
Labium
Labium
(ii)
(i) Heteropteran bug: labium arises from
front of head.
(ii) Auchenorrhynchan bug: labium arises
from rear of head.
(iii) Sternorrhynchan bug: labium arises
Position of labium in the
three main bug suborders. Rostrum nearly between front legs.
This allows much greater flexibility and a larger choice of food. In the Auchen-
orrhyncha and Sternorrhyncha, the labium, which arises from the posterior
part of the head, or seemingly from between the front legs, is permanently
directed backwards.
With the exception of non-feeding male scale insects and the sexual forms
of a few aphids, whose mouthparts are vestigial or lacking, the bug labium
is similar throughout the order. The outer covering of the labium, (with 1–4
segments), is grooved for most of its length and surrounds the slender, tough-
ened feeding stylets. The stylet bundle is made up of a pair of mandibular and
a pair of maxillary stylets. The mandibular stylets enclose the maxillary stylets
and can be closely connected by means of longitudinal ridges and grooves on
their surfaces fitting together like the seal of a zip-lock plastic bag. The two
pairs of stylets can slide freely on each other but are difficult to pull apart. The
mandibular stylets have saw-like serrations, teeth, and, sometimes, barbs to
penetrate plant and animal tissues. Predacious bugs penetrate the cuticle of
their prey through a weak spot and use their long stylets and saliva to mac-
erate the internal tissues before they are sucked out. The inner surfaces of
the maxillary stylets are folded into longitudinal ridges and grooves, which
firmly unite the two and provide two very fine, parallel canals running along
their entire length. The ventral canal is the salivary canal, which carries diges-
tive enzymes from the salivary glands in the anterior part of the thorax, the
other is the food canal. Bug saliva is a complex mixture of a number of differ-
ent enzymes, toxins, lubricants, and other substances. Herbivorous bugs need
enzymes, such as pectinases, to break down plant cell walls, while the saliva
HEMIPTERA 139
of carnivorous heteropteran bugs may contain powerful venoms causing the

instantaneous paralysis, death, and internal liquefication of prey.
Maxilla-interlocking
(ii) ridge and groove
Head Muscles of
(i) cibarial pump
Food canal
Salivary pump Left mandible Right mandible
Oesophagus Left maxilla Right maxilla
Salivary canal
Salivary duct Labrum
X
Stylet bundle
a b c d e
Labium (iii)
Tip of stylet bundle

Left Right Maxillae
mandible mandible pushed
advances advances between
mandibles
(i) Diagrammatic section
through a bug’s head.
(ii) Cross-section through In plant sap-feeding bugs, such as aphids, the site of the phloem vessels
stylet bundle at X. in the plant may be some distance from the surface, and the stylet bundle
(iii) Stages in the insertion of
has to wander between the tough-walled cells of the plant’s epidermis be-
a bug’s stylet bundle into food
source. The process may be fore reaching a feeding site. The stylets are protected by the formation of a
repeated many times until the proteinaceous sheath formed by the hardening of special salivary gland se-
required location is reached.
cretions, which are produced throughout the course of penetration. Inside
the head, powerful muscles operating the sucking or cibarial pump draw the
liquid or pre-dissolved food up the stylet bundle and pass it into the pharynx.
Bugs feeding on the sap in phloem vessels do not require very strong cibarial
pumps as their diet is under slight positive pressure.
As plant material is not generally very nutritious, having at most 4% of its
dry weight as nitrogen, there are two options open to herbivorous bugs. They
can consume large volumes or grow slowly. The sap of phloem vessels, fed
on by the great majority of herbivorous bugs, usually contains much less than
0.5% weight to volume of soluble nitrogen, but up to 9–10% weight to volume
of sugars. Xylem sap, fed on by cicadas, froghoppers, and some leafhopper
species, is even poorer in nutrients than phloem sap, and some bugs that spe-
cialize on this diet may take anything from several months to many years to
complete their nymphal development.
A modification of bugs’ guts seen in many sap suckers, such as the Cica-
didae, Cercopidae, some Cicadellidae, and virtually all the coccoid families,
is the filter chamber, formed by the intimate association of epithelial tissues
from the rear of the midgut or the front of the hindgut with those of the an-
terior foregut. This allows large amounts of water and low molecular weight
molecules to bypass the midgut entirely and pass straight into the rectum,
while the rest of the food carries on through the midgut to be processed more
efficiently.
Fore gut
Malpighian tubule
Anterior mid gut
Posterior mid gut
Hind gut
Posterior mid gut
Schematic representation of
the digestive tract and filter
chamber of an
auchenorrhynchan bug.
Most of the water in its diet
passes directly from the Rectum
interior midgut into the
posterior midgut and
hindgut via the filter
chamber.
Some heteropteran bugs rely on symbiotic microorganisms to provide them

with essential nutrients, and these may be contained within special bod-
ies in the haemocoel or inside the lumen of the midgut. In blood-feeding
species, bacteria are responsible for providing the bugs with B vitamins. How-
ever, auchenorrhynchan, coleorrhynchan, and sternorrhynchan bugs that
feed generally on a diet low in nutrients, are especially well known for their
bacterial (Proteobacteria) symbionts, which are contained inside special cells
called mycetocytes. The mycetocytes, which may be aggregated into bodies
known as mycetomes, are free in the haemolymph of the body cavity or as-
sociated with the fat body. The symbionts are important in the recycling of
nitrogen, providing their hosts with vitamins, certain essential amino acids
such as tryptophan, which are absent from the diet, and certain lipids, such as
the sterols, which insects are unable to manufacture themselves and are vital
to the production of ecdysone (the insect moulting hormone). The symbionts
of a few bug groups have been examined. Aphid symbionts belong to the
Buchnera aphidicola complex and these differ from those seen in whiteflies or
scale insects. Others, such as whiteflies and coleorrhynchans, share the same
type of endosymbionts, indicating that their association is very ancient. The
success and radiation of sap-feeding auchenorrhynchan and sternorrhynchan
bugs was made possible by the acquisition of bacterial symbionts. A unique
case of insect–plant coevolution can be found in the whitefly Bemisia tabaci,
a major agricultural pest. This species acquired a detoxification gene from its
host plant through horizontal gene transfer (non-sexual movement of DNA
between two different species) millions of years ago. This ‘stolen’ plant gene
allows B. tabaci to neutralize the chemical compounds produced by plants to
ward off pests, and to feed on a variety of species that other insects cannot.
HEMIPTERA 141
Predatory bugs can either be active hunters, like many assassin bugs that
stalk and pounce on their prey, or ambushers, that wait for their prey. Timid
predators, like anthocorids and water bugs such as hydrometrids, do not use
their legs to hold prey, preferring to stab very small or slow-moving insects,
larvae, eggs, and pupae with their rostrum. Predatory stink bugs (Pentatomi-
dae) have been shown to locate their caterpillar prey by responding to specific
chemical odours as well as the vibrations produced when the caterpillars are
chewing on leaves. Some intriguing examples of prey specialization can be
seen in several bugs associated with webs. The reduviid subfamily Emesinae
contains a number of elongate, thread-legged species that feed on insects
caught in spider webs or on the spiders themselves. Stenolemus species have
specially modified antennae, which are used to send and receive vibrational
signals through the web. When the bug moves on to a web, it causes the web
to sway in a manner to simulate wind motion, therefore being acoustically
invisible to its spider prey. The bug also actively manipulates the web by cut-
ting silk threads in its path, therefore covering its tracks. Other interesting
prey-capturing techniques are found in assassin bugs that may have special
hairs on various parts of their bodies, which are designed to secrete sticky
substances. In the nymphs of many genera, the dorsal body surface hairs are
used to hold debris or the sucked-out bodies of their prey as a camouflage.
Some adult assassin bugs produce sticky secretions on their front legs and use
them to catch small insects. Others, particularly those belonging to the sub-
family Apiomerinae, go one step further and collect certain plant resins on
brush-like organs on their front tibia, and use this to trap stingless bees and
beetles. Even more interesting is the use of tools by some assassin species in
the subfamily Salyavatinae that prey on termites. Nymphs of the neotropical
species Salyavata variegata disguise themselves heavily by sticking debris to
glandular hairs on their backs and go fishing for their prey using the sucked-
out bodies of previous victims as bait. The nymphs dangle the bait in front
of holes in the termite nest, and the workers that come out to investigate and
often grasp the lure are seized and eaten by the assassin bug.
(i) Hind leg Opening of

Front leg attaches here scent gland (ostiole)
attaches here
Prosternum
(i) View of the underside of Mesosternum
an adult heteropteran bug’s
Middle leg
thorax, showing location of
attaches here
metathoracic scent gland
openings. Metasternum Peritreme
(ii) Magnification of the
surface sculpturing of the
‘evaporative’ or scent
retentive area around the
gland opening.
(iii) Heteropteran nymph (ii)
with three dorsal abdominal (iii)
gland openings (Lygaeidae).
A characteristic defence system of heteropterans is the production of mix-

tures of noxious or repellent organic compounds from scent glands in the
metathorax of adults and the abdomen of nymphs. The secretions may also
contain components which function as aphrodisiacs and sexual attractants, or
promote aggregation, dispersal, and alarm. The secretions produced by adult
bugs are generally more complex and varied than those produced by nymphs,
and fall into several major chemical categories, such as the aliphatic aldehy-
des (alkanals), aliphatic alcohols (alkanols), and dicarbonyl compounds. The
compounds can be active as vapours, soluble liquids, sprayed into the eyes or
mouth of an attacker, or wiped by the bug on to the surface of the attacker,
where they may penetrate beneath the cuticle or skin. Some scents panic
ants by imitating their own alarm pheromones. Of course, evolution can turn
the tables on the best laid plans, and certain parasitic flies (Tachinidae) and
a number of hymenopteran parasitoids use the defensive scents and sexual
odours of nymphal and adult bugs as cues to locate and thus parasitize them
or their eggs.
Many seed-feeding lygaeid bugs, such as the red-and-black-coloured
Milkweed bug (Oncopeltus fasciatus), are able to store toxic cardenolides ob-
tained from their host plants in special compartments of their body, which
they release rapidly when attacked.
Many bugs have evolved mutualistic relationships with ants. Among the
heteropterans, ground-living nymphs and sometimes adults of a few fami-
lies, mimic ants and the similarity enables them to prey on their host’s larvae
and pupae. Plant-feeding heteropterans, such as plataspids (Plataspidae) and
leaf-footed bugs (Coreidae), are known to provide food for ants in return for
protection from predators and parasites. However, it is among the auchen-
orrhynchan and sternorrhynchan families that we find some of the most
complicated ant relationships. Many of these interactions, known as tropho-
bioses, are based on the simple principle of reward (usually honeydew) for
services rendered (usually protection). When sap-sucking bugs feed, they ex-
tract the dilute nitrogen-containing nutrients they require, and their watery
wastes are high in sugars. This therefore represents a very valuable food re-
source to ants, and even to humans. The manna that sustained the Israelites
on their mythical flight from Egypt is very likely to have been the honeydew
of scale insects feeding gregariously on Tamarisk bushes. The ants, in return
for honeydew, act as bodyguards, and when protecting their particular colony
of aphids or scale insects, they will attack any predator or parasitic wasp that
comes near. Some ants even treat their colonies of bugs like domesticated an-
imals, building carton nests over them or taking them into their own nests
to protect them from frosts. Some mealy bug species (Pseudococcidae) will
climb voluntarily onto the backs of their bodyguards or are carried to safety
in the ants’ jaws.
For a long time, the only insects thought to show sociality in varying de-
grees were social wasps and bees (Hymenoptera) and termites (Isoptera). The
discovery, some 20 years ago, of a special non-reproducing soldier caste in
species belonging to the aphid subfamily Eriosomatinae was of great interest,
for it showed that eusociality also occurred in the Hemiptera. For example,
HEMIPTERA 143
first instar nymphs of the Sugar Cane Woolly aphid (Ceratovacuna lanigera)
serve as colony defenders by using sharply pointed frontal horns to attack
predators like hover fly larvae. In addition, the soldiers secrete droplets from
their abdominal cornicles, which are spread on the attacker’s body. The se-
cretion contains an alarm pheromone, which causes all other aphids in the
colony to back away and recruits many more of the sterile soldier nymphs.
Several gall-inhabiting aphid species have also been shown to have a first in-
star soldier caste. In these species, the soldiers may have distinctly thickened
or lengthened legs, which are used to kick and pierce the bodies of predators,
such as anthocorid bugs, ladybirds, and hover fly larvae, as well as herbivo-
rous competitors such as moth caterpillars. The fighting techniques vary from
species to species: some use their hind legs to tear cuticle, others have sharp
horns, and many use their short, stout rostrum to stab. Although soldiers may
spend much longer in the first instar and are able to feed, in the vast majority
of species that have them, they do not ever live past the first instar and are
thus effectively sterile. In the aphid Pemphigus spirothecae, spirally shaped
galls are formed on the leaf petioles of the host plant, Black Poplar (Popu-
lus nigra). Unusually, the soldiers that survive their first instar develop into
wingless adults and give rise parthenogenetically to more soldier first instar
nymphs, as well as normal first instar nymphs that will ultimately become the
winged generation with the job of producing the sexual forms.
Most of the soldier aphids that attack enemies die in the process. Other
aphid species can protect themselves, and many species are able to produce
secretions from a pair of abdominal cornicles. Droplets are exuded when
the aphids are attacked, and these can spread onto the predator’s body and
mouthparts, where it hardens into a thick glue. Some aphids are polymor-
phic, the red and green forms of the pea aphid (Acyrthosiphon pisum) being
an example. Recently, it has been shown that red morphs are less vulnerable
to parasitic wasps, but more likely to be eaten by predators, and green morphs
were safer from predators than from parasitoids.
Cornicle Many sternorrhynchan bugs protect themselves by secreting waxes or
resins. The well-known Woolly Apple Aphid (Eriosoma lanigerum) is easily
recognizable by the tangled masses of white wax that it secretes. Scale in-
sects in several families yield commercial quantities of waxes and resins in
a very pure state. The wax is produced from special wax glands located in the
epidermis. The products of these glands are carried to the outside through
ducts to variously shaped pores. Scale insect wax covering may take the form
of short, hollow or solid curls with a circular cross-section. Others may pro-
duce shapeless masses, and some produce very delicate solid threads less than
Wingless female aphid 0.1 µm in diameter, which can be formed into hollow cylinders themselves.
giving birth to a first instar
Macaroni-like or even more complicated extrusions with a diameter of up to
nymph.
7 µm are also produced. In the Lac insect (Kerria lacca) nearly 80% of the se-
cretions are resins, whereas in some species of ground pearl (Margarodidae),
the secretions are essentially pure wax.
Most bugs reproduce bisexually, but a few species of scale insect, such as
the Cottony Cushion Scale (Icerya purchasi), are hermaphrodites. Due to their
profound impact on commercially important plants, and their phenomenal
powers of reproduction and dispersal, aphids have been the subject of consid-
erable study over the years. Life cycles and reproduction are usually relatively
simple, but around 10% of aphid species have complex life cycles involving
sexual and asexual forms and alternations of generations between two un-
related host plants (one woody and one herbaceous) at different times of the
year. In sexual forms, wings and mouthparts may be entirely lacking. In a sim-
ple, single host plant case, overwintering eggs hatch in the spring and give
rise to winged or wingless, parthenogenetically reproducing females. These
females, known as fundatrices, produce nymphs that develop into partheno-
genetic adult females called virginoparae. Feeding continues and a variable
number of generations exhibiting asexual reproduction are produced until
the beginning of the autumn, when the nymphs will develop into wingless
reproductive females and males. If the host plant has developed a very high
aphid population, many will have already flown off to seek out other, less-
crowded host plants of the correct species. Sexual reproduction and mating
take place and the resultant eggs are laid on the host plant.
Mate finding in bugs can simply involve males locating females at ovipo-
sition or feeding sites, or it may require sound or vibrational signalling or
pheromones. Often difficult to see due to their cryptic colouration and be-
haviour, cicadas (Cicadidae) are well known for their ability to produce very
loud acoustic songs. The songs, which are produced by males, originate from a
pair of organs called tymbals located on either side of the abdomen. The tym-
bal organs are resilin-containing cuticular membranes stiffened by a number
of sclerotized ribs. As the tymbal muscle, attached to the inside of each tym-
bal, contracts, the tymbal is deformed inwards and each of the ribs buckle
producing a loud click. The resonating tymbal organs drive an abdominal
air sac resonator, and the resultant sound is radiated to the outside via the
tympanic membranes. The tympanic membranes, or ‘hearing organs’, which
are present in males and females, are on the underside of the abdomen, pro-
tected by a cuticular flap called the operculum. Another fascinating aspect
of the biology of some cicadas that has intrigued entomologists for more
than 300 years is the synchronous mass emergence of periodical cicadas in
the genus Magicicada. Like all cicadas, the females lay eggs on the twigs of
trees and hatching nymphs drop to the ground where they enter the soil. The
nymphs spend the next 13 or 17 years feeding on the very poor diet of xylem
sap. The three species of periodical cicada in North America are different
from each other in terms of morphology and behaviour, and all have both
13- or 17-year life cycle forms, the longer life cycle forms occurring more in
northerly regions. Four-year accelerations can occur, changing a 17-year life
cycle form to a 13-year life cycle form. When the adults emerge en masse,
often within a matter of a few days of each other, the densities reached can
be as high as 3.5 million insects per hectare. The mechanisms by which the
cycle is timed, and emergence is triggered, are unknown but might involve
annual cycles of food quality, temperature, or a molecular clock. The reasons
why these cicadas take so long to develop remain unknown. One hypothe-
sis suggests that the mass emergence of billions of cicadas may overwhelm
predators, which leads to less overall mortality for adult cicadas, and also
HEMIPTERA 145
prevents their enemies from becoming accustomed to hunting cicadas every

year. Another hypothesis states that such long life cycles evolved to prevent
the hybridization with other species of cicadas.
(i) Tymbal cover
Operculum covering tympanal membrane
(ii) Tymbal
Tymbal cover muscle
(i) Location of the tymbal Tymbal relaxed Tymbal
(sound-producing) and
pulled in
tympanal (hearing) organs
on one side of a cicada.
(ii) Schematic cross section
through cicada at base of Tympanum
abdomen. Large air space
Most auchenorrhynchans, coleorrhynchans, and heteropterans produce

substrate-borne vibrational songs that cannot be heard by the human ear, us-
ing various abdominal organs, some of which being smaller, ‘silent’ versions
of the tymbals of cicadas. The vibrational energy of these small bugs is trans-
mitted as mechanical vibrations through their legs to the foliage, while the
acoustic signals of cicadas are primarily transmitted through the air. Water
bugs, such as gerrids, catch prey by detecting the ripples they make and also
attract mates by sending out pulsed surface ripples with their legs.
When copulation occurs, males introduce sperm by means of an aedeagus
or penis via the female’s vagina. Copulation usually begins with the male bug
on top of the female and holding her back with his front legs; however, the
final positions taken by mating bugs can be quite different. Once the genitalia
are locked in place, sperm transfer can begin. To avoid other males displacing
sperm with their own, males may remain in copula for a long time, or may
guard their mate until just before or even after she lays the eggs. In many bugs,
A final instar cicada nymph the female (or less commonly the male) parent may invest quite a great deal of
crawls up a stem where it energy in protecting their eggs and nymphs from attack. The occurrence of fe-
will moult into the adult
form. male brood guarding is documented in several families, but it is characteristic
of treehoppers (Membracidae) and certain stink bugs (Acanthosomatidae).
Females guard eggs or young nymphs with vigorous wing fanning, buzzing,
rapid movements, physical contact, and butting. In some species of shield bug
(Acanthosomatidae), female guarding, while effective in deterring predators,
does not seem to reduce the levels of attacks from parasitoids. If a female bug
protects her offspring, she can lay fewer eggs, but the survivorship of those
eggs is much greater.
While it is usually the job of the female to protect the brood, males of some
assassin bugs take over. In Rhynocoris (Reduviidae), when the female has fin-
ished laying, the male takes his position astride the egg mass. Over the course
of the next few days, the female will mate several times with the guarding male
and add to the original egg mass by laying additional egg batches. During the
time that the male is guarding, he will chase off parasitic wasps and other en-
emies, but may not always be completely successful in defending his progeny
for, when guarding a large egg mass, some of the eggs on the outside of the
mass may be left exposed and become parasitized. When the eggs have all
hatched, the male loses interest and moves away. The most famous cases of
sexual role-reversal in bugs are to be found in certain giant water bugs (Be-
lostomatidae). The females of many species, such as those belonging to the
genera Belostoma and Abedus take a major part in courting and competing
for available males, onto whose folded front wings they glue their eggs. The
cost to the male of this brooding service may be quite high for, while carrying
a heavy egg mass, he is less mobile, cannot fly, and is less able to catch prey.
It is therefore important that the male makes sure that the eggs he is aerating
and protecting from going mouldy are his. There would be no point in putting
himself at risk for another male’s genes. Due to the widespread phenomenon
of sperm precedence, the sperm from the last mating is generally going to
fertilize a much larger proportion of the eggs laid by the female. Males will
only allow the females to glue eggs on their backs once they have mated with
them and further enhance their assurance of paternity by letting the female
glue only a few eggs at a time before copulating again. When all the eggs are
laid the female leaves the male to look after their brood. There is recent evi-
dence of this kind of egg brooding behaviour to protect against predators and
parasites in one species of terrestrial bug, Phyllomorpha laciniata (Coreidae).
Both males and females carry eggs on their backs, but males carry twice as
many as females.
Two notorious human blood

feeders. Approximate body
lengths are shown beside
each bug. Triatoma infestans Cimex lectularius
Bed bugs (Cimicidae) have had a long history of association with people, and
probably became intimately dependent on human blood when early humans
settled down to live in caves with fires for warmth during cold spells. Bed bugs
HEMIPTERA 147
are a very old group, being dated to about 100 mya, meaning that they prob-
ably were parasitic on both dinosaurs (including birds) and early mammals.
The most common bed bug feeding on humans, Cimex lectularius, initially
parasitized bats and switched to hosts of the genus Homo sometime between
99,000–867,000 years ago. Adult bed bugs are flattened, reddish brown, wing-
less, nocturnally active insects with a body length of 4–6 mm. During the day
they hide in bedding, mattresses, or cracks in floors and walls. There are five
nymphal stages, and the entire life cycle can take anything from 2–10 months.
Hosts are located at close range by body temperature, odours, and carbon
dioxide. The adults can live for several months, and sometimes longer than
a year without a blood meal. When they feed, adult bed bugs can consume
more than five times their own weight in blood in a matter of 15 minutes,
after which they hide away.
Bugs are not generally thought of as disease carriers, but there are excep-
tions. Chagas’ disease, prevalent in Central and South America, is caused by
the microscopic protozoan organism, Trypanosoma cruzi, that is carried by
certain kissing bugs (Reduviidae: Triatominae). The disease organism and the
bugs that carry it have been present in Central and South America for a long
time and, before the arrival of people, infection was confined to a variety of
other mammalian species, birds, and a marsupial. At the beginning of the
twenty-first century, Brazilian doctor Carlos Chagas first recognized and de-
scribed the disease in humans and investigated the life cycle and other aspects
of the protozoan and its insect vectors. Certain estimates indicate that up to
8 million humans may be affected by this disease, and that many more are
at risk. Species of Triatoma, Panstrongylus, and Rhodnius have become inti-
mately associated with humans, their houses, and their domestic animals, and
the three most important in human disease transmission are Triatoma infes-
tans, Triatoma dimidiata, and Rhodnius prolixus. The disease is much more
common in rural areas where houses may harbour up to 1,000 bugs. During
the day the bugs hide in thatch or in cracks and crevices in walls and emerge
to feed at night, biting sleeping people, especially around the face and mouth,
hence the common name of kissing bugs. An adult triatomine bug can ingest
anything up to ten times its own weight in a single blood meal. The risk of
infection from a single bite is low, but as people may receive tens of bites a
night and thousands in a year, the risk of infection by Trypanosoma cruzi are
considerable. When they feed, the bugs leave excrement behind on the skin
and transmission of the protozoan occurs when people scratch their bites. As
many as 28,000 people may contract Chagas’ disease annually, most of these
being children (congenital transmission), who may die in the early critical
phase. Those that survive are infected for life, and the parasite, circulating in
the bloodstream, infects, multiplies, and ultimately destroys muscle and nerve
cells all around the body, particularly in the heart and digestive system. The
medical problems associated with the parasite are many and varied, ranging
from general weakness to serious problems of the alimentary tract, cardiac ar-
rhythmia, heart failure, and death. It has been suggested that Charles Darwin
contracted Chagas’ disease while working in the New World.
While some heteropteran bugs, such as the Southern Green Stink Bug
(Pentatomidae: Nezara viridula), can be widespread and polyphagous plants
pests, it is auchenorrhynchan and sternorrhynchan bugs that are responsible
for the biggest crop losses worldwide. Their feeding causes physical dam-
age, metabolic disorders, and directly infects or makes the plants prone to
a plethora of viral, rickettsial, mycoplasmal, and fungal diseases. Pest white-
fly species (Aleyrodidae) attack citrus fruit trees and crops grown under glass.
Scale insects and mealy bugs (superfamily Coccoidea) attack a wide range of
economically important plant species. The viral diseases that are carried and
transmitted to plants by sucking bugs probably do much more damage than
the removal of nutrients. Aphid pests, for instance, are capable of transmitting
a number of different viruses to their host plants simultaneously. The incred-
ibly polyphagous species Myzus persicae have been shown to act as a vector
for more than 110 different plant viruses. Aphids, commonly called blackfly
or greenfly, are incredibly abundant and in one acre (0.4 hectare) there may
be 2,000 million individuals on green parts of plants, and a further 260 mil-
lion underground on roots. Aphids evolved 280–300 mya, but really became
a significant group with the rise of the flowering plants in the Cretaceous (60–
100 mya). Aphids are small and do not fly very fast (1–3 km per hour), and,
as wind speeds are typically more than 3 km per hour, they have little con-
trol over where they go. They can, however, travel hundreds of miles. During
the day, as the ground heats up, convection currents are set up which carry
huge numbers of aphids up into the air. They have been recorded at 600 m
(2,000 feet) or more. In the evening, as the ground cools, the air above the
ground gets colder than the air higher above it. Convection ceases and the
aphids come down wherever they happen to be. Billions may perish at sea or
never find a host plant, but those pest species that find themselves settling on
a suitable crop can build up to damaging numbers very quickly. The number
of aphids flying each day is determined by factors such as how many are be-
coming adult, how overcrowded they are, the quality of the food plant, and
climatic conditions like temperature and light intensity.
More than 21,000 world species of leafhopper (Cicadellidae) have been
named and many are serious crop pests. Leafhopper saliva is toxic to the
plants and phloem-feeding species are the next most notorious vectors of
plant viral diseases to aphids. Many major crop plants, such as cotton, pota-
toes, beans, rice, and alfalfa, are at risk and resistant plant strains are being
developed. Some hemipterans are consumed by humans, and in fact the stink
glands of many heteropterans are prized food items. The ancient Greeks had
a taste for egg-filled female cicadas, while giant water bugs (Belostomatidae)
are regularly eaten in Southeast Asia, and their extract is often used as season-
ing for other dishes. Some stink bugs (e.g., Edessa mexicana) are a delicacy in
certain areas of Mexico and are said to taste like cinnamon.
Hemiptera are a remarkably old group, with origins in the Carbonifer-
ous (about 360 mya), perhaps even earlier. Most modern families started
diversifying in the Jurassic–Cretaceous. Extinct Hemiptera, like their mod-
ern relatives, were among the most abundant insect groups in virtually every
habitat, and their fossils are remarkably well represented.
HEMIPTERA 149
Key reading
Akhoundi, M., Sereno, D., Durand, R., Mirzaei, A., Bruel, C., Delaunay, P., Marty, P., and Izri,
A. (2020). Bed bugs (Hemiptera, Cimicidae): overview of classification, evolution and
dispersion. International Journal of Environmental Research and Public Health 17(12):
4576.
Andersen, N. M. (1982). The semiaquatic bugs (Hemiptera, Gerromorpha). Entomonograph
series 3. Scandinavian Science Press, Denmark.
Bennet-Clark, H. C. (1997). Tymbal mechanics and the control of song frequency in the
cicada Cyclochila australasiae. Journal of Experimental Biology 200: 1681–94.
Blackman, R. L. and Eastop, V. F. (2000). Aphids on the world’s crops: an identification and
information guide (2nd edn). John Wiley, Chichester.
Bramer, C., Friedrich, F., and Dobler, S. (2016). Defence by plant toxins in milkweed bugs
(Heteroptera: Lygaeinae) through the evolution of a sophisticated storage compartment.
Systematic Entomology 42(1): 15–30.
Buckley, R. C. (1987). Interactions involving plants, Homoptera and ants. Annual Review of
Ecology and Systematics 18: 111–35.
Cobben, R. H. (1968). Evolutionary trends in Heteroptera. Part 1: Eggs, architecture of the
shell, gross embryology and eclosion. Centre for Agricultural Publishing and Documen-
tation, Wageningen.
Cobben, R. H. (1978). Evolutionary trends in Heteroptera. Part 2: Mouthpart structures and
feeding strategies. V. Veenman and Zonen bv, Wageningen.
Davranoglou, L.-R., Cicirello, A., Taylor, G. K., and Mortimer, B. (2019). Planthopper bugs
use a fast, cyclic elastic recoil mechanism for effective vibrational communication at small
body size. PLoS Biology 17(3): e3000155.
Dixon, A. F. G. (1997). Aphid ecology: an optimization approach. Chapman and Hall,
London.
Foster, W. A. (1990). Experimental evidence for effective and altruistic colony defence
against natural predators by soldiers of the gall forming aphid Pemphigus spirothecae
(Aphididae: Pemphigidae). Behavioral Ecology and Sociobiology 27: 421–30.
Itô, Y. (1989). The evolutionary biology of sterile soldiers in aphids. Trends in Ecology and
Kaitala, A. (1997). Oviposition on the backs of conspecifics: an unusual reproductive tactic
in a coreid bug. Oikos 77: 381–9.
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151
Thysanoptera
(thrips)
Common name Thrips Metamorphosis Incomplete but with

Derivation Gk. thysanos–fringe, tassel; resting pupa-like stages
pteron–a wing Distribution Worldwide
Size Body length 0.5–12 mm. Number of families 9
Mostly under 3 mm Known world species 6,200 (0.55%)
Antenna
Ocelli
Compound eye
Pronotum
Small
and slender- Short legs
bodied
Narrow, hair-fringed
wings
Frankliniella occidentalis — Thysanoptera
Key features
• small to very small, slender
• narrow wings with dense fringes of hairs
• mouth with only a single mandible (left one)
• arolia modified into protrusible vesicles
• many species are serious crop pests
Thrips are very small insects
© Manfred Ulitzka
with oar-like hairy wings,
and asymmetrical
mouthparts. Megalurothrips
sjostedti can be a serious pest
of cowpeas. Saudi Arabia.
Thrips are generally small or very small, slender-bodied, readily recognizable

insects. The two pairs of wings are very narrow and, in addition to bearing
long fringes of hairs on their margins (known as ‘cilia’), they have little or no
obvious venation.
Thrip wings can be of normal size, reduced, vestigial, or totally lack-
ing. The head carries a pair of short antennae, each with between four and
nine segments, and conspicuous compound eyes, and three ocelli in winged
individuals.
The piercing and sucking mouthparts of thrips are unusual in that they are
asymmetrical. The right mandible is extremely reduced and non-functional,
while the left mandible is sharp and stylet-like, and it is used to penetrate plant
A typical adult
thysanopteran. tissue or the bodies of minute insects. The remaining mouthparts form long
hemiptera-like stylets and are used to suck up the liquid foodstuff or, in some
cases, fungal spores. The three pairs of legs are characterized by the presence
of specialized devices that are used for attachment to plant and other surfaces:
the tarsi of all species have a large adhesive, bladder-like protrusible vesicle
contained in a hollow between the claws. When the tarsus contacts a surface,
the bladder is pumped up with haemolymph. The front pair of legs of most
thrips are usually greatly enlarged, making thrips look like the Popeye of the
insect world.
Although these insects have a hemimetabolous mode of development and
are most closely related to the Hemiptera, they are unusual in that there are
one or more pupa-like resting stages between the two, true nymphal stages
THYSANOPTERA 153
and the adult. In some cases, there are three pre-adult stages, of which the first
may still be capable of feeding. The next two pre-adult stages become more
antenna pupa-like with a degree of tissue reorganization, and a cocoon may even be
spun.
Like the Hymenoptera, thrips are haplodiploid. That is, female thrips,
compound
which result from fertilized eggs, are diploid, however, males are haploid. This
eye
means that unfertilized virgin females can give birth to males, in a process
known as arrhenotoky. However, there are also some thelytokous species,
ocellus where females are produced from unfertilized eggs.
Detail of the head of a thrip. Although Thysanoptera is a fairly small order, their life cycle can be quite
complex. A few species show sub-social behaviour and parental care. In the
arrhenotoky
Australian species, Oncothrips tepperi and Oncothrips habrus (Phlaeothripi-
parthenogenesis where
haploid males are produced dae), sub-fertile soldier morphs defend their colony and siblings from attack
from unfertilized eggs by other insects, as well as take-overs by other thrips species.
Even in small insects like thrips, sex and violence can be inextricably
linked. In some species of Hoplothrips (colonial fungus-feeding thrips), males
fight, often to the death, using enlarged front legs to grapple and stab each
other in order to control access to prime egg-laying sites and therefore ac-
cess to females. Victorious males, which tend to be larger than vanquished
males, get to mate with the females just before they lay eggs, and are able
to assess the reproductive potential of their mates by ‘measuring’ the females’
abdomen with their front legs. Fights between males usually result from chal-
lenging males sneaking in to mate with females. Fighting is clearly worth the
risk because defending males may be defeated through accumulated injuries
from previous fights, and the rewards of becoming a defender are high in
terms of inclusive fitness. Development from egg to adult stage may be very
rapid, taking less than a fortnight, and there may be more than one genera-
tion during a single year. Herbivorous and pollen-feeding species can be very
destructive and are responsible for the transmission of many plant diseases,
while predacious species may be of benefit in controlling pests. Some phy-
tophagous species are of use in the control of weeds. Around 300 species in
more than 50 genera induce galls on their host plants. In these species, galls
are formed by the action of feeding by the thrips on growing tissue, but the
causal mechanism is not known.
There are two suborders: the Terebrantia, which includes large families
such as the Aeolothripidae and the Thripidae, and the Tubulifera, which con-
tains a single family, the Phlaeopthripidae (tube-tailed thrips). In Terebrantia,
the wings are held parallel at rest, and females lay eggs in plant tissue with ser-
rated, blade-like ovipositors. In Tubulifera, the wings, when present, overlap
each other when folded, and females lay their eggs in crevices or on plants.
Aeolothripidae are commonly called banded thrips due to the fact that
the wings may have cross bands or mottled patches. Some unusual species
are nearly wingless and may look very ant-like. They are generally yellowish-
brown to dark brown in colour, and in the females, the ovipositor is curved
upwards. Aeolothripids can be found on cruciferous and leguminous plants,
grasses, and conifers. Although also known collectively as predacious thrips,
most species are pollen feeders. The best-known and commonest genus is
Aeolothrips. The species Aeolothrips fasciatus has a very wide distribution, is

yellow to dark brown with banded wings, and is commonly found in the heads
of clovers. The nymphs are yellow or orange. The fully developed nymphs
form a silken cocoon underground before emerging as adults. Predacious
species are of use in controlling natural populations of other thrips, aphids,
and other small insects and mites.
The ovipositor in common thrips (Thripidae) bends downwards, not up-
wards, and these insects can be found on the leaves and flowers of a vast range
of plants. The vast majority are plant sap suckers, but some feed on fungal
spores, or are predacious. Reproduction can occur parthenogenetically, and
in some pest species (e.g., Common Greenhouse Thrip, Heliothrips haemor-
rhoidalis), males have been collected rarely. Females lay their smooth eggs
inside plant tissues. This family contains several genera (Limothrips, Thrips,
and Taeniothrips) with many pest species, some of which can do serious
damage by their feeding and the transmission of viruses to a wide range of im-
portant crops. Crops affected include peas, beans, citrus trees, cereals, coffee,
tea, tobacco, onions, cultivated flowers, and greenhouse produce. The Grain
Thrip (Limothrips cerealium) is a widespread pest that breeds in the ears of
cereal crops and emerges in vast numbers when the crop is ripe. Frankliniella
contains several pest species such as Frankliniella occidentalis (the Western
Flower Thrip). This species, which attacks vegetables and ornamental crops
grown under glass, is thought to be the most significant pest thysanopteran in
Europe. Thrips tabaci is a widespread and polyphagous pest that attacks crops,
such as cotton, tomatoes, and, as the name implies, tobacco. Apart from be-
ing the vector for a number of plant viruses, this species feeds mainly in the
young flowers and causes the young fruit to abort.
Tube-tailed thrips, which can be recognized by their tubular, pointed
abdominal end, are rather larger and stouter-bodied than aeolothripids or
thripids, and most are dark brown or black. This family contains some of the
world’s largest thrip species; a few tropical genera such as Phasmothrips, have
species measuring up to 12 mm in length. Tube-tailed thrips can be found
on a wide range of herbaceous plants, shrubs, and trees, in flowers, on twigs,
under bark, in soil, and in leaf litter. The majority of species feed on fun-
gal threads or spores, but there are predacious species that feed on mites and
small, soft insects like whiteflies. Liothrips vaneeckei damages lily bulbs, while
other species attack olives and grasses.
As expected, the parasitoids that use thrips as hosts are small species. Lar-
val thrips are parasitized mainly by wasp species in the Eulophidae, and thrip
eggs by wasp species in the Trichogrammatidae and Mymaridae. Some of
these parasitoids may have a significant role in controlling greenhouse pest
thrip species.
Thrips are a very old group, with putative fossils from the Permian (about
290 mya). The earliest definitive thrip fossils come from the Late Triassic,
about 200 mya. Some remarkable amber fossils from the Cretaceous of Spain
(100 mya) have trapped thrips covered in pollen grains of a cycad or gingko
tree. The thrips deliberately covered themselves with pollen, which they car-
ried off along the plant to be used as nourishment for their larvae. As the
THYSANOPTERA 155
thrips moved along the flowers, they pollinated them, suggesting that these
insects played a vital role in these ancient ecosystems—in fact, this fossil is
the earliest record of pollination in existence. In addition, the fact that these
thrips were deliberately collecting pollen for their young shows that they were
likely sub-social, like many thrips species today.
Key reading
Ananthakrishnan, T. N. (1993). Bionomics of thrips. Annual Review of Entomology 38:
71–92.
Crespi, B. J. (1988). Risks and benefits of lethal male fighting in the colonial, polygynous
thrips Hoplothrips karnyi (Insecta: Thysanoptera). Behavioral Ecology and Sociobiology
22: 293–301.
Crespi, B. J. (1992). Eusociality in Australian gall thrips. Nature 359: 724–6.
Crespi, B. J., Carmean, D. A., and Chapman, T. W. (1997). Ecology and evolution of galling
thrips and their allies. Annual Review of Entomology 42: 51–71.
Grimaldi, D., Shmakov, A., and Fraser, N. (2004). Mesozoic thrips and early evolution of
the order Thysanoptera (Insecta). Journal of Paleontology 78: 941–52.
Van der Kooi, C. J. and Schwander, T. (2014). Evolution of asexuality via different mecha-
nisms in grass thrips (Thysanoptera: Aptinothrips). Evolution 68: 1883–93.
Lewis, T. (ed) (1997). Thrips as crop pests. CAB International, Wallingford.
Loomans, A. J. M. and Van Lenteren, J. C. (1995). Biological control of thrips pests: a review
of thrips parasitoids. Wageningen Agricultural University Papers 95: 89–201.
Mound, L. A. and Kibby, G. (1998). Thysanoptera: an identification guide (2nd edn). CAB
International, Wallingford.
Mound, L. A., Heming, B. S., and Palmer, J. M. (1980). Phylogenetic relationships between
the families of recent Thysanoptera (Insecta). Zoological Journal of the Linnean Society
69: 111–41.
Palmer, J. M., Mound, L. A., and Du Heaume, G. J. (1989). CIE Guides to insects of importance
to man: 2. Thysanoptera. CAB International, Wallingford.
Peñalver, E., Labandeira, C. C., Barrón, E., Delclòs, X., Nel, P., Nel, A., Tafforeau, P., and
Soriano, C. (2012). Thrips pollination of Mesozoic gymnosperms. Proceedings of the
National Academy of Sciences of the United States of America 109: 8623–8.
Tommasini, M. G. and Maini, S. (1995) Frankliniella occidentalis and other thrips harmful
to vegetable and ornamental crops in Europe. Wageningen Agricultural University Papers
95: 1–42.
156
Division Endopterygota
(= Holometabola)
·····························································································
This monophyletic group contains the most successful insect orders. The
young stages are called larvae and look very different from the adults they
will become. Their wings develop internally (endopterygote) and metamor-
phosis is complete (holometabolous). The total transformation from larva to
adult takes place during a pupal stage.
Neuropterida
This superorder comprises the Raphidioptera (snakeflies), the Megaloptera
(alderflies, dobsonflies, and fishflies), and the Neuroptera (lacewings,
antlions, and allies). The Neuropterida are generally considered as one of
the earliest lineages of Holometabola, and likely represent the sister-group to
the Coleopterida, the superorder that includes Coleoptera (beetles) and the
Strepsiptera (twisted-wing insects). With more than 10,000 described species,
Neuropterida represents the fifth-largest superorder of Holometabola.
157
Raphidioptera
(snakeflies)
Common name Snakeflies Metamorphosis Complete (egg, larva,

Derivation Gk. Rhaphis–a needle; pupa, adult)
rhaphe–a seam; pteron–a Distribution Northern hemisphere
wing Number of families 2
Size 6–28 mm (body length) Known world species 248 (0.022%)
Forward-facing mouthparts
Head tapers
Ocelli
Elongate pronotum
Front and hind

wings similar Pterostigma
in size and
venation
Ovipositor (female only)
Phaeostigma notata — Raphidioptera
Key features
• prothorax very elongate, appearing as a long ‘neck’
• head elongate and slightly flattened
• ovipositor about as long or longer than body
• wings transparent
Snakeflies such as this

female Atlantoraphidia
© Paul Brock
maculicollis take their name
from their elongate head
and prothorax. UK.
These moderately shiny, dark-greyish, or dark reddish-brown insects are the

sister-group to all other Neuropterida. With fewer than 300 species, snakeflies
represent the smallest holometabolan order. The name snakefly comes from
the snake-like way in which the adults catch their prey. The head, which is
slightly flattened with biting, forward-facing mouthparts, is supported by an
elongate prothorax and can be raised and extended forward to seize food. The
head is broadest across the eyes and tapers behind to meet the prothorax.
The antennae are half as long (Raphidiidae) or as long (Inocellidae) as the
body.
Ocelli are present in the Raphidiidae but are absent in the Inocellidae.
The two pairs of wings are similarly sized, transparent, and have veins that
are forked close to the wing margins. Both pairs of wings have a small dark
or pale mark called the pterostigma on the front edge towards the wing tip.
The females, which are a little larger than males, have a long, slender, and
conspicuous ovipositor, which is about as long as the body, or distinctly
longer.
Snakeflies are unusual among most insects in that they reach their greatest
The typical slightly flattened diversity in cool, temperate regions of the Northern Hemisphere, particu-
head with biting, larly in the Mediterranean, Central Asia, and North America. The reasons
forwards-facing
underlying this peculiar distribution are unknown, although the observation
mouthparts, supported by
an elongate prothorax of that snakefly larvae require cool temperatures to pupate might represent an
the Raphidiidae. adaptation to a temperate climate.
RAPHIDIOPTERA 159
Raphidioptera are usually found in wooded areas especially among rank

vegetation. Adults and larvae are active during the day, and they are vo-
racious predators of aphids and other small soft-bodied arthropods such
as spiders and springtails. However, adult Raphidiidae may occasionally
consume nectar and pollen, whereas the Inocellidae may not feed at all.
Courtship in these insects involves prolonged vibrational signal duets be-
tween males and females. These ‘songs’ are produced by sustained abdominal
percussion and wing fluttering. Inocellid males may also fasten their head on
the female’s abdomen during mating, using unique abdominal organs. Sev-
eral hundred eggs are laid, in groups of up to 100, in slits and cracks in tree
bark or in rotting timber. The long, slender larvae have well-developed heads
and legs, are very agile, and hunt for food under loose bark, in rotting wood,
or in leaf litter. The short, curved, and pointed mandibles are similar to those
of the adults.
Pupation takes place in a simple chamber in rotten wood or leaf litter, and,
as there is no cocoon, the pupa can move around. The commonest genera are
Snakefly larvae are Raphidia, Agulla, and Xanthostigma.
terrestrial and live under Although definitive snakefly fossils are scarce, molecular estimates sug-
bark or in rotting wood gest that the earliest ancestors of extant species appeared during the early
(Raphidia sp.).
Cretaceous (about 136 mya). One would expect snakeflies to have appeared
much earlier, given that they are the sister-group to all other Neuropterida,
which are thought to have evolved more than 239 mya. However, diverse and
ancient lineages of tropical, warm-adapted snakeflies may have lived in the
Mesozoic but did not survive the climatic catastrophes of the Cretaceous–
Tertiary extinction event caused by the meteor impact that also killed the
non-avian dinosaurs. As a result, only their poorly diverse, cool-adapted
relatives survived, resulting in the temperate snakefly fauna we observe today.
The economic importance of Raphidioptera has not been sufficiently re-
searched. However, given that snakeflies are effective predators of aphids and
other arthropod pests, they may act as natural agents of pest control in certain
parts of the world.
Key reading
Aspöck, H. (1998). Distribution and biogeography of the order Raphidioptera: updated facts
and a new hypothesis. Acta Zoologica Fennica 209: 33–44.
Aspöck, U. and Aspöck, H. (2007). Verbliebene Vielfalt vergangener Blüte. Zur Evolution,
Phylogenie und Biodiversität der Neuropterida (Insecta: Endopterygota). Denisia 20:
451–516.
Aspöck, U., Aspöck, H., and Rausch, H. (1994). Die Kopulation der Raphidiopteren:
eine zusammenfassende Übersicht des gegenwärtigen Wissensstandes (Insecta: Neu-
ropteroidea). Mitteilungen der Deutschen Gesellschaft für Allgemeine und Angewandte
Entomologie 9: 393–402.
Engel M., Pérez de la Fuente, R., Peñalver, E., and Delclòs, X. (2012). Snakefly diversity
in Early Cretaceous amber from Spain (Neuropterida, Raphidioptera). ZooKeys 204:
13–40.
Henry, C. S. (2006). Acoustic communication in neuropterid insects. In: Insect sounds
and communication. Drosopoulos, S. and Claridge, M. F. (eds), pp. 153–66. Taylor and
Francis, Boca Raton.
Oswald, J. D. (2015). Neuropterida species of the world: a catalogue of the species-group

names of the extant and fossil Neuroptera, Megaloptera, Raphidioptera and Glosse-
lytrodea (Insecta: Neuropterida) of the World. Lacewing Digital Library v. 4.0. Retrieved
from: http://lacewing.tamu.edu/SpeciesCatalog/Main.
Vasilikopoulos, A., Misof, B., Meusemann, K., Lieberz, D., Flouri, T., Beutel, R., Niehuis,
O., Wappler, T., Rust, J., Peters, R., Donath, A., Podsiadlowski, L., Mayer, C., Bartel, D.,
Böhm, A., Liu, S., Kapli, P., Greve, C., Jepson, J., and Aspöck, U. (2020). An integrative
phylogenomic approach to elucidate the evolutionary history and divergence times of
Neuropterida (Insecta: Holometabola). BMC Evolutionary Biology 20: 64.
161
Megaloptera
(alderflies, dobsonflies, and fishflies)
Common name Alderflies, dobsonflies, Metamorphosis Complete (egg, larva,

and fishflies pupa, adult)
Derivation Gk. megalo–large, great; Distribution Widespread but mainly
pteron–a wing temperate regions
Wingspan 18–170 mm Known world species 373 (0.033%)
Thread-like
antenna
Compound eye
Meso- and
metathorax
Wings broad similarly sized
at base
Sialis lutaria — Megaloptera
Key features
• aquatic larvae with distinct lateral abdominal gills
• wings of roughly equal size
• medium-sized to very large insects
• adults always near water
Dobsonflies, such as this

Protohermes motuoensis
represent some of the largest
© George McGavin
Neuroptera. Although they

are voracious aquatic
predators as larvae, adults
probably do not feed at all.
Northern India.
There are two families, the alderflies (Sialidae) and the dobsonflies and fish-
flies (Corydalidae). The head carries a pair of prominent compound eyes and
unmodified, thread-like antennae. Ocelli are present in corydalids but absent
in sialids. Despite their conspicuous jaws, adult megalopterans do not feed.
In some male Corydalidae, the jaws may be hugely enlarged to several times
the length of the head and are used in male-to-male combat or for grasp-
ing the female during mating. These are known as the dobsonflies. Despite
the alarming proportion of their jaws, dobsonflies are harmless, although it is
just conceivable that their larvae, known as hellgrammites or toe-biters, might
occasionally earn their vernacular name. Male Corydalidae of the subfamily
Chauliodinae are known as the fishflies and lack the distinctly enlarged jaws.
Megalopterans do not fly readily, and when they do, they flutter rather
weakly and alight readily. There are two pairs of wings with a branching
pattern of venation. The hind wings are broad at their bases, due to the pres-
ence of large anal lobes, and are usually larger than the front wings. At rest,
the wings may be held roof-like over the body or may appear to be almost
wrapped around the body. Megalopteran adults are found near cool, clean
MEGALOPTERA 163
streams. Dobsonflies prefer running water, while alderflies can be found in

ponds and canals as well as streams.
Mating in Megaloptera takes place on marginal vegetation or on the
ground. In Sialidae, both sexes locate each other and communicate by rhyth-
mically tapping their abdomen and wings on the substrate, which produces
species-specific low-frequency vibrations that they sense through their legs.
In Corydalidae, only males produce these vibrational signals, as part of a pre-
mating ritual. The males of certain male dobsonfly (Protohermes) and fishfly
species (Parachauliodes, Neochauliodes) are known to attach a jelly-like exter-
nal spermatophore to the genitalia of the female. The spermatophore contains
bundles of sperm in seminal fluid, which are transferred to the female over
the course of several hours. Males remain in copula throughout this time, for,
if they separate, the female may eat the spermatophore before all the sperm
have been transferred successfully. Females produce large numbers of eggs,
which they stick to reeds and other plants near or hanging over the water.
The egg masses may frequently resemble bird droppings, which provides an
effective camouflage against many predators.
At rest, the wings are held The larvae of all megalopterans are aquatic, highly predacious, with simple
together roof-like over the or branched abdominal gills, and can be very similar in general appearance to
body.
larvae of some water beetle families. When the young larvae hatch, they drop
or crawl into the water using their well-developed legs. In alderflies, the larval
proleg abdomen has a single terminal filament and seven pairs of lateral filaments.
a fleshy abdominal limb of The abdomen of a dobsonfly larva has a terminal pair of small hook-like
insect larvae prolegs.
The larvae hide or burrow, and ambush a range of arthropod prey, particu-
larly small crustaceans, and will also take worms, amphibians, and even small
fish. Larval development goes through a maximum of 11 instars and can take
anything from 12 months in alderflies, but sometimes more than 48 months
in dobsonflies. The time spent as a pupa is relatively short and can be as little
as two weeks. Unlike species in the closely related order Neuroptera, the fi-
nal larval stage does not spin a cocoon in which to pupate. Instead, pupation
takes place on land within a simple chamber made in moist soil, sand, mossy
vegetation, or under rotting wood. The pupae have functional jaws, can move
around freely, and may even protect themselves if attacked.
The fossil record of Megaloptera is rather poor, with specimens that can be
confidently assigned to Megaloptera being known from the Early and Mid-
dle Jurassic (about 200–170 mya). Molecular divergence estimates suggest
that the common ancestor of extant Megaloptera appeared in the Trias-
sic about 239 mya. Most morphological and molecular studies suggest that
Megaloptera represent the sister-group to the Neuroptera. Megaloptera are
of no direct economic importance, although they may serve as bioindicators
The aquatic larva of an of freshwater ecosystems.
alderfly, showing the single
terminal filament and seven
pairs of lateral filaments, Key reading
Sialis sp.
Hayashi F. (1996). Insemination through an externally attached spermatophore: bundled
sperm and post-copulatory mate guarding by male fishflies (Megaloptera: Corydalidae).
Journal of Insect Physiology 42: 859–66.

and communication. Drosopoulos, S. and Claridge, M. F. (eds), pp. 153–66. Taylor and
Liu, X., Wang, Y., Shih, C., Ren, D., and Yang, D. (2012). Early evolution and historical
biogeography of fishflies (Megaloptera: Chauliodinae): implications from a phylogeny
combining fossil and extant taxa. PLoS ONE 7: e40345.
Mangan, B. P. (1994). Pupation ecology of the dobsonfly Corydalus cornutus (Corydalidae:
Megaloptera) along a large river. Journal of Freshwater Ecology 9: 57–62.
Merritt, R. W. and Cummins, K. W. (eds) (1996). An introduction to the aquatic insects of
North America (3rd edn). Kendall/Hunt, Dubuque.
New, T. R. and Theischinger, G. (1993). Megaloptera (alderflies, dobsonflies). Handbuch der
Zoologie Band IV, Arthropoda, Insecta, Tome 33. de Gruyter, Berlin.
Rupprecht, R. (1975). Die Kommunikation von Sialis (Megaloptera) durch Vibrationssig-
nale. Journal of Insect Physiology 21: 3057–313, 3157–320.
Vasilikopoulos, A., Misof, B., Meusemann, K., Lieberz, D., Flouri, T., Beutel, R., Niehuis,
Böhm, A., Liu, S., Kapli, P., Greve, C., Jepson, J., and Aspöck, U. (2020). An integrative
Whiting, M. F. (1994). Cladistic analysis of the alderflies of America North of Mexico
(Megaloptera: Sialidae). Systematic Entomology 19: 77–91.
165
Neuroptera
(lacewings, antlions, and mantidflies)
Common name Lacewings, antlions, and Metamorphosis Complete (egg, larva,

mantidflies pupa, adult)
Derivation Gk. neuron–sinew, nerve; Distribution Worldwide
pteron–a wing Number of families 18
Size Body length 2–90 mm. Known world species > 5,820 (0.51%)
Wingspan 1.8–150 mm
Thread-like antenna Conspicuous compound eye
Front and
hind wings Characteristic
similar in net-like
size and venation
venation
Thoracic segments Veins forked

of similar size at wing margin
Chrysoperla carnea — Neuroptera
Key features
• elongate bodied
• prominent eyes
• wing venation complex and net-like
• larval mandibles and maxillae form sucking mouthparts
© Rupert Soskin
Ascalaphids are among the

most beautiful neuropterans.
Libelloides coccajus, France.
Members of this order have a worldwide distribution, although many families

are restricted to particular regions. The vast majority of adult neuropterans
are predatory (with a few nectar-feeding exceptions) and are mainly active in
the evening or after dark. The larvae of all species are highly predacious and
can be found in a wide range of habitat types, even freshwater, where larvae
of the Sisyridae predate on sponges.
The species of some families exhibit remarkable convergent similarities
to other insect groups. Adult Mantispidae have enlarged, raptorial front legs
similar in design and function to those of praying mantids. Owlflies (As-
calaphidae) are efficient aerial predators which look similar to dragonflies in
general appearance, and also catch prey while flying.
In general, adult neuropterans have biting, chewing mouthparts and a pair
of conspicuous, laterally placed compound eyes, which, in some species, ap-
pear to shine. Ocelli may or may not be present. The antennae can be short,
about as long as the head and thorax combined, but are generally much longer
and thread-like. In some owlflies and antlions, the end of the antennae may be
swollen to form a club. The adults of some families have prothoracic glands
NEUROPTERA 167
© George McGavin
Lacewing larvae consume
vast amounts of aphids and
other soft-bodied insect
during their lifetime. UK.
capable of producing substances that repel some predators. There are usu-
ally two pairs of similarly sized wings held roof-like over the body at rest. The
venation in most families is net-like with the main veins forking at the wing
margins in many families. Exceptions include the Coniopterygidae, which
comprise very small species with a white, powdery covering, where the hind
wings are often reduced, and the Nemopteridae, whose hind wings are devel-
oped into long, tail-like structures. In these species, it has been shown that
the long hind wings may deter smaller predatory insects such as robber flies
(Asilidae) by making the nemopterid appear bigger than its actual size.
Although courtship and mating have been described in relatively few
species, it seems that all families produce species-specific vibrational songs
and pheromones. Male and female green lacewings produce low-frequency
vibrations by vertical abdominal vibration. These vibrations, which are pro-
chordotonal organs duced in short bursts every second with a frequency typically less than 80
subcuticular Hz, are transmitted through the legs to the foliage. The legs of lacewings have
mechanoreceptors that pick
up vibrational signals—made different types of chordotonal organs, of which the subgenual organ in the
up of one or more units called tibiae, for the reception of substrate-borne vibration, is the most important.
scolopidia
The males usually sing first, and the females respond at species-specific time-
frames. These courtship duets are a prelude to mating and have been shown to
play a major role as a reproductive isolation mechanism in the genus Chrysop-
erla and may be the primary force driving the rampant sympatric speciation
observed in this genus.
Larval morphology is varied and very much linked with lifestyle. Free-
living hunters are slender with longish legs, and many have specialized
adhesive disc-like structures on the last two abdominal segments for clinging
A lacewing larva (Chrysopa onto foliage. The bodies of pit-building or ambushing species tend to be fat
spp.) sucks the body fluids
from an aphid using sharp, and squat with short legs, very large jaws, extended necks, and other modifica-
curved mandibles. tions. The strong, sickle-shaped mouthparts of the larvae are highly modified
as sucking needles, a character that clearly separates this order from the Mega-
loptera. The sharply pointed and curved mandibles have a groove that runs
along their internal surface. Pressed tightly against the groove, an elongated
maxilla forms a hollow tube, through which the larvae can inject enzymes
and suck the juices of their prey.
The alimentary tract of larval neuropterans is peculiar in that the hindgut
is not united with the midgut. During larval life, only fluids are expelled and
any solid material that accumulates remains in the midgut, only to be passed
meconium as a meconium when the adult first emerges from the pupa with its gut joined
the first excrement passed by up. This characteristic is also found in most members of the Hymenoptera.
a newly emerged adult
The larvae of Mantispidae are parasitic on spider egg sacs or the larvae of
hymenopterans, such as social wasps or bees. In Mantispa uhleri, the mobile
first instar larvae find an adult spider and climb aboard to await the pro-
duction of an egg sac, inside which they will feed. To keep going until this
happens, the young mantispid sucks the spider’s haemolymph. Once inside
the egg sac or host cell, the larva moults and becomes more maggot-like with
short legs. The larvae of Ascalaphidae are ‘sit-and-wait’ predators that lie in
debris or rest on plants to seize anything suitable that comes within range
of their jaws. A unique predatory strategy may be found in the larvae of the
beaded lacewing (Berothidae) species Lomamyia latipennis. This species lives
in the colonies of the termite Reticulitermes hesperus. When a hungry larva
encounters a termite, it raises the tip of its abdomen, and waves it in front of
the termite’s head, releasing an unknown chemical. The effect of this chemi-
cal is to paralyse the termite, which allows the larva of Lomamyia to feed on it
undisturbed. This phenomenon has fascinated entomology enthusiasts, who
refer to this predatory strategy as the ‘death fart’. However, observation of this
strategy has never been repeated since the original study, and the chemical
involved is unknown.
In the majority of Neuroptera, larvae are mature after three instars and
pupate inside a fragile, spherical, silk cocoon spun from special organs
(modified malpighian tubules) at the end of their abdomens. Sometimes the
cocoon will incorporate small sand grains or soil particles. The pupae are
similar to those of the Megaloptera in that they have free appendages and
functional mouthparts, which are used to open the cocoon before the adult
emerges from its pupal exuvia.
Many neuropterans are directly or indirectly beneficial in controlling pop-
ulations of mites and small, soft-bodied animals, such as aphids, mealybugs,
and scale insects. The two biggest neuropteran families are the common or
green lacewings (Chrysopidae) and antlions (Myrmeleontidae). Common or
green lacewings are generally green in body colour, the wings often with del-
icate tints of pink, green, and blue. The wings of chrysopids are distinctive in
that they have two seemingly zigzag veins and lack any dark patterning. The
eyes are bright golden, brassy, or reddish. Both adults and their larvae are vo-
racious predators of aphids, thrips, psyllids, scale insects, coccids, and mites,
although, in some species, the adults will eat pollen, nectar, and honeydew.
The adults are mainly nocturnal in habits, and species of the largest subfam-
ily, the Chrysopinae, have bat-detecting, ultrasonic sound receptors located
NEUROPTERA 169
in the large veins of the wings. Chrysopid eggs are laid on vegetation and, like
the eggs of most species, are supported by distinctive long, delicate stalks.
The larvae, which are often pale yellow, white, or greenish, with brown or
black markings, can disguise themselves by attaching the sucked-out bodies
of their victims on to special, hooked dorsal hairs. The disguise helps them
to move about, camouflaged in the colony and may protect them from birds.
A typical lacewing showing Some larvae that are predacious on mealy bug colonies gather the mealy bugs’
the complex wing venation. waxy coverings and use them to build up a protective cover on their own
backs. Although soft-bodied and weak, aphids can fight back. Many species
of aphid produce a sticky substance from their abdominal cornicles, which
is designed to harden inside and block the hollow jaws of lacewing larvae.
Green lacewings pupate in pea-shaped, silk cocoons attached to the under-
sides of leaves. Adults will often enter houses to hibernate and are attracted to
lights. Chrysopa, the largest genus in the entire family, is very common and
widespread. Chrysoperla carnea and other species have been reared in large
numbers for use as a biological control agents in various crops.
Brown lacewings (Hemerobiidae) are less common than green lacewings,
and can be found in deciduous woodlands, gardens, and hedgerows. The eggs
are laid on plants but are not stalked. The larvae do not have any body warts
or tubercles, and their body hairs are simple and not hooked.
Adult antlions (Myrmeleontidae) superficially resemble large damselflies
Chrysopid eggs supported but are softer and have short knobbed or club-ended antennae, which are
above the leaf surface by
long delicate stalks. about twice as long as the head. The wings are long and narrow and may
be clear or have distinctive brown or black patterns. The abdomen is elon-
gate and, in males, has terminal sexual clasping organs that look a bit like the
forceps of earwigs. Open woodland, scrub grassland, dunes, and warm, dry
sandy areas are typical habitats for antlions. Although a few eat pollen, most
adult antlions are carnivorous, and seize insects from vegetation or on the
wing. The females lay single eggs in soil or sand. Some larvae construct insect-
trapping pits in loose sand, while others live as ambush prey on the trunks
of trees in soil or under stones and debris. All antlion larvae, sometimes
called doodlebugs, are carnivorous. They are broadly oval, with very large
jaws armed with sharp spinous teeth for preying on a wide range of insects
and spiders. In the best-known species, such as those of the genus Myrmeleon,
the larvae live in sand at the bottom of conical pits with only their sharp jaws
showing. Where the antlions construct their pit is dependent on a number of
abiotic factors, such as shelter and substrate type. If population densities are
high, some species will space each other by means of sand flicking and other
forms of display. If too much sand is flicked into a pit from a neighbouring
pit, the antlion will usually relocate. Ants or any other insect stumbling into
the pit will tumble to the bottom or be showered with sand grains flicked up
by the antlion. Once at the bottom of the pit, the prey is seized and, when
sucked dry, is flicked out before the larva renovates its pit. The size of the
sand grains is important. Ants are much more likely to escape from the pit
if the grains are too big. It is not surprising that, in such a specific interac-
tion, ants have learned to avoid antlions, or areas where the larvae are likely
to be lurking. Some experiments on ground-foraging ant species show that
experienced ants will not be tempted into high-risk areas even by attractive
baits and will continue to avoid areas from which antlions have been removed.
The larvae of some Australian antlions, Callistoleon manselli and Callistoleon
illustris, have taken the art of pit building a stage further by adding furrows or
trenches radiating from the central pit. The extra construction work is worth
the effort as the furrows increase the prey capture rate. However, the building
behaviour is plastic, and because the furrows are added after the central pit is
dug, the larvae will not bother to make them if they are supplied with plenty
of prey.
The evolutionary history of Neuroptera goes very far back in time, with
phylogenomic estimates suggesting an origin in the Permian (about 280 mya),
which is in agreement with the available fossil evidence. Many of the mod-
ern families and their behaviours appeared early on. For example, lacewing
larvae from Early Cretaceous Lebanese amber (about 130 mya), were found
to be camouflaged with bits of debris attached on their body through special-
ized hairs. The debris particles formed a protective dome-like structure that
shielded the larvae from attacks (presumably from their hosts), a behaviour
that is found in many lacewing larvae today.
Key reading
Canard, M., Semeria, Y and New, T. R. (eds) (1984). Biology of the Chrysopidae. Entomo-
logica series 27. Junk Publishers, The Hague.
Brooks, S. J. and Bernard, P. C. (1990). The green lacewings of the world: a generic review.
Bulletin of the British Museum (Natural History) Entomology 63: 137–210.
Devetak, D. and Pabst, M. A. (1994). Structure of the subgenual organ in the green lacewing,
Chrysoperla carnea. Tissue and Cell 26: 249–57.
Engel, M. S., Winterton, S. L., and Breitkreuz L. C. V. (2018). Phylogeny and evolution
of Neuropterida: where have wings of lace taken us? Annual Review of Entomology 63:
531–51
Gotelli, N. J. (1996). Ant community structure: effects of predatory antlions. Ecology 77:
630–8.
Henry, C. S. (1977). The behaviour and life histories of two North American ascalaphids.
Annals of the Entomological Society of America 70: 117–95.
Henry, C. S. (1979). Acoustical communication during courtship and mating in the green
lacewing Chrysopa carnea (Neuroptera: Chrysopidae). Annals of the Entomological
Society of America 72: 68–79.
Henry, C. S. (1980). Acoustical communication in Chrysopa rufilabris (Neuroptera:
Chrysopidae), a green lacewing with two distinct calls. Proceeding of the Entomological
Society of Washington 82: 1–8.
Henry, C. S. (1985). Sibling species, call differences, and speciation in green lacewings
(Neuroptera: Chrysopidae: Chrysoperla). Evolution 39: 965–84.
Henry, C. S. (1994). Singing and cryptic speciation in insects. Trends in Ecology and
and communication. Drosopoulos, S. and Claridge, M.F. (eds), pp. 153–66. Taylor and
Johnson, J. B. and Hagen, K. S. (1981). A neuropterous larva uses an allomone to attack
termites. Nature 289: 506–7.
New, T. R. (1975). The biology of the Chrysopidae and Hemerobiidae (Neuroptera),
with reference to their usage as biocontrol agents: a review. Transactions of the Royal
Entomological Society of London 127: 115–40.
NEUROPTERA 171

Pérez-de la Fuente, R., Peñalver, E., Azar, D., and Engel, M.S. (2018). A soil-carrying
lacewing larva in Early Cretaceous Lebanese amber. Scientific Reports 8: 16663.
Redborg, K. E. (1982). Interference by the mantispid Mantispa uhleri with the development
of the spider Lycosa rabida. Ecological Entomology 7: 187–96.
Vasilikopoulos, A., Misof, B., Meusemann, K. Lieberz, D., Flouri, T., Beutel, R., Niehuis,
Böhm, A., Liu, S., Kapli, P., Greve, C., Jepson, J., and Aspöck, U. (2020) An integrative
Matsura, T. and Kitching, R. L. (1993). The structure of the trap and trap building be-
haviour in Callistoleon manselli New (Neuroptera: Myrmeleontidae) Australian Journal
of Zoology 41: 77–84.
Wells, M. M. and Henry, C. S. (1992). The role of courtship songs in reproductive iso-
lation among populations of green lacewings of the genus Chrysoperla (Neuroptera:
Chrysopidae). Evolution 46: 31–42.
172
Coleopterida
This grouping, which includes beetles (Coleoptera) and twisted-wing insects
(Strepsiptera), is the most diverse animal lineage, with more than 400,000
species.
173
Coleoptera
(beetles)
Common name Beetles Metamorphosis Complete (egg, larva,

Derivation Gk. Koleos–sheath; pupa, adult)
pteron–a wing Distribution Worldwide
Size Body length 0.1–180 mm. Number of families 170
Mostly under 25 mm Known world species About 400,000 (35.34%)
Terminal segments
forming an
antennal fan
Compound
eye
Front tibia Pronotum
Scutellum
Front wings
toughened
as elytra
Elytra meet
down body
Hindleg midline
Melolontha melolontha — Coleoptera
Key features
• very abundant and ubiquitous in all terrestrial and freshwater ecosystems
• front wings modified as rigid elytra covering hind wings
• prothorax usually large, and distinct from the head and rest of the body
• many species are crop pests
• the largest order
© George McGavin
The stag beetle’s (Lucanus
cervus) impressive
mandibles are used in
male-to-male combat. UK.
Click beetles (Elateridae) are

able to escape from
© Zestin Soh
predators by using a
jumping mechanism on the
underside of their thorax.
Singapore.
COLEOPTERA 175
Giraffe weevils
(Trachelophorus giraffa)
from Madagascar need no
explanation as to their
common name. The long
‘neck’ of the male is an
extension of the head, used
© Zestin Soh
to construct nests for the
females and in male-to-male
combat.
Chitin, the major component of arthropod cuticle, was first described as pro-
teinaceous in nature from a study of the wing cases of the cockchafer Melolon-
tha published by Odier in 1823. The protective front wing cases have con-
tributed greatly to the success of beetles, and their tough nature has a useful
spin-off for archaeologists and palaeoecologists. The wing cases, or even just
fragments of wing cases, and other body parts in sediments and deposits are
often recognizable to the level of species. As the host plants and habits of these
beetles are unlikely to have changed much in the last 1.5 million years, build-
ing up a picture of exactly which species were present can provide invaluable
details of what plants were growing at the time and what the habitat was like.
Identifying beetle communities in archaeological excavations serves as an in-
valuable tool for understanding past climate, habitat modification by humans,
their diet and subsistence methods, as well as methods of food storage.
Many male beetles such as

this cockchafer, have large
comb-like antennae, which
© Anne Riley
are covered with olfactory

receptors that allow them to
locate females at a distance.
UK.
Beetles play an essential role

in the recycling of nutrients
in virtually every terrestrial
ecosystem. Cockchafer
larvae decompose plant
© Rupert Soskin
debris, allowing for living
plants to extract the
nutrients from decaying
matter. UK.
© Rupert Soskin
The corn weevil (Sitophilus

zeamais) is a notorious pest
of cereal products. France.
COLEOPTERA 177
There is now some doubt as to whether the British polymath J. B. S. Hal-

dane really said ‘an inordinate fondness for beetles’, in reply to being asked
what could be inferred of the Creator by studying the works of creation, but
it is a good story. There can be no doubt that beetles really are the most suc-
cessful multicellular animals. They comprise at least 40% of all known insect
species and about 25% of all known animals. There are more species of beetles
than there are species of plant, and, like all other insect groups, many species
await discovery and description. It is estimated that the total number of beetle
species (including those yet undescribed) ranges between 1.5 to 2.1 million.
© Rupert Soskin
Cerambycid beetles like
Rhagium sycophanta are the
primary decomposers of live
and decaying wood. France.
Ladybirds are voracious

predators of aphids and
other agricultural pests, and
for this reason they can be
© Anne Riley
exceptionally beneficial for

gardens and eco-friendly
crops. Coccinella
septempuncta, UK.
The species comprising the order are classified into four very unequal
suborders (in terms of species number): the Archostemata, Myxophaga,
Adephaga, and Polyphaga (see Box).
Coleopteran suborders
Suborder Major families Habits

Archostemata 3 Primitive families: Cupedidae, Mostly found in decaying
Micromalthidae (1 sp.), Omatidae wood, fungivorous
·········································································································································································
Myxophaga 4 Families: Cyathoceridae, Hydroscaphi- Aquatic or associated with
dae, Microsporidae, Torridincolidae wet habitats, algal feeders
·········································································································································································
Adephaga 11 Families including Carabidae, Terrestrial and aquatic,
Dytiscidae, Gyrinidae mostly predacious
·········································································································································································
Polyphaga > 149 Families including Anobiidae, Variable: mostly herbivo-
Bruchidae, Buprestidae, Cerambyci- rous, some saprophagous,
dae, Chrysomelidae, Coccinellidae, wood-boring, predacious,
Curculionidae, Dermestidae, Elateridae, fungivorous, etc.
Scarabaeidae, Scolytidae, Staphylinidae,
Tenebrionidae
Beetles can be found in every conceivable terrestrial and freshwater habi-

tat, from the equator to the polar regions. One of the features that has enabled
them to become so successful, and the most dominant insect taxon, is the
front pair of wings, which are toughened and act as wing cases or elytra to pro-
tect the much larger, membranous hind wings, which are folded underneath
at rest. In all species, the elytra meet down the midline of the body, and, in
some species, they are permanently fused together. In most species, the elytra
cover most of the thorax and abdomen, but, in some (e.g., the Staphylinidae),
they are shortened (as in earwigs) to allow greater abdominal flexibility. In
these species, the hind wings are folded double, and the dorsal surface of the
abdominal segments are specially toughened to compensate. The strength of
the body and the mechanical protection afforded by the sclerotized elytra has
enabled the beetles to safely burrow, dig, and squeeze into places that other in-
sects cannot reach. Wherever you look, under bark and stones, in the ground,
inside plants, in leaf litter, in caves, in dung, and in carcasses, beetles have
been able to colonize successfully. Additionally, the elytra provide protection
from predators, and, as they also cover the abdominal spiracles, they cut down
on water loss. The space beneath the elytra allows aquatic species to maintain
an air store while submerged. Beetles have chewing-biting mouthparts and
usually have antennae with fewer than eleven segments. They are generally
compact and tough-bodied, and some species can be so armoured that en-
tomologists find great difficulty in pushing even the sharpest stainless-steel
pin through their body. The joints where body and limb segments meet are
often specially strengthened, recessed, or retractile, so that not even the tini-
est chink is available for predatory jaws or an ovipositor to gain access. The
body shape has evolved to suit a multitude of functions and environments.
Long-legged species for desert life; smooth, streamlined species for aquatic
life; robust, tank-like species for earth moving; seed-shaped mimics to deceive
predators; and so on.
COLEOPTERA 179
Among the beetle multitudes there are many scavengers, predators, and a
few specialized parasites, but the vast majority of beetle species are herbivo-
rous and here lies the second major reason for their great success. The first
beetles that arose in the Permian were detritivores or fungivores, but it did
not take them long (50 million years or so) to move on to conifers and cycads,
which were spreading in the Triassic and Jurassic. The early herbivorous bee-
tles chewed particular plant parts, such as the pollen-bearing structures, and
the numbers of these tissue-feeding taxa rose steadily. The rise to dominance
of the flowering plants (Angiospermae) in the Cretaceous provided beetles
with a multiplicity of opportunities to radiate. Already well adapted to eat-
ing plant tissues, more and more herbivorous beetles appeared. As quickly as
the evolution equipped the plants with chemicals and other defences against
herbivores, the herbivores evolved counter-measures. Enzymes appeared to
deal with plant poisons, or the beetles simply shifted their attention to a dif-
ferent, less well-defended plant part. Each evolutionary thrust and parry led
to the appearance of more and more species. The 100-million-year-long co-
evolutionary contest between beetles and flowering plants has produced very
species-rich superfamilies, such as the leaf beetles (Chrysomeloidea) and the
weevils (Curculionoidea), and today their interactions represent the most
familiar biological association on the planet.
Most beetles reproduce sexually. Mate attraction and courtship involves
sound production, pheromones, and light displays. Fireflies and glow worms
belong to the family Lampyridae. The most distinctive feature of these beetles
is their ability, shared with some click beetles (Elateridae) and other related
families (e.g., Phengodidae), to produce light from special luminous organs.
The adults of some species and the larvae of all species are predacious, feeding
mostly on land snails. The luminous organs on the underside of the abdomen
are covered by transparent cuticle, and the beetles regulate the flash interval
and duration by controlling the oxygen supply to the organs, where a chemi-
cal reaction involving a substance called luciferin and the enzyme luciferase,
produces a cold, greenish light.
Mating in beetles always takes place with the male holding on to the
female’s back. Sometimes the male will remain in copula for many hours
or days, even though sperm transfer has been completed. This post-
insemination mate guarding is just one form of paternity assurance seen in
beetles. One odd example concerns the incredibly long front legs of the male
Harlequin Beetle, Acrocinus longimanus (Cerambycidae). Before naturalists
saw living specimens, they imagined that the beetles used their legs to brachi-
ate through vegetation like miniature gibbons, but their real purpose is just as
unusual. As in most insects, sperm is used on a last-in, first-out basis, and it
is important that the female does not mate with another male before she lays
her eggs. The last male to mate before egg laying will fertilize the lion’s share
of the eggs. To protect his mate from rivals, and therefore his investment, the
male Harlequin Beetle uses his long front legs as a moveable corral until the
eggs are laid.
The females of most beetles lay eggs. Some species are ovoviviparous, and
the eggs hatch inside the reproductive tract of the female and she gives birth to
larvae. Beetle larvae are very variable in appearance, but all have toughened
head capsules with forward-facing, biting mouthparts, and short antennae
having a maximum of four segments. There are four distinctive larval types:
campodeiform larvae, like those of ground beetles or water beetles, are ac-
tive predators with well-developed legs and large jaws; eruciform larvae, like
those of the leaf beetles, are herbivorous with short or very short legs; soft,
white, C-shaped, scarabaeiform grubs are typical of scarabaeid beetles and
related groups and have three pairs of thoracic legs and are found in rotting
plant tissue or animal dung; and legless apodous grubs are typical of weevils,
click-beetles, and other beetles that bore or gall plants. There are never any
leg-like structures on the abdominal segments, and, when larval development
is complete, pupation occurs and the transformation to the adult stage be-
gins. Beetle pupae have immobile mouthparts and free appendages (they are
not firmly attached to the body surface). Many species pupate inside a cham-
ber underground or inside the larval food plant, and, sometimes, a cocoon is
made.
Beetle larvae can be herbivorous, carnivorous, or saprophagous, and many
species develop in wood, dung, carrion, and fungi. There are a number of in-
teresting parasitic families. Fewer than 60 species belong to the Rhipiceridae,
whose larvae are endoparasites of cicada nymphs. The young active beetle
larvae grab hold of a cicada nymph before it goes underground. The 2,000
or so world species of oil or blister beetles (Meloidae) are parasitic, or rather
predacious, on eggs of grasshoppers, or the larvae of some Hymenoptera. The
larvae of bee parasites wait on flowers for the correct species of host to visit,
and then cling to the body hairs to enable transportation back to the nest.
Fewer than 500 species belong to the Rhipiphoridae, the larvae of which are
endoparasites of cockroaches, wasps, and bees.
The vast majority of beetles are terrestrial and less than 3% are aquatic.
Beetles in some families (Gyrinidae, Haliplidae, Hygrobiidae, Noteridae, and
Dytiscidae) spend most of their life history in aquatic environments, while
others may be aquatic only as larvae or as adults. Pupation is terrestrial.
Species in the Hydraenidae are only aquatic as adults, and some beetles be-
longing to families such as the Curculionidae and Chrysomelidae are only
aquatic as larvae.
There are more than 4,000 species of diving beetles (Dytiscidae), a
widespread family of water beetles where both larvae and adults are carniv-
orous. The adults have a streamlined body shape and long, slightly flattened,
hair-fringed hind legs for efficient swimming. The males of many species have
special, swollen front tarsi, which are used to grip the female during mating.
These beetles store air under their wing cases and when they need to obtain
more air, they rise to the water’s surface tail first (the tip of their abdomen).
Detail of the front leg of the Diving beetles have fully developed wings and can fly very well.
Dytiscus marginalis male
showing the enlarged tarsal The larvae and adults of whirligig beetles (Gyrinidae) eat mosquito larvae.
segments. There are about 800 species of these small beetles, which get their common
name from the characteristic circular, jerky way in which they move over the
surface of ponds and slow-moving streams. At first glance, they look as if they
only have front legs as the middle and hind legs are very short, stout, and
COLEOPTERA 181
paddle-like. The front pair of legs is used for catching food. The eyes of these
beetles are interesting in that they are divided horizontally into two zones:
one for above-water vision and one for below-water vision.
In the same suborder (Adephaga), the Carabidae, or ground beetles, are a
large family of well over 40,000 species worldwide.
Although most of these beetles are dark coloured, some species can be
metallic or brightly coloured. As major predators in many natural and man-
aged habitats, ground beetles are important elements in food chains. They
regulate pests and disseminate insect viruses and other entomopathogens,
and they also serve as useful indicator species in conservation studies. Many
carabids are specialist hunters and can, for example, follow the mucus trails
Larva of Dytiscus marginalis left by snails and slugs. The larvae of tiger beetles (Cicindellinae, which are
taking air at the water often considered a family in their own right), live in vertical tunnels in soil
surface.
or sand. From these tunnels they ambush passing prey and drag them un-
derground. The larvae have a flattened top of the head and pronotum, which
form the lid of the burrow.
More than 500 species of bombardier beetles (Brachininae) are known
worldwide. The most common genus is Brachinus. These beetles have a highly
unusual and impressive defence mechanism. Accidentally standing barefoot
on two bombardier beetles simultaneously on a shower room floor in the
tropics was, for one of us (G McG), an unforgettable lesson. Like both barrels
of a miniature shotgun being discharged, the beetles escaped but left the au-
thor with two patches of discoloured skin, which persisted for weeks. This is
due to the fact that bombardier beetles are able to direct a stream of boiling-
hot quinones from the rear of their abdomens to ward off predators. In some
species, the stream consists of up to 500 pulses per second. The hot quinones
result from a chemical reaction between hydroquinones and hydrogen perox-
ide in the abdomen. These compounds are secreted by cells and fill a reservoir
from where they are passed through a non-return valve to a cuticle-lined re-
action chamber. Here the mixture is acted on by oxidative enzymes, which
A typical ground beetle causes a very rapid exothermic reaction that releases oxygen, and, in doing
(Carabidae). so, boils part of the contents. The rapid rise in pressure due to the production
of gases forces the liquid out of the chamber with a relatively loud popping
sound—like a mini-explosion. The effect on predators, both small and large, is
immediate. Creationists have stated gleefully that these beetles provide them
with an irrefutable example of ‘special creation’. However, their argument is
easily dismissed in much the same way as their creationist view of the evolu-
tion of the vertebrate eye. They argue that it is impossible to see how such a
complex system, which requires more than two essential components, could
have evolved by chance alone. They point out that the reaction chamber or
either of the two chemicals are useless in isolation, but their inability to see
how the system could have evolved by natural selection stems as much from
a lack of imagination as a lack of knowledge.
This is how the system may have evolved by natural selection. In fact,
The larvae of tiger beetles quinones are not at all uncommon in arthropods and, as they are non-
(Cicindellinae) live in palatable, they commonly have a defensive function. From these, hydro-
vertical tunnels in soil or
sand. quinones evolved as a superior line of defence; indeed, many arthropods
accumulate defensive compounds for later use. Predation is an immensely

strong selective pressure, and any small adaptation that improves survivabil-
ity will be passed on to future generations. Many insects, including beetles,
have a simple defensive system of secretory glands that fill a reservoir, the
end of which is closed by a valve so that the stored contents do not leak
to the outside before they are needed. Hydrogen peroxide might easily be-
come mixed with hydroquinones in any secreted defensive liquid because it
is a normal metabolic by-product and would increase the liquids’ predator-
deterring quality. A mixture of hydroquinones and hydrogen peroxide reacts
very slowly to produce quinines, which would further improve the defensive
function, but the reaction would be faster in the presence of enzymes. Cata-
lases and peroxidases, the enzymes involved, are commonplace and, were
they able to act on the secretions as they were expelled from the reservoir, they
could increase the output temperature of the liquid and produce even more
quinones. This is yet another small improvement to the system which could
easily have evolved by natural selection, and which would give those beetles
possessing it a significant adaptive advantage. The tube through which the se-
cretions leave the reservoir increases in size and gets more enzyme-secreting
capacity. The temperature of the defensive secretion goes up. The tube be-
comes a thick-walled reaction chamber and so on. Each little stage along the
way is advantageous, and thousands upon thousands of generations of bee-
tles, each one put to the ultimate test of leaving offspring before they get eaten,
will produce better and still better defences.
Another remarkable lineage of ground beetle, with about 600 species, is
the Paussinae—commonly known as the ant-nest beetles. Most of these bee-
tles look nothing like other carabids, as they are generally very robust, with
flattened antennae that may take various shapes—they can even be flat and
rounded, like elephant ears, or with horns, like deer antlers. As their name
suggests, they are obligate or facultative parasites of ant nests, where they
feed on ant larvae and even ant workers. Specialized glands on their body
surface of ant-nest beetles exude chemical compounds that imitate the ant
pheromones that allow the latter to communicate with each other. In this
way, ant-nest beetles are chemically invisible to their ant hosts and are not
attacked. Using stridulatory organs, ant-nest beetles are also able to speak
three different ‘languages’ by imitating the sounds produced by different ant
castes (workers, soldiers, queen). In this way, the ant-nest beetles are treated
as royalty by their hosts, being constantly taken care of, and gaining access to
any chamber of the ant nest.
Ninety per cent of all beetle families are placed in the suborder Polyphaga,
and some of these are extremely species rich. More than 63,000 species belong
to the family Staphylinidae. Rove beetles, as they are called, have a character-
istic, elongate, parallel-sided shape and very short wing cases. Most species
are small predators or scavengers, and live in a vast range of habitats, from
With its jaws held wide seaweeds to the fur of mammals. Being the dominant beetle lineages in most
open and abdomen raised, leaf litter habitats, rove beetles come in frequent contact with ants. This has
a staphylinid beetle displays
a typical threat posture. led numerous lineages, such as the Aleocharinae and many Pselaphinae, to
COLEOPTERA 183
evolve into obligate parasites of ant nests. Staphylinidae contain the largest
myrmecophilous number of myrmecophilous species than any other insect group.
Greek for ant-loving. Any Species of Nicrophorus, which belong to a small, closely related family of
species that lives with and
depends on ants for food, fewer than 300 species, the burying beetles (Silphidae) are well known for
protection, and care their habit of locating a carcass of an animal and burying it by digging out the
soil beneath. Other insects are highly attracted to carcasses of dead animals,
and the burying behaviour has evolved to prevent competition from fly mag-
gots. It has been shown that the cooperation of the two parent beetles leads
to greater breeding success. As in many species, single mothers who have to
compete for resources with conspecifics do worse than those with partners.
Usually, a monogamous pair will work together, but in cases where the car-
casses are big enough it might pay a male to try to attract a second mate. In
this way, he will father more offspring, but more brood means that the repro-
ductive success of the first female declines. Males in this position do try to
attract other females by emitting sexual odours, but their partners do all they
physically can to stop them from signalling. Sometimes, where there is a large
corpse and likelihood of intense competition from flies and other carrion in-
sects, it is advantageous for silphids to breed communally. There appears to be
an agreed truce between females who would normally compete, and, in these
examples, cooperative behaviour extends to females caring for each other’s
offspring. Take-overs are common at the height of the breeding season. Los-
ing pairs will be ejected from the carcass, and if any eggs have been laid, they
will be killed before the new female lays her own eggs. The guts of burying
beetles host symbiotic microbes, such as certain yeasts. When these microbes
are deposited on carrion, they prevent the growth of harmful microbes that
would otherwise lead to further putrefaction of the carcass and the build-up
of toxic substances. Using this approach, parent burying beetles are able to
maintain the carcass in a fresh state that provides a high-quality nutritional
resource for their larvae.
More than 30,000 species of scarab beetles, chafers, and related species
make up the Scarabaeidae, which includes some of the largest insects of all.
The family is divided into several important subfamilies: Scarabaeinae (dung
beetles or scarabs), Aphodiinae (small dung beetles), Cetoniinae (fruit and
flower chafers), Dynastinae (Hercules and rhinoceros beetles), Rutelinae (leaf
chafers), and Melolonthinae (leaf chafers). Adults and larvae feed on food-
stuffs ranging from dung and carrion to living or dead plant material, and
many herbivorous species are pests. Rhinoceros beetles (Oryctes rhinoceros),
for instance, can be a serious pest in coconut palm plantations. Insecticides
are too expensive and difficult to apply, but the problem has been solved by
the discovery of a suitable baculovirus biocontrol agent.
Despite enormous variation in shape, colour, and size between species, a
single character, the distinctive antennal club, unites these species. The club is
made up of three to seven flat, expanded, moveable plates, which can be closed
or opened out. Scarabaeid larvae are typical white grubs with well-developed
jaws, and many live in the soil and feed on roots.
Dung-burying beetles are extremely important recyclers in many parts
of the world and can clear vast quantities of dung in a relatively short time,
returning valuable nutrients to the soil; a job they have been doing for a long
time. Cretaceous trace fossils found have been interpreted as the remains of
herbivorous dinosaur dung with evidence of dung beetle workings. Species in
a dung beetle assemblage vary greatly in size and the manner in which they
utilize the resource. Several size classes of species roll balls of dung away using
their hind legs, bury it, and lay an egg inside the ball. The dung is used as a
food source by the larvae and the adults alike. Species in the genera Scarabeus
and Kheper are well-known dung rollers and can move balls as big as 50 mm
across. Others tunnel into and under the dung, some breed inside the dung
pad, and a few (called kleptocoprids) lay their eggs in the dung buried by
other species. Competition for dung can be intense and, in some species, the
females cling to the anal hairs of mammals and are thus in a good position
to deposit their eggs in fresh dung. In Australia, settler introduction of non-
native animals like cattle, horses, and sheep caused immense problems. The
Australian dung beetle community could not handle the vast quantities of
dung produced, which lay on the ground, degrading pastureland and provid-
ing a breeding ground for enormous numbers of flies. The introduction of 16
or so species of European and African dung beetles alleviated the problem.
The females of some species (Copris) stay with their developing brood un-
til they are ready to emerge as adults themselves. In these beetles, the parental
care has an important outcome for the survivorship of the young. In particu-
lar, balls deserted by females prior to pupation are likely to become infected
by species of fungi, which cover the outside and invade the interior.
Treatment of cattle with avermectins to control parasitic infections is
having a serious negative effect on dung beetles and other insects of dung
communities in many parts of the world. These pesticides are excreted in the
faeces and remain active for a long time. The result is the greatly delayed dis-
appearance of the dung beetle and unknown knock-on effects in associated
communities.
The scarab beetle, one of the most important religious symbols of ancient
Egypt, has been depicted in countless hieroglyphs, paintings, amulets, and
pieces of jewellery. Khepri (The Being), depicted as a man with a scarab as his
head, was the god of the morning sun.
The first scarab beetle to be worshipped by the ancient Egyptians was prob-
ably the large and brightly coloured species Kheper aegyptiorum. Initially, the
symbolism interpreted the movement and daily rebirth of the sun in terms of
the beetle rolling its ball. It has now been suggested that priests dug up the
dung balls, and probably observed the process of metamorphosis 5,000 years
before it was described by the French entomologist Jean-Henri Fabre. They
developed their ideas along the lines that if the sun and the scarab could ‘die’
(enter the ground), be transformed, and then be reborn, the same might be
possible for humans. It is conceivable that complex procedures of mummi-
fication represent an imitation of the dung beetles’ pupal stage, designed to
protect the body underground during its transformation and ultimate rebirth.
The 15,000 or so species of jewel beetles (Buprestidae) are typically inhabi-
tants of woodland and forest, particularly in tropical regions. The majority are
very attractive metallic green, blue, and red, with contrasting markings. The
COLEOPTERA 185
adults are tough, slightly flattened, and taper towards the rear of the body. The
larvae have a distinctive shape with a large, expanded prothorax and tapering
abdomen.
They are commonly called flat-head borers, and most species chew oval
cross-sectioned tunnels in the wood of dead or dying trees. Some species lay
their eggs exclusively in freshly burned wood, and the adults are attracted
from long range to forest fires. Species of the genus Melanophila have been
shown to have paired infra-red detectors very close to the coxae of their
middle legs. These incredible heat-sensing pit organs are each made up of
50–100 sensillae located at the bottom of a pit in the cuticle. The sensitivity
of the organs, which can respond to fires generating temperatures between
The expanded and flattened 425–1150◦ C, and their ventral location enables flying beetles to detect suit-
thorax of jewel beetle larvae able mating and egg-laying sites from a distance of several kilometres. A few
is a very characteristic
feature, and results in the buprestid species bore in the pith or roots of their host plants, and some are
tunnels they bore having an gall formers. The common name of the family refers to the fact that, in many
oval cross-section. parts of the world, the highly coloured and beautifully patterned wing cases
are used as decoration in embroidery and as jewellery.
The Cerambycidae is a large family with about 37,000 species worldwide.
Longhorn beetles, as they are often known, are typically elongate with long or
very long antennae.
The larvae of almost all cerambycids feed by boring into live or dead wood,
and several species can be forestry and timber pests. The larvae have symbi-
otic bacteria within special cells in the haemolymph that provide them with
essential nutrients lacking in their diet, and in many cases larval development
takes several years.
The leaf beetles (Chrysomelidae) are a large family consisting of more than
37,000 species. Leaf beetles tend to have a large, fairly smooth, rounded ap-
pearance, and many are very brightly coloured. All species are herbivorous,
the adults chewing leaves and flowers and the larvae feeding externally or
A typical longhorn beetle boring into stems, roots, and leaves. A great many species, particularly the
(Cerambycidae). flea beetles (subfamily: Alticincae), are damaging to crops. The notorious
Colorado Potato Beetle, Leptinotarsa decemlineata, which feeds on potatoes,
tomatoes, and aubergines, and spreads plant diseases, is a very serious pest.
Criocerus asparagi, the Asparagus Beetle, and Oulema melanopa, the Cereal
Leaf Beetle, are examples of two serious pest species that were introduced ac-
cidentally to North America. A few chrysomelids have their uses and have
been successful in biological control programmes against weeds. St John’s
Wort or Klamath Weed (Hypericum perforatum), brought to North America
by early European settlers, became a serious weed but was eventually con-
trolled by a European chrysomelid beetle. Related to the leaf beetles are the
Bruchidae, sometimes called pea and bean weevils. The adults of all species lay
their eggs on seeds, and the larvae burrow inside, feed, and pupate just below
A typical ladybird beetle
the surface. Many species such as Bruchus pisorum, Callosobruchus chinensis,
(Coccinellidae). Callosobruchus maculatus, and Acanthoscelides obtectus are serious pests of
stored peas, beans, and leguminous field crops.
Ladybirds or ladybug beetles (Coccinellidae) are favourite insects with
people of all ages. They have inspired poetry, become characters in folk
stories, and are symbols of health and happiness. Most of these small, brightly
coloured beetles are predacious on soft-bodied insects, such as aphids, mealy-
bugs, and scale insects, but there are some herbivorous species (Epilachna),
which can sometimes be crop pests. The predacious species can consume vast
quantities of prey during the course of their life cycle. Ladybird larvae are very
active with well-developed legs and seek out colonies of aphids on their host
plants using visual and olfactory cues.
When fully grown, the larvae pupate on foliage. Adult ladybirds show
a phenomenon known as reflex bleeding, whereby toxic fluids ooze out of
joints, especially the knees, and serve to deter would-be predators. Ladybirds
are very beneficial in controlling the natural populations of many pest insects,
and several species have been used as biological control agents. Rodolia car-
A typical ladybird larva.
dinalis, for instance, has been used in many parts of the world to control the
serious citrus pest, Cottony Cushion Scale (Margarodidae: Icerya purchasi).
However, one species, the Harlequin Ladybird (Harmonia axyridis), has be-
come a major invasive insect worldwide, replacing native ladybird species in
the areas it invades, and occasionally harming agriculture. This species is par-
ticularly harmful to viticulture, as large aggregations of ladybirds alter the
taste of wine.
The 2,200 or so small, light-brown to black beetle species that comprise
the Anobiidae feed on a wide range of animal and plant materials, includ-
ing wood, fungi, and dried organic foods like biscuits, bread, and foodstuffs.
A typical ladybird pupa.
Some anobiid species develop on stored, dry plant products, and can be pests
in stores, herbaria, and houses. Well-known cosmopolitan pest species in-
clude the Drugstore Beetle (Stegobium paniceum), the larvae of which feed
on a wide range of organic materials, Lasioderma serricorne, the cosmopoli-
tan Cigarette Beetle, which attacks stored tobacco, spices, and drugs, and
Anobium punctatum, the Furniture or Cabinet Beetle, whose larvae (wood-
worm) can destroy structural wood and wooden objects of all kinds. The
adult beetles normally emerge from pupa just below the surface of the wood
during the summer. Sometimes, larvae boring through wooden shelving will
The wood-boring larvae of
the Deathwatch Beetle can continue through books (bookworms). The gut of all these larvae contains
cause immense structural symbiotic yeasts which are responsible for breaking down the otherwise indi-
damage to timber.
gestible cellulose to sugars. The Deathwatch Beetle (Xestobium rufovillosum)
gets its name from the sexual signals that both sexes of this species generate
in late spring. They prefer hardwood timbers previously attacked by fungus,
and the adults, bracing themselves with their legs, tap their heads against
the substrate to attract mates. Males are able to orientate themselves to fe-
males by tapping and waiting for a reply before moving in the direction of
the vibrations. The origin of the common name derives from the fact that
the faint sounds can be heard in a quiet room or church where the dead are
laid out.
Of all the phytophagous beetles that have had a significant impact on
humans since agriculture first developed, the weevils (Curculionidae), with
more than 86,000 species worldwide, are the most economically important.
These beetles are characterized by the possession of a snout-like extension
A typical weevil of the head called the rostrum, which carries the jaws and the characteristi-
(Curculionidae). cally elbowed antennae.
COLEOPTERA 187
Sometimes, the rostrum can be as long as the rest of the body. Nearly
all species are herbivorous, and the larvae, depending on the species, can be
found eating every plant part from roots to seeds. Very many species are se-
rious pests, and cause great economic losses, some of which deserve special
mention. The Granary Weevil, Sitophilus granarius, has been recorded from
the time of the Pharaohs. A related species, the Rice Weevil, Sitophilus oryzae,
is also a serious pest of grains. Anthonomus grandis, the Cotton Boll Weevil,
is a scourge in cotton plantations. As for many crops, there is an increas-
The front view of a weevil ing demand for non-chemical pest control, and the use of parasitoids and
showing the elongate
rostrum and the elbowed
other natural enemies is meeting with some success. Massive releases of the
antennae. pteromalid wasp Catolaccus grandis, for example, have killed over 90% of Boll
Weevils in certain areas.
Bark beetles (Scolytidae), which are closely related to weevils, are the most
important pests of forests. Adult beetles bore into the cambium of trees just
under the bark, where they fed and lay eggs. The developing larvae tunnel
outwards from the egg chamber leaving characteristic tracks as they go.
The adults of some species (Xyleborus spp.) carry fungal spores, and in-
oculate the tunnels where the fungus grows, providing food. Bark beetles of
the genus Scolytus were the vectors of the Dutch elm disease fungus (Cer-
atostomella ulmi) that killed elm trees in many parts of Britain and North
America. Species of the genera Dendroctonus and Ips are widespread pests of
coniferous plantations across the northern hemisphere.
The darkling beetles (Tenebrionidae) are a large family of around 20,000
species whose larvae are associated with soil, rotting wood, fungi, and litter.
Some species, such as those of the genus Tenebrio, can be pests in grain, and
The patterns made by species of Tribolium are pests of flour, cereals, dried fruit, and meal.
developing bark beetle
larvae as they tunnel away In the Namib desert, some tenebrionid species climb to the tops of dunes
from the egg chamber. in the morning and dip their heads into the moist prevailing wind. Tiny
droplets of water condense on their elytra, which are specially sculptured with
hydrophobic grooves and hydrophilic bumps. These droplets coalesce to pro-
duce larger droplets that run forward. A bed of hairs at the junction of the
prothorax and the head channels the water from the dorsal surface to the ven-
tral surface and thus towards the mouth. This, one among many adaptations
for survival, is why beetles are the most abundant and successful of all insects.
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Stork, N. E., McBroom, J., Gely, C., and Hamilton, A. J. (2015). New approaches narrow
global species estimates for beetles, insects, and terrestrial arthropods. Proceedings of the
National Academy of Sciences of the United States of America 112: 7519–23.
Strong, L. (1992). Avermectins: a review of their impact on insects of cattle dung. Bulletin
of Entomological Research 82: 265–74.
Vondran, T., Apel, K.-H., and Schitmz, H. (1995). The infrared receptor of Melanophila
acuminata De Geer (Coleoptera: Buprestidae): ultrastructural study of a unique insect
thermoreceptor and its possible decent from a hair mechanoreceptor. Tissue and Cell 27:
645–58.
189
Strepsiptera
(strepsipterans)
Common name Strepsipterans, Metamorphosis Complete (egg, larva,

twisted-wing insects pupa, adult)
Derivation Gk. Streptos–twisted; Distribution Worldwide
pteron–a wing Number of families 10
Size Body length 0.4–35 mm Known world species About 630 (0.056%)
(usually under 6 mm)
Branched antenna
Small strap-like
Berry-like eye
front wings
Little
obvious
vention
Fan-shaped
hind wing
Stylops melittae (male) — Strepsiptera
Key features
• endoparasites of insects
• rarely seen
• sexes extremely sexually dimorphic
A male Stylops ater mating

with a female that is
parasitising an Andrena vaga
© John Smit
bee (female is the yellow
spot at the end of the bee’s
abdomen). Netherlands.
The first members of this strange order that were described belonged to the
family Stylopidae, and, for this reason, the common name ‘stylops’ is often
used to refer to any strepsipteran species, and the insects they parasitize are
described as having been ‘stylopized’. However, the common name twisted-
wing insects is becoming more widespread in recent years.
Strepsipterans are highly specialized endoparasites of other insects in at
least seven orders and 34 insect families belonging to the orders Thysanura,
Blattodea, Mantodea, Orthoptera, Hemiptera, Diptera, and Hymenoptera.
There has even been a report of a microcaddisfly being used as a host. De-
spite this broad host range, the insects most commonly stylopized are bugs,
wasps, and bees. Males and females may parasitize different insect hosts.
The order is divided into two suborders—the Mengenillidia (one small
family, the Mengenillidae) and the Stylopidia (eight families)—as well as a
family of uncertain subordinal placement, the Bahiaxenidae. In the Menge-
nillidae, the fully grown male and female larvae leave the hosts, which are
species of silverfish (Lepismatidae), and pupation takes places outside. In
this family, both adult males and females are free-living, a unique condition
STREPSIPTERA 191
© John Smit
A male Stylops ater at rest.
Netherlands.
neoteny among the Strepsiptera. The males are winged, but the females are neotenous,
the retention of immature
grub-like, and wingless, although they possess legs and antennae.
characteristics in the adult
stage The females of all other Strepsiptera are totally endoparasitic, as they never
leave the confines of their host’s body, and they are surrounded by the cuticle
of their own pupal stage. The females do not move and have no eyes, anten-
nae, mouthparts, legs, wings, nor external genitalia. They do not go through a
pupal stage, and remain as legless larviform adults, their body almost entirely
filled with eggs. The only bit of them that is visible and protrudes through
the body wall of the host is the relatively featureless, chitinized cephalotho-
rax. Adult males, on the other hand, are dark coloured, free-living, winged, do
not feed, and are short lived (typically for just a few hours). Their heads have
prominent, raspberry-like eyes and strange, branched antennae, and the tho-
rax bears a pair of small strap-like front wings and much larger, fan-shaped
hind wings with little obvious venation. The legs of some males lack claws.
Interestingly, the front wings of male strepsipterans, which are reduced to
flattened stalk-like or strap-like structures, have been shown to function in
a manner analogous to dipteran halteres. As the front wings oscillate rapidly
in flight, any rotational movement of the insect will induce inertial (Coriolis)
forces in the wings, perpendicular to the plane of oscillation. These forces are
thought to be sensed by two fields of campaniform sensillae at the bases of
the front wings, and the nerves from these run to the thoracic ganglia. Me-
chanical stimulation of the front wings produces compensatory movements
of the head and abdomen.
Males are able to detect and respond to sexual pheromones emitted by

virgin females and, after locating them, the male clings to the body of the
host and mates with her. The precise way mating is achieved in Strepsiptera
is contentious, but the few known examples are fascinating. In Mengenill-
idae, the male uses his sharp penis to pierce the female in any body part
other than the head, and ejaculates directly in her blood stream (haemo-
coel), where fertilization takes place. At least in some Stylopidia, mating also
takes place via traumatic insemination, but the male pierces a specific invagi-
nation of the female cephalothorax to inject his semen. This invagination
is specifically adapted for this purpose, and likely serves to protect the fe-
male from the damage caused by being pierced by the male’s penis. Other
Stylopidia may mate somewhat more conventionally by pushing their penis
through an opening in the pupal skin that surrounds the female, although
this requires further confirmation. The production of the male-attracting
pheromone ceases when copulation takes place, and in some species the males
are able to detect whether a female has mated previously with more than
one partner. One species avoids the hassle of being marauded by males by
following a parthenogenic approach, where only females are born. The num-
ber of eggs a female strepsipteran produces varies from about a thousand to
many hundreds of thousands, depending on her size, which has been deter-
mined by the size of the host insect inside which she has developed. In one
Mobile first instar larva species, a single egg may give rise to more than one embryo by subdividing
(triungulin) of a stylopid.
(polyembryony). The first instar larvae develop within the body of the female
and emerge via the brood canal opening around the cephalothorax. Females
may give birth to many thousands of tiny, six-legged, first instar larvae called
primary larvae (also known by certain authors as triungulins or planidia),
whose main job is to find immature stages of the appropriate host insect
species.
If their hosts are exopterygote insects, such as grasshoppers or bugs, the
Normal male
primary larvae are shed as the host moves about. In the case of species that
parasitize social Hymenoptera, such as wasps and bees, the primary larvae
emerge from the brood canal of their mother when the host visits a flower.
Stylopized male Here they will sit, ready to grab hold of a suitable adult who will unwittingly
transport them back to the nest, where they will find the right sort of eggs
or larvae inside which to develop. The primary larvae produce enzymes that
Normal female soften the cuticle of the host and, once inside, they moult immediately to a
legless endoparasite. After five or more grub-like instars, the fully grown lar-
vae pupate, with one end of the pupa sticking out from the body wall of the
host. If the pupa contains a male, the cephalothorax end of the pupal cuticle
Stylopized female
is pushed off and the male flies away, whereas females remain in situ. Hosts of
Reversal of secondary males usually die when the strepsipteran emerges but hosts of females live on
sexual characteristics in a
bee (Andrena chrysoceles) until she has mated and given birth. In all cases, hosts suffer what is known
parasitized by a as parasitic castration and are unable to reproduce. However, stylopized in-
strepsipteran.
sects live longer than usual, and have larger fat stores as well, which allow
their strepsipteran parasites to complete their life cycle and have abundant
resources throughout their development. Furthermore, stylopized social in-
sects like wasps display altered gene expression patterns in their brain, which
STREPSIPTERA 193
suggests that Strepsiptera are able to manipulate the behaviour of their host
for their own benefit.
The type of host and the position in which the female strepsipteran pupates
is characteristic of particular families. Members of the Elenchidae are often
found on grasshoppers; Halticophagidae on treehoppers, leafhoppers, spittle
bugs, and mole crickets; and Stylopidae on bees (Adrenidae and Halictidae)
and wasps (Sphecidae and Vespidae).
Female in
Stylops sp. (to same scale) abdomen of bee Male
In most known life cycles, both males and females of any strepsipteran
species parasitize the same host species. However, there are some exceptions,
where males and female utilize very different hosts. The family Myrmeco-
lacidae contains a little over 120 species, mostly known from the collection
of male specimens. The females of only a handful of species have been found,
and, in the few cases where males and females have been linked, they have
been found in completely different hosts; the females attacking mantids,
grasshoppers, or crickets, and the males using ants as their hosts. The differ-
ences in size, both of the sexes and their respective hosts, is remarkable. Male
myrmecolacids are less than 1.5 mm in length, whereas the females are much
larger, some species, ranging in length from 20–35 mm, being the largest
strepsipterans in the world. The females are endoparasitic in several species of
phytophagous cricket, belonging to the genera Segestidea and Segetes that can
cause damage to oil and coconut palms in Papua New Guinea. The introduc-
tion of a strepsipteran, Stichotrema dallatorreanum, to islands and areas where
it is absent may be an effective means of biologically controlling pest crickets
in plantations. One of the effects of stylopization in female crickets concerns
the host’s eggs. More than half of the eggs from parasitized females are darker
and misshaped compared to normal eggs. In addition, electron microscopic
examination of the eggs reveals that their shells lack the fine structures neces-
sary for normal gaseous exchange. In this way, the populations of these pest
crickets are greatly reduced.
One species that has a significant role in pest control is Elenchus japonicus
(Elenchidae). This species is parasitic in two important pests of rice in Asia,
the White-backed Planthopper, Sogatella furcifera, and the notorious Brown
Planthopper, Nilaparvata lugens. The planthoppers are obviously killed when
males emerge, but the effects of stylopization on the host are greater because
the genitalia of the bugs are affected, or sometimes even lacking. The plan-
thoppers, therefore, cannot reproduce, but are still able to disperse to new
areas, further spreading the parasite. The species Stichotrema dallatorreanum
is also used as a pest control agent in Papua New Guinea against bush crickets,
as they are parthenogenic and can be readily mass reared.
Due to the highly modified morphology of both sexes of Strepsiptera, the
relationships of this order had been highly controversial ever since its dis-
covery. Due to morphological characteristics these insects share with some
families of beetles, such as the parasitic Rhipiphoridae, the Strepsiptera have
sometimes been regarded as a superfamily of the Coleoptera. Early molecular
sequence data instead indicated a close relationship with the Diptera. How-
ever, detailed morphological and phylogenomic studies have conclusively
demonstrated that Strepsiptera are the sister group to Coleoptera, and any
similarities with the Rhipiphoridae arose independently, due to their similar
parasitic lifestyle.
The fossil record of Strepsiptera is relatively scarce, with the oldest fos-
sils of either adult males or primary larvae dating to the Cretaceous. This
shows that this group has been following a remarkably constant parasitic way
of life for at least 100 million years. Molecular estimates suggest that Strep-
siptera arose at least 120 mya, although the ultimate origin of the group may
go back to the Permian, when their lineage split from the common ancestor of
beetles.
Key reading
Beani, L., Dallai, R., Cappa, F., Manfredini, F., Zaccaroni, M., Lorenzi, M. C., and Mercati, D.
(2021). A Strepsipteran parasite extends the lifespan of workers in a social wasp. Scientific
Reports 11: 7235.
Geffre, A. C., Liu, R., Manfredini, F., Beani, L., Kathirithamby, J., Grozinger C. M., and Toth,
A. L. (2017). Transcriptomics of an extended phenotype: parasite manipulation of wasp
social behaviour shifts expression of caste-related genes. Proceedings of the Royal Society
B 284: 20170029.
Kathirithamby, J. and Hamilton, W. D. (1992). More covert sex: the elusive females of the
Myrmecolacidae. Trends in Ecology and Evolution 7: 349–51.
Kathirithamby, J. (2009). Host-parasitoid associations in Strepsiptera. Annual Review of
Kathirithamby, J. (2018). Biodiversity of Strepsiptera. In: Insect biodiversity: science and so-
ciety, volume II. Foottit, R. G. and Adler, P. H. (eds), pp. 673–703. John Wiley & Sons
Ltd., Hoboken.
Kinzelbach, R. (1990). The systematic position of the Strepsiptera (Insecta) American
Entomologist 36: 292–303.
Kinzelbach, R. and Pohl, H. (1994). The fossil Strepsiptera (Insecta: Strepsiptera). Annals of
the Entomological Society of America 87: 59–70.
Misof, B., Liu, S., Meusemann, K., Peters, R. S., Donath, A., Mayer, C., Frandsen, P. B., et al.
7639–7767.
Niehuis, O., Hartig, G., Grath, S., Pohl, H., Lehmann, J., Tafer, H., Donath, A., Krauss, V.,
Eisenhardt, C., Hertel, J., Petersen, M., Mayer, C., Meusemann, K., Peters, R. S., Stadler, P.
F., Beutel, R. G., Bornberg-Bauer, E., McKenna D. D., and Misof, B. (2013). Genomic and
morphological evidence converge to resolve the enigma of Strepsiptera. Current Biology
23: 1388.
Peinert, M., Wipfler, B., Jetschke, G., Kleinteich, T., Gorb, S. N., Beutel, R. G., and Polh, H.
(2016). Traumatic insemination and female counter-adaptation in Strepsiptera (Insecta).
Scientific Reports 6: 25052.
Pix, W., Nalbach, G., and Zeil, J. (1993). Strepsipteran forewings are haltere-like organs of
equilibrium. Naturwissenschaften 80: 371–4.
STREPSIPTERA 195
Pohl, H., Batelka, J., Prokop, J., Müller, P., Yavorskaya, M. I., and Beutel, R. G. (2018). A nee-
dle in a haystack: Mesozoic origin of parasitism in Strepsiptera revealed by first definite
Cretaceous primary larva (Insecta). PeerJ 6: e5943.
Solulu, T. M., Simpson, S. J., and Kathirithamby J. (1998). Effects of strepsipteran para-
sitism on a tettigoniid pest of oil palm in Papua New Guinea. Physiological Entomology
38: 388–98.
196
Antliophora
This superorder comprises scorpionflies (Mecoptera), fleas (Siphonaptera),
and true flies (Diptera). While scorpionflies are modestly diverse, flies are
among the richest insect orders, with at least 158,000 species.
197
Mecoptera
(scorpionflies)
Common name Scorpionflies Metamorphosis Complete (egg, larva,

Derivation Gk. Mekos–long; pteron–a pupa, adult)
Known world species About 750 (0.066%)
Head prolonged
forming a rostrum
Thread-like antenna
Ocelli Large compound eye
Front and hind

wings of similar
size and venation
Slender legs
Elongate body
Bulbous genitalia (male only)
Panorpa germanica — Mecoptera
Key features
• distinctive elongated face
• abdominal tip of males often resembling a scorpion sting
• mostly in damp wooded areas
A male scorpionfly browsing

on pollen. The scorpion-like
abdominal tip of many
mecopterans gives them
© Anne Riley
their common name.

Panorpa communis,
Hungary.
The common name of the order is derived from the occurrence in the males of
some species of swollen genitalia, which are positioned at the end of a slender
abdomen and held curved upwards in a scorpion-like fashion.
Many features of these insects, such as the two pairs of equally sized
and richly veined membranous wings, render them superficially similar to
neuropterans, but, in fact, they are most closely related to the Diptera and
Siphonaptera. Some mecopteran families (Choristidae, Nannochorsitidae,
Notiothaumidae, and Meropeidae) comprising only dozen or so primitive
species are found only in the southern hemisphere. The Bittacidae (hang-
ingflies), a larger family, is also confined to southern continents, while the
Panorpidae (common scorpionflies) and the Boreidae (snow scorpionflies or
snow fleas) occur only in the northern hemisphere. With the exception of
boreids, scorpionflies are generally found in damp, wooded, or shady areas
MECOPTERA 199
with an abundance of vegetation. The body is generally elongate and cylin-

drical, and the large, narrow wings may be clear or marked with spots or
bands of dark colour. Some species are short-winged or completely wingless.
The head has large compound eyes, three ocelli, and is characteristically ex-
tended downwards in the form of a beak or rostrum, at the end of which are
biting mouthparts. The antennae, which are thread-like and have as many
as 60 segments, are typically longer than half the length of the body. The
Female legs are long and slender in most species, but, in the hangingflies (Bittaci-
dae), they are very long indeed and used for prey capture. Bittacids, which
resemble crane flies (Tipulidae), hang from vegetation, holding on with their
front legs, or fly, trailing their slender, hind legs in the air. The fifth tarsal
segments of the hind legs are enlarged and raptorial to seize small insects.
The majority of other scorpionflies, typified by the Panorpidae, feed on dead
or dying insects, and will also consume carrion, nectar, sap flows, and fruit
juice. There may be intense interspecific competition for food, and it is the
larger species which get the biggest share. An unusual feeding strategy, seen
Male in panorpids, is the robbing of prey from orb webs. These scorpionflies are
Details of the abdominal
able to walk over webs to feed on insects that have been trapped, or even feed
tips of male and female on silk-wrapped insects that the spider has stored. Despite being careful, they
scorpionflies showing do sometimes get trapped, but are able to free themselves using salivary se-
external genitalia.
cretions. Snow scorpionflies (Boreidae) feed on mosses, and members of the
family Panorpodidae are herbivorous. Unusually, the larvae of the southern
hemisphere family Nannochoristidae are aquatic and live around the margins
of lakes and in swampy areas, where they prey on larval non-biting midges
(Chironomidae).
Prior to mating, which often takes place after dark, Mecoptera frequently
engage in complex courtship rituals that involve slow wing movements that
display species-specific wing pigmentation patterns, as well as rhythmic ab-
dominal vibrations, which generate low-frequency vibrational songs. Mating
often involves pheromones and the offer of nuptial gifts. In some species, sex-
ual pheromones only attract conspecific females, while in others, both males
and females are attracted. Mating behaviour has been studied in depth in a few
species, mostly in the genus Panorpa. Males use a variety of courtship tech-
niques, of which the commonest is the offering of gifts, which may take the
form of a dead insect, or a mass of saliva produced by the males which is stuck
to the surface of a leaf. The male then emits his sexual pheromone to attract
a mate to his offering. In scorpionflies, size is everything, and females will
quickly reject males whose gifts are too small. In the case of salivary masses,
the size of the gift is a good indication of how well the male has fed prior to
mating. As would be expected, females prefer males who provide large nutri-
tious gifts and, as a result, they are able to lay more eggs. Having accepted the
gift, the female will feed on it during the process of copulation, which may last
more than an hour. Competition for females, and the gifts to offer them, can
be fierce. Males who do not find good enough gifts may take females by force,
and after grabbing a potential mate using their genital claspers, they employ
the notal organ (a clamp-like structure located on the dorsal surface of the
abdomen) to restrain one of the female’s front wings during mating. The no-
tal organ is also used during normal mating. Males will often try to wrestle
rival males from females while they are copulating. Females may mate with
several males during their adult life. In hangingflies (Harpobittacus and Bitta-
cus), the picture is further complicated by males stealing each other’s nuptial
gifts, either aggressively or by stealth. In the latter case, a male will imitate the
behaviour of a responsive female, and before the duped male discovers his
mistake, they seize the nuptial gift for their own use. In some species, part of
the male genitalia takes the shape of a long, coiled ‘penisfillum’, and it is likely
this structure might be used to remove sperm from previous matings.
Eggs are laid in small groups in the soil. The larvae of most species re-
semble small caterpillars or beetle grubs. The larvae usually have compound
eyes (not a collection of simple eyes), which vary in the number of om-
matidia from fewer than 10 to more than 30, but some species are eyeless.
There are three pairs of thoracic legs and a number of abdominal prolegs
in those larvae resembling caterpillars (Panorpidae, Bittacidae, and Cho-
ristidae). The soil surface-dwelling larvae of some hangingflies are able to
camouflage themselves by eating soil and then covering themselves with
the excrement produced. After three or four moults, pupation takes place
in an underground cell, or among damp soil or vegetation. The pupae can
move their jaws, and their legs and antennae are not stuck down to the body
surface.
Snow scorpionflies (Boreidae) are dark brown or black in colour and can
be seen in autumn and winter on the surface of snow, among mosses, or under
stones, particularly in cold or mountainous regions. These insects are rarely
encountered, but, when present, are noticeable on account of their colour and
rapid walking. Males use their reduced, hook-like wings to hold and carry
the females during mating. The eggs are laid in moss, and the larvae, which
have well-developed thoracic legs, feed on moss and other plant matter. Su-
perbly adapted for life in cold conditions, these insects contain antifreeze
compounds and can tolerate temperatures down to −6◦ C, and, when walking
on the surface of snow, the body of species such as Boreus brumalis absorbs
radiation from the sun. When threatened, snow scorpionflies use a catapult-
like mechanism on their mid- and hind legs to make long-distance jumps that
allow them to escape from potential predators.
Scorpionflies are a particularly old order, with fossils dating to the Early
Permian (about 290 mya). It appears that extinct mecopterans were up to
three times more diverse than their living relatives, although this could be
artefactual, as many of these fossils might represent early Antliophora and not
true Mecoptera. The relationships of scorpionflies are debated, especially with
regards to fleas (Siphonaptera), which may represent their sister group, or a
lineage within Mecoptera (discussed in more detail in Siphonaptera (fleas)).
What is certain, is that flies (Diptera) are the sister group to Mecoptera and
Siphonaptera.
The economic importance of Mecoptera is negligible, although they might
have a useful ecological role as recyclers in many ecosystems.
MECOPTERA 201
Key reading
Burrows, M. (2011). Jumping mechanisms and performance of snow fleas (Mecoptera,
Boreidae). Journal of Experimental Biology 214: 2362–74.
Byers, G. W. and Thornhill R. (1983). Biology of the Mecoptera. Annual Review of
Courtin, G. M., Shorthouse, J. D., and West, R. J. (1984). Energy relations of the snow
scorpionfly Boreus brumalis (Mecoptera) on the surface of snow. Oikos 43: 241–5.
Hartbauer, M., Gepp, J., Hinteregger, K., and Koblmüller, S. (2015). Diversity of wing pat-
terns and abdomen-generated substrate sounds in 3 European scorpionfly species. Insect
Science 22: 521–31.
Penny, N. D. (1975). Evolution of the extant Mecoptera. Journal of the Kansas Entomological
Society 48: 331–50.
Ren, D. and Shih, C. K. (2005). The first discovery of fossil eomeropids from China (Insecta,
Mecoptera). Acta Zootaxonomica Sinica 30: 275–80.
Thornhill, R. (1980). Rape in Panorpa scorpionflies and a general rape hypothesis. Animal
Thornhill, R. (1981). Panorpa (Mecoptera: Panorpidae) scorpionflies: systems for under-
standing resource-defense polygyny and alternative male reproductive efforts. Annual
Review of Ecology and Systematics 12: 355–86.
Thornhill, R. and Sauer, K. P. (1991). The notal organ of the scorpionfly (Panorpa vulgaris):
an adaptation to coerce mating duration. Behavioural Ecology 2: 156–64.
Willman, R. (1987). The phylogenetic system of the Mecoptera. Systematic Entomology 12:
519–24.
Zhong, W. and Hua, B. (2013). Mating behaviour and copulatory mechanism in the
scorpionfly Neopanorpa longiprocessa (Mecoptera: Panorpidae). PloS ONE 8: e74781.
202
Siphonaptera
(fleas)
Common name Fleas Metamorphosis Complete (egg, larva,

Derivation Gk. Siphon–pipe, tube; pupa, adult)
a+pteron–wingless Distribution Worldwide
Mostly under 5 mm Known world species About 2,190 (0.19%)
Backward Tough, laterally flattened body

facing bristles
Pronotal comb
Simple lateral eye Sensillum
Genal
comb
Maxillary
palp
Mouthparts
Enlarged hind leg

Ctenocephalides felis — Siphonaptera
SIPHONAPTERA 203
Key features
• small, wingless ectoparasites on mammals and some birds
• blood feeders
• characteristic jumping ability
• many species are vectors of disease
© Martin Gore
Cat fleas do not live
permanently on humans but
will take blood meals.
Ctenocephalides felis, UK.
While feeding, fleas excrete

© Martin Gore
droplets of digested blood.

These are among the
principal food items for flea
larvae. UK.
Found wherever there are suitable hosts, even in Arctic and Antarctic re-
gions, fleas are a distinctive and readily recognizable group. The vast majority
of flea species are parasitic on land mammals (well over 90%), the remain-
der being attached to bird species. Aquatic mammals are flea-free, as are
roaming or thick-skinned mammals, such as elephants and rhinoceroses. As
holometabolous insects, the larvae are very different from the adults, and have
a different feeding ecology. Fleas have evolved as parasites of animals that reg-
ularly use lairs, burrows, dens, or nests, because it is here that the larvae feed
and develop. There are, however, some strange and fascinating exceptions
to this rule such as the Arctic Hare Flea, Euhoplopsyllus glacialis (Pulicidae),
where the larvae live in the host’s fur, and the Tasmanian Devil Flea (although
it is found also in quolls and the extinct thylacine), Uropsylla tasmanica (Py-
giopsyllidae), where the eggs are stuck to the hair of the host, and the larvae
actually develop inside the host’s skin. As adults, these blood-sucking insects
are wingless and highly modified for an ectoparasitic life on the bodies of
their hosts. The tough, shiny, brownish or reddish body is streamlined and
flattened from side to side with a variety of backward-pointing spines and
bristles. Comb-like structures on the cheeks on the posterior edge of the
pronotum of many species, genal and pronotal ctenidia, respectively, are very
useful identification features. It is thought that ctenidia prevent the flea from
being pushed backwards and help it stay on the host’s fur.
Fleas living on spiny hosts, such as porcupines and hedgehogs, have fewer,
stouter, and more widely spaced comb teeth than those living on furry mam-
mals, as their hosts cannot effectively scratch themselves. The head typically
has very short, three-segmented antennae, which fit into special grooves, and
short, robust mouthparts for piercing skin and sucking blood. A spring-like
system allows the stylets to be rapidly and repeatedly driven through the skin
to reach the blood vessels just below the surface. Fleas may have a pair of
simple, lateral eyes similar to ocelli, and are negatively phototactic, but many
Head of the Dog Flea
(Ctenocephalides canis). nest-living species are blind. The legs are stout, and the enlarged hind legs are
part of the flea’s unique and powerful jumping mechanism. Before the flea
jumps, the hind legs are flexed and rotated by the trochanter-levator muscles
past a pivot point into a cocked position. In this position, a block of highly
elastic rubber-like protein called resilin, located between the pleuron and the
notum, is compressed and acts as an energy store. The jump is started by con-
traction of the trochanter-depressor muscles, which bring the legs back again,
past the pivot point. At this stage, the energy from the resilin block is released
and, together with the force generated from the contracting jumping mus-
cles (notum-trochanter muscle), the legs are rotated rapidly and down against
the substrate. Many fleas can jump considerable distances—the Cat Flea can
achieve a high jump of over 30 cm (12 in) at up to 130 times the acceleration
of gravity. Hungry fleas may jump hundreds of times an hour for several days
to find a host.
Head of the Human Flea
(Pulex irritans) lacking genal The dorsal surface of the last visible abdominal segment bears a con-
and pronotal ctenidia. spicuous sensillum, which can be flat or rounded in outline. This plate-like
structure is covered with a variable number of sensory hairs arising from
dome-like bases. Although it certainly has a sensory role, it is hard to believe
that the exact function of such a large structure has remained unknown.
Unlike lice, which live permanently on their hosts, most fleas do not live
on their host all the time. As adults, many species feed intermittently, and
respond to the presence of an animal by sensing movement, temperature,
humidity, carbon dioxide, and odours. Some species, collectively known as
SIPHONAPTERA 205
nest fleas (as compared with fur fleas which spend most of their time on the
host), only move onto a host to feed and spend the rest of their time in the
lair or burrow. Fleas regularly move from host to host, and in some species,
can survive for well over a year without a blood meal. The life cycle is rela-
tively simple in most species. The eggs are dropped from the fur or laid in
the host’s burrow, lair, or nest. The blind, pale, and worm-like larvae feed
on a range of organic materials, which include dried blood and the excre-
ment of the adult fleas, which they sometimes take directly from the anus.
Incredibly, the larva of one species, The European Rat Flea (Nosopsyllus fas-
ciatus), is known to beg for food by biting an adult on one of its posterior
bristles. This action stimulates the flea to release a droplet of liquid excre-
ment. Flea larvae do not have any thoracic legs or abdominal prolegs and
move around by wriggling. There are three larval instars, and pupation takes
place within a silken cocoon, which typically has bits of debris and small par-
ticles woven into its structure. Adult fleas will not emerge immediately once
pupation is over; they will usually remain dormant inside until vibrations
or other cues alert them to the presence of a host nearby. Workers clearing
empty houses or new owners moving into property where pets have lived pre-
viously can be attacked by thousands of starving fleas that have matured in
the absence of hosts. The life cycles of some fleas can be dependent on the
development of their hosts, such as fleas whose hosts return annually to nest-
ing sites. In some cases, the levels of hormones associated with the pregnancy
of a host animal will stimulate the flea to reproduce in anticipation of new
hosts.
Ceratophyllidae is the biggest of the 19 flea families, comprising about 20%
of all flea species. More than 80% of the 455 species in this family are asso-
ciated with small rodents, and the remainder with birds. Insectivorous bats
are hosts to about 127 species that belong to the Ischnopysllidae. The species
belonging to each genus of bat flea are confined to host species within a single
bat family. Most bat fleas are found on cave-dwelling bats, where the environ-
ment provides a living for the larvae. Some specialized species in the family
Malacopsyllidae use armadillos as hosts.
About 165 species make up the Pulicidae (common fleas), which, although
a relatively small family, contains many species of medical or veterinary
importance (see Box). This family includes species parasitic on dogs, cats,
humans, hedgehogs, and porcupines, as well as many other carnivores and
rodents.
Some pulicids are cosmopolitan in distribution, and among the best-
known species are the Cat Flea (Ctenocephalides felis) (also a worldwide pest
of dogs), the Dog Flea (Ctenocephalides canis), and the Human Flea (Pulex ir-
ritans). Apart from being very itchy and irritating, the bites of fleas can cause
allergic reactions and transmit a variety of diseases and parasitic worms. Pet
owners may only find a few dozen adult fleas on the bodies of their animals,
but many tens of thousands of eggs, larvae, and pupae will be present in the
home.
Fleas can act as vectors of a number of bacterial, rickettsial, and viral dis-
eases of animals and humans (see Box). The best-known disease is Black
Death (or bubonic plague) caused by the bacterium Yersinia pestis, and is ac-
tually primarily a disease of rodents, which can be passed to humans when
the fleas’ hosts die, and the fleas are forced to seek blood from other sources.
A flea feeding on an infected brown or black rat (or other rodent) might pick
up the bacteria in a blood meal. These bacteria multiply in the midgut of the
flea, and, in some cases, may completely block the tract. When such a flea
attempts to feed on the next host—rodent or human—the powerful pharyn-
geal pump tries to work but cannot pass the blood meal into the to gut. When
it stops trying to feed, pressure and elastic forces cause the blood in the oe-
sophagus, which is now mixed with some of the plague bacterial culture, to
be pumped back into the host. In this way, a blocked flea may infect many
hosts before it dies of starvation. There have been three great world outbreaks
or pandemics of plague. The first is believed to have originated from Central
Asia and spread through North Africa to Europe during ad 550–600. The
second pandemic in the Middle Ages is thought to have caused 75 million
human deaths throughout Europe and Asia. Outbreaks of the third plague
pandemic in parts of India, Asia, Africa, and South America took place in
the twentieth century, and plague can still occur today under certain condi-
tions. The disease and its effects have been considerably reduced by the use of
pesticides and modern drugs. Flea species in 50 genera across several fami-
lies are vectors of plague in many parts of the world. Seventeen species in the
genus Xenopsylla (Pulicidae) are disease vectors, the best-known of which is
the cosmopolitan Rat or Plague Flea (Xenopsylla cheopis).
Stick-tight fleas belonging to the genus Echidnophaga (Pulicidae) are
unusual in that they remain firmly attached to the skin of their host by
their needle-like mouthparts. About 23 species are known, of which, one
(Echidnophaga gallinaceus) is widespread and a serious pest of poultry. Even
more unusual than stick-tight fleas are 13 species of sand-fleas, jiggers, or
chigoes belonging to the Tungidae. Originally, an American species, Tunga
penetrans is believed to have been spread to tropical Africa by the Atlantic
slave trade. Another tungid species, common on rats, occurs in the eastern
parts of China. Walking about in bare feet in dry, sandy areas of tropical
Africa and Central and South America is not generally a good idea as, quite
apart from the dangers of sharps and thorns, Tunga penetrans is a significant
human parasite. The tiny females burrow into the skin of the foot, often under
toenails, and remain encysted there, whereas males are free living. Gaseous
exchange, excretion, mating, and egg-laying all take place through a small
aperture at the rear of the embedded female. After fertilization, the abdomen
swells enormously as the eggs develop, and the legs degenerate. Jiggers can
grow to the size of pea causing pain, intense itching, and inflammation. In
some cases, infection of the skin lesions can be serious and severe ulceration
can lead to the loss of toes and even death through blood poisoning or tetanus
infection.
SIPHONAPTERA 207
Some fleas of medical or veterinary importance
Family Species Medical or veterinary importance

Pulicidae Xenopsylla cheopis Widespread vector of plague and murine
or endemic typhus in rats and humans
·········································································································································································
Ctenocephalides felis Cat Flea
·········································································································································································
Ctenocephalides canis Dog Flea. Intermediate host for Dipylidium
caninum, a tapeworm of cats, dogs and,
sometimes, humans
·········································································································································································
Pulex irritans Human Flea*
·········································································································································································
Tungidae Tunga penetrans Jiggers, chigoes, sand fleas. Cause tungiasis
in humans
·········································································································································································
Ceratophyllidae Ceratophyllus gallinae European Chicken Flea
* Although called the Human Flea, this species is also commonly found on goats, pigs, and some other
animals such as foxes and badgers. Early humans would have quickly acquired this flea from another
species, perhaps a previous occupant of a cave or shelter. Pulex irritans can act as a vector of bubonic
plague and has also been implicated in the transmission of a number of other human diseases. Fleas
have been implicated in the transmission of tularaemia and many other diseases than those already
mentioned.
Host specificity in fleas is less strong than that of lice, and many species will
readily feed on a range of related, or even unrelated, hosts. Some flea species
may be found on more than 30 host species, and some unfortunate animals
are host to more than 20 different flea species. The main reason for this is that,
being holometabolous insects, where the larval stages have a totally different
ecology, it is the nest or lair where the larvae develop, rather than the host,
that is important to the flea’s survival. Even though the host range may be
broad, in some cases, where a flea is forced to take blood meals from a species
that is not normally used as a host, they may lay fewer eggs. After killing and
eating a rabbit, the ears of a domestic cat may become encrusted with dozens
of feeding rabbit fleas, but even if they laid eggs, it is unlikely that the cat’s
bedding would provide as suitable a larval habitat as a rabbit burrow.
Insecticidal control of adults combined with treatment of the resting place
and bedding of domestic animals is common, but the use of growth regula-
tors, such as lufenuron, in feedstuffs is very effective. Adult fleas take up the
substance from the blood and it is transferred to the eggs where it interferes
with the formation of chitinous structures in the larvae.
Myxoma virus (Fibromavirus myxomatosis) causes the disease myxomato-
sis in rabbits and is spread by blood-sucking vector insects (e.g., mosquitoes).
In Britain, the main vector in a control programme in the 1950s was the Eu-
ropean Rabbit Flea (Spilopysllus cuniculi). This species was also introduced to
Australia to try to control the huge rabbit populations, the rabbit having been
introduced previously by European settlers for meat and skins.
The fossil record of fleas is sparse and challenging to interpret, as it is
difficult to distinguish what is a true flea or an extinct unrelated ectopar-
asite that simply looks like siphonapteran without actually being one. The
earliest unambiguous flea fossil is Tarwinia australis from the Early Creta-
ceous of Australia. Molecular studies also support a Cretaceous origin for
fleas and suggest that they diversified on mammal hosts and infected birds
and monotremes secondarily.
The relationship of fleas to other Antliophora has remained the last un-
solved puzzle in reconstructing the insect tree of life. A close relationship of
fleas to Mecoptera is supported by the vast majority of morphological and
molecular studies. However, the great debate lies in whether fleas represent
the sister group to Mecoptera, or a lineage within the latter order. If fleas are
highly modified Mecoptera, then the rank of Siphonaptera as a distinct order
is no longer necessary. However, phylogenomic data are currently conflicting.
One study suggests with confidence that fleas originate within Mecoptera,
and are most closely related to Nannochoristidae, while another study sug-
gests that both phylogenomic and morphological data are inconclusive for
a flea–nannochoristid relationship. At the moment, it might be prudent to
maintain Siphonaptera as a distinct order until more data are available, and
it is this approach we follow in this book.
Key reading
Bennet-Clark, H. C. and Lucey E. C. A. (1967). The jump of the flea: a study of the energetics
and a model of the system. Journal of Experimental Biology 47: 59–76.
Bibikova, V. A. (1977). Contemporary views on the interrelationships between fleas and the
pathogens of animal and human diseases. Annual Review of Entomology 22: 23–32.
Damgaard, P. D. B., Marchi, N., Rasmussen, S., Peyrot, M., Renaud, G., Korneliussen,
T., Moreno-Mayar, J. V., Pedersen, M. W., Goldberg, A., Usmanova, E., Baimukhanov,
N., Loman, V., Hedeager, L., Pedersen, A. G., Nielsen, K., Afanasiev, G., Akmatov, K.,
Aldashev, A., Alpaslan, A., Baimbetov, G., Bazaliiskii, V. I., Beisenov, A., Boldbaatar,
B., Boldgiv, B., Dorzhu, C., Ellingvag, S., Erdenebaatar, D., Dajani, R., Dmitriev, E.,
Evdokimov, V., Frei, K. M., Gromov, A., Goryachev, A., Hakonarson, H., Hegay, T.,
Khachatryan, Z., Khaskhanov, R., Kitov, E., Kolbina, A., Kubatbek, T., Kukushkin, A.,
Kukushkin, I., Lau, N., Margaryan, A., Merkyte, I., Mertz, I. V., Mertz, V. K., Mijiddorj,
E., Moiyesev, V., Mukhtarova, G., Nurmukhanbetov, B., Orozbekova, Z., Panyushkina,
I., Pieta, K., Smrčka, V., Shevnina, I., Logvin, A., Sjögren, K. G., Štolcová, T., Taravella,
A. M., Tashbaeva, K., Tkachev, A., Tulegenov, T., Voyakin, D., Yepiskoposyan, L., Un-
drakhbold, S., Varfolomeev, V., Weber, A., Wilson Sayres, M. A., Kradin, N., Allentoft,
M. E., Orlando, L., Nielsen, R., Sikora, M., Heyer, E., Kristiansen, K., and Willerslev,
E. (2018). 137 ancient human genomes from across the Eurasian steppes. Nature 557:
369–74.
Hirst, L. F. (1953). The conquest of plague. A study of the evolution and epidemiology.
Clarendon Press, Oxford.
Holland, G. P. (1964). Evolution, classification and host relationships of Siphonaptera.
Huang, D. (2015). Tarwinia australis (Siphonaptera: Tarwiniidae) from the Lower Cre-
taceous Koonwarra fossil bed: Morphological revision and analysis of its evolutionary
relationship. Cretaceous Research 52: 507–15.
Meusemann, K., Trautwein, M., Friedrich, F., Beutel, R. G., Wiegmann, B. M., Donath, A.,
Podsiadlowski, L., Petersen, M., Niehuis, O., Mayer, C., Bayless, K. M., Shin, S., Liu, S.,
Hlinka, O., Minh, B. Q., Kozlov, A., Morel, M., Peters, R. S., Bartel, D., Grove, S., Zhou,
X., Misof, B., and Yeates, D. K. (2020). Are fleas highly modified Mecoptera? Phyloge-
nomic resolution of Antliophora (Insecta: Holometabola). bioRxiv. doi: http://dx.doi.
org/10.1101/2020.11.19.390666
Kwak, M. L., Madden, C., and Wicker, L. (2017). The first record of the native flea Acan-
thopsylla rothschildi Rainbow, 1905 (Siphonaptera: Pygiopsyllidae) from the endangered
Tasmanian devil (Sarcophilus harrisii Boitard, 1841), with a review of the fleas associated
with the Tasmanian devil. The Australian Entomologist 44: 293–6.
SIPHONAPTERA 209
Linardi, P. M., Beaucournu, J. C., de Avelar, D. M., and Belaz, S. (2014). Notes on the genus
Tunga (Siphonaptera: Tungidae) II—neosomes, morphology, classification, and other
taxonomic notes. Parasite 21: 68.
Rothschild, M. and Ford, B. (1964). Breeding of the rabbit flea (Spilopsyllus cuniculi Dale)
controlled by reproductive hormones of the host. Nature 201: 103–4.
Rust, M. K. and Dryden, M. W. (1997). The biology, ecology and management of the Cat
Flea. Annual Review of Entomology 42: 451–73.
Tihelka, E., Giagomelli, M., Huang, D.-Y., Pisani, D., Donoghue, P. C. J., and Cai, C-Y. (2020).
Fleas are parasitic scorpionflies. Palaeoentomology 3: 641–653.
Zhu, Q., Hastriter, M. W., Whiting, M. F., and Dittmar, K. (2015). Fleas (Siphonaptera) are
Cretaceous and evolved with Theria. Molecular Phylogenetics and Evolution 90: 129–39.
210
Diptera
(flies)
Common name True or two-winged flies Metamorphosis Complete (egg, larva,

Derivation Gk. di–two, pteron–a wing pupa, adult)
Size Body length 0.5–60 mm. Distribution Worldwide
Wingspan up to 75 mm Number of families About 180
Known world species > 158,000 (13.96%)
Sarcophaga carnaria — Diptera
DIPTERA 211
Key features
• abundant and ubiquitous
• one pair of wings used for flight (although some wingless)
• some species have immense economic impact through disease transmis-
sion
• the fourth-largest order
Flies dominate most

ecosystems, being predators,
herbivores, parasitoids, or
© Anne Riley
decomposers. Dolichopus
ungulatus feeding on an
enchytraeid worm. UK.
Nuptial gifts are frequently

used to entice the opposite
sex, as seen here in two
© Anne Riley
mating empidid flies. While

the female is eating a dead
fly, the male mates with
her. UK.
Dipterans are important

pollinators. This Stylogaster
sp. from Malaysia, is about to
feed on nutritious nectar.
Females of this genus are
© Zestin Soh
endoparasites, laying eggs
into insects driven out by
army ant raids.
Flies are one of the dominant insect orders in most habitats. Most of the
species that make up this huge and diverse order are beneficial to the func-
tion of ecosystems as pollinators, parasites, and predators, and vital to the
processes of decomposition and nutrient recycling. However, the word ‘fly’
usually conjures up images of dirt, disease, and death. The activities of rela-
tively few species have a greater impact on humans and other animals than
any other insect group. Enormous numbers of wild and domestic animals,
and perhaps as many as one person in six, are affected by fly-borne diseases.
Tremendous losses of crops and other important plants are also caused by
flies.
Insects in other orders have ‘-fly’ as part of their common names but, in
these cases, the name is written as one word, for example caddisfly, scorpi-
onfly, and mayfly. If the common name does refer to a dipteran, then two
words should be used (e.g., horse fly, house fly, and hover fly). In keeping
with the other very large orders, such as the Hymenoptera and Coleoptera,
it is estimated that the number of species that have still not been described
equals the number already named, if not considerably more.
Fly species are recognized by the possession of a single pair of membra-
nous, front wings, although secondary winglessness occurs in some ectopara-
sitic species, such as the Sheep Ked, Melophagus ovinus (Hippoboscidae). The
Wing hind wings are reduced to form a pair of ‘gyroscopic’ organs called halteres.
Haltere
These elegant gyroscopes, which beat at the same frequency as the wings,
but out of phase, provide vital information to the flight system to keep the
insect flying straight. Groups of surface stress receptors called campaniform
sensillae are located in dorsal and ventral groups at the base of halteres and
respond to the minute distortions in the cuticle created during flight. The sen-
sillae are connected directly to the flight control systems by nerves and allow
Dorsal view of a tipulid the fly to respond instantly to changes in pitch, roll, or yaw. Visual informa-
thorax. The hind wings are
reduced to form a pair of tion from compound eyes and the brain is sent directly to the muscles that
halteres. control the halteres, not to muscles that control the wings. In turn, signals
DIPTERA 213
from the halteres are relayed to the wing muscles. In this manner, the hal-
teres, acting as flight stabilizers, can be fine-tuned by images from the eyes.
The brain does not have a flight control centre, and the task is undertaken
peripherally by a system of super-fast reflexes.
The thorax is modified with a very reduced front and hind segment. The
middle segment, or mesothorax, is enlarged and packed with the flight mus-
cles. The muscles operating the wings are not attached directly to the wing
bases as in more primitive fliers, such as the dragonflies. Instead, the muscles
alter the shape of the highly elastic, box-like thorax, and the wing hinges are
arranged in such a way that rapid oscillations of the thoracic components are
translated into wing motion (up to 1,000 wing beats per second). The flight
systems that allow this phenomenal degree of aerial agility are complex and
are not fully understood. Hovering, backward flight, 360 degree turns, and
even upside-down flight and landing are nothing unusual to these insects.
These aerial skills are essential to survival and are even brought into play dur-
ing mating. Copulation in horse flies (Tabanidae), for example, takes place on
the ground after bouts of hovering and high-speed interception manoeuvres
by the males.
The order is divided into two suborders (see Box): the more plesiomorphic
Nematocera (26 families) and the Brachycera (104 families). The Nematocera,
or ‘thread-horned’ flies (the horn refers to the antenna), such as crane flies,
mosquitoes, black flies, midges, and fungus gnats, are delicate and often have
A crane fly larva (or slender bodies, legs, and wings. Nematoceran antennae are elongate, thread-
leatherjacket).
like, and multi-segmented. The larvae of many nematocerans are aquatic, and
the adult females of several families are blood feeders. The Brachycera, or
short-horned flies, are more robust with short, stout antennae of fewer than
six segments.
Within the Brachycera, two divisions are recognized (see Box). The Or-
thorrhapha (19 families) typified by the horse flies (Tabanidae), robber
flies (Asilidae), and long-legged flies (Dolichopodidae), and a large group
of ‘higher’ flies called the Cyclorrhapha (85 families), containing families
such as the hover flies (Syrphidae), house flies (Muscidae), and blow flies
(Calliphoridae).
Typically, flies have a mobile head with a pair of large compound eyes,
and three ocelli are usually arranged in a triangular array on top of the head
between the eyes. Dipteran mouthparts, which are designed for the inges-
tion of liquid foods, ranging from blood and nectar to the products of plant
or animal decay, form a more or less elongate rostrum or proboscis. There
are two basic types of mouthparts: biting/sucking or licking/sponging. The
first type is typical of primitive flies like mosquitoes, while the second type is
more characteristic of the higher Diptera, such as muscid or calliphorid flies.
The rostrum usually incorporates the labrum, which may be small and flap-
like, but is usually elongated to form the roof of the canal along which food
is sucked. The floor of the food canal is made up of either the elongate hy-
popharynx or the overlapping mandibles (mainly in predatory species). The
hypopharynx is very deeply grooved along its length and carries the salivary
secretions. The slender, elongate portions of the maxillae, which are not fused
into the proboscis, are called the lacinia. The labium, which is usually the
largest component of the mouthparts, terminates in two sensory labial palps
(labella). In the higher flies, the labial palps, which are traversed by fine pseu-
dotracheae, function as sponges. To suck up food, flies have two muscular
pumps: the cibarial pump at the base of the food canal and the pharyngeal
A typical brachyceran fly. pump between the pharynx and the gut.
Principal dipteran taxa
Suborder Principal Larvae: pupae Adults

(division) families
Nematocera Tipulidae Well-defined head Antennae thread-
Culicidae capsule (eucephalic). like with at least 6
Chironomidae Mandibles present segments (usually a lot
Ceratopogonidae moving in horizontal more). Maxillary palps
Simuliidae plane. Many species long (3–5 segments)
Bibionidae detritivorous or filter
Cecidomyiidae feeders, but some
Mycetophilidae fungivorous, gall for-
mers and predators.
Pupa not enclosed in
larval cuticle
·········································································································································································
Brachycera Tabanidae Head capsule re- Antenna short, fewer
(Orthorrhapha) Stratiomyidae duced (hemicephalic). than 6 segments. Last
Asilidae Mandibles present, segment elongate or
Bombyliidae moving in vertical with bristle-like arista.
Empididae plane. Many species Maxillary palps short
Dolichopodidae predacious or par- (1–2 segments)
asitic. Pupa not
enclosed in larval
cuticle
·········································································································································································
Brachycera Phoridae Head capsule ves- Antenna short, fewer
(Cyclorrhapha) Syrphidae tigial, retractable than 6 segments. Last
Conopidae (acephalic). Mandibles segment elongate or
Tephritidae absent; function re- with bristle-like arista.
Ephydridae placed by specialized Maxillary palps short
Drosophilidae mouth hooks. Larval (1–2 segments)
Chloropidae feeding types very
Anthomyiidae varied. Most species
Muscidae saprophagous (includ-
Calliphoridae ing dung and carrion
Sarcophagidae feeders); others her-
Tachinidae bivorous, predacious,
Oestridae parasitic or para-
Glossinidae sitoids. Pupa enclosed
Hippoboscidae in last larval cuticle
(puparium)
DIPTERA 215
Antenna Maxillary palp
Maxillary palp
Maxillary palp
Hypopharynx
Labellum Labrum Labrum
(a)
Maxilla Mandible (c)
Labium (b) Labellum
Labellum Hypopharynx
Mouthparts of some species
of the Diptera.
.
(a) Tipulidae (crane fly)

‘primitive’ mouthparts;
(b) Culicidae (mosquito) The mouthparts of a mosquito are made up of a bundle of slender, elon-
piercing-sucking mouthparts; gate stylets surrounded by the labium. Only females suck blood and have
(c) Muscidae (house fly)
sponging-lapping fully developed mouthparts. Before feeding, the female uses her labella to test
mouthparts. the host’s skin. To penetrate, the whole proboscis is pressed onto the surface,
and the stylet bundle (made up of the labrum, mandibles, maxillae, and hy-
popharynx) is sawed through the tissue towards a blood vessel. The ends of
the mandibles and maxillae have serrated edges. To allow deep stylet penetra-
tion the labium folds back on itself. Saliva is pumped down the hypopharynx
and the blood meal is sucked back up the food canal.
The mouthparts of a house fly, which are made up from fusion of the
clypeus, maxillae, and labium, can be extended by means of muscles and
haemolymph pressure. Paired labial palps, located at the end of the proboscis,
form a sponge-like structure (the labellum) traversed by grooves called pseu-
dotrachea. Salivary secretions are pumped down the hypopharynx and flow
on the undersides of the labellum. The pseudotracheae allow saliva to act on
a large area of food, and the dissolved liquid flows into the pseudotracheae by
means of capillary action to then be drawn back up the food canal. In these
flies, mouthparts, such as the mandibles and maxillary lacinia, are lost.
Warning colouration and mimicry of other insects, particularly hy-
menopterans, is common in many dipteran families. Bees and wasps can
be equipped with powerful stings, so the ability to look or behave like a
bee or wasp affords flies protection from some of their enemies. Hover flies
(Syrphidae) are especially well known for their very wasp or bee-like ap-
pearance. Sometimes flies imitate hymenopterans not only to avoid attack,
but also in order to prey on their larvae. Species of African carpenter bees
(Xylocopa, Anthophoridae) are perfectly mimicked by species of robber fly
(Hyperechia, Asilidae). The female robber flies lay their eggs inside the nests
of the carpenter bees. The fly larvae eat the bee larvae and their provisions.
Diptera are mostly all bisexual, and parthenogenesis is rare. The majority
of adult flies do not live very long, and their main task is to find a mate
and lay eggs. Courtship, which varies enormously between families and be-
tween species, can involve swarming displays, sound production, dances,
nuptial gifts, pheromones and vibrational or acoustic songs. Drosophilid and
tephritid flies (Drosophilidae and Tephritidae) are well known for the species-
specific, short-range courtship songs that they make by vibrating their wings.
The males in several families of flies fight for access to females and may have
exaggerated physical traits that are sexually selected. The incredible stalked-
eyed flies (Diopsidae) are an example. In several families, the males of some
species have strange head modifications. In some, only the eyes, or the eyes
and antennae, are placed on stalks, while, in others, the head bears bizarre
antler-like outgrowths. In all these cases, the males display to females and
engage in male-to-male ritual combat. Females seem to prefer to mate with
larger males (indicated by the width between the eyes) and, like deer, smaller
males lose out in any fights over territory.
Dance flies (Empididae) are predaceous on other insects, particularly
other flies. The males offer nuptial gifts of food to their mates, and the size
of the gift, and therefore how long the females feed, is related to how much
Larva and pupa of mosquito sperm a male can transfer to the female during that time. Males have to be
Culex pipiens. careful not to offer too large a prey item, as they fly in tandem with the female
while she feeds, and thus have to support their own weight plus the combined
weight of their mate and the nuptial gift. In some dance flies, the females form
swarms and await the approach of a gift-bearing male.
Copulation often takes place on the wing, and eggs are laid in the vicin-
ity of larval food, as they generally hatch quickly. In tsetse flies and some
ectoparasitic fly families, the eggs hatch inside the mother and the larvae de-
velop internally, feeding on special secretions through a pseudoplacenta. Fly
Larva and pupa of a larvae, or maggots as some are called, are cylindrical and often elongate, grub-
chironomid midge.
like or worm-like, and, although they are highly variable in appearance, all
lack thoracic legs. Nematoceran larvae go through four developmental stages
or instars, and have a conspicuous, often dark, sclerotized head with biting
jaws that move in a horizontal plane.
Brachyceran flies have between five and eight larval instars. Their larvae
have a reduced head capsule with vertically biting mouthparts. Larvae of the
cyclorrhaphan flies do not have any distinct head capsule. The tapered, ante-
rior end is a dark, sclerotized cephalopharyngeal skeleton, of which the most
conspicuous part is a pair of mouth hooks. The transformation from larva
to adult takes place during the pupal stage and, in the higher flies (Cyclor-
Larva and puparium of a rhapha), pupation occurs inside the skin of the puparium, the third and last
cyclorrhaphan fly (e.g., larval stage.
blow fly). The immense diversity of larval habitats and lifestyles is one of the major
factors in the success of the order as a whole. Dipteran larvae can be herbiv-
orous, predacious, parasitic, saprophagous, coprophagous, or fungivorous in
all manner of freshwater and terrestrial habitats. The larvae of certain non-
inquiline biting midges (Chironomidae) even live as inquilines inside the trap leaves
a species that lives in the nest, of pitcher plants and are responsible for accelerating the breakdown of prey
gall, or home of another items, and the uptake of nitrogen by the host plant.
DIPTERA 217
Species such as shore flies (Ephydridae) can even be found in some very
harsh environments, such as pools of crude oil, hot springs, and the highly
alkaline or saline water of marshes and mangroves.
Until the seventeenth century, it was thought that maggots in dead ani-
mals arise there by spontaneous generation. In 1668, Francisco Redi proved
that maggots hatched from the eggs of flies. Flies and other insects feeding
on carrion arrive and use the resources in a predictable succession associated
with each stage of decay. Knowledge of the ecological succession under differ-
ent conditions is of great use to forensic scientists, especially when deciding
on the time of death in murder cases. The enormous scale of the recycling
services carried out by flies should not be underestimated.
Although broad larval feeding patterns exist within each family, there
are many exceptions. The larvae of most fungus gnats (Mycetophilidae and
Keroplatidae) are fungivorous, but the larvae of Arachnocampa luminosa
(Keroplatidae) catch prey in an unusual manner in New Zealand’s Watiomo
Cave. The larvae of these and other Arachnocampa species in New Zealand
and Australia live in caves and catch flying prey on sticky threads that hang
from the roof. The larvae lure their prey, which emerge from streams flowing
through the caves, by being luminescent. The adults are also able to glow and
use this for mate location in the darkness of the caves.
At least 20% of all fly species, representing more than 20 families, are par-
asitoids, that is, their larvae develop inside the bodies of hosts, which are
killed in the process. Larvae of flies belonging to the family Conopidae are
mostly endoparasitoids of aculeate Hymenoptera, although some species at-
tack crickets, cockroaches, and other flies. Adult conopids are usually found
on flowers, and often look very wasp-like, but this is for protection from
predators and not to fool their hosts. In line with the findings in many
parasite–host interactions, conopid larvae are able to manipulate the be-
haviour of their hosts. In one case, a conopid parasitoid of the bumble bee
(Bombus terrestris) causes its host to burrow into the soil just before it dies.
This is highly advantageous to the fly, because it now has a safe and protected
pupariate site in which to pupariate and overwinter.
a term used of higher flies. The The bumble bees can fight back a bit, and it has been shown that parasitized
fly pupates inside a puparium
which is the hardened skin of worker bees do not return to their nest as usual but stay outside all night.
the last larval stage This slows up the growth of the parasitoid inside and may increase the bee’s
chances of survival by decreasing the chances of the parasitoid’s successful
development.
More than 8,200 species belong to the widespread family Tachinidae.
These flies are typically stout bodied and many species resemble bristly house
flies, but some can be much larger, very hairy, and almost bee-like. Adults
feed on carbohydrate-rich food sources, such as honeydew, and males often
congregate on hilltops, waiting for receptive females. Larval tachinids are par-
asitic on other insects and very rarely on other arthropods. The range of insect
hosts used by these flies is extensive and includes the adult and immature
stages of Lepidoptera, Coleoptera, Orthoptera, Hemiptera, Hymenoptera,
Diptera, and some other orders. The method by which the host is parasitized
varies from the simple to the complex. Eggs can be laid directly on the host
whereupon they hatch and burrow inside, or they can be laid through the
skin of the host. The females of some species lay their eggs on an appropriate
plant, which is then eaten by the host, and, in some cases, the tachinid female
lays directly into the mouth of the host insect while it is feeding. While the
larvae consume the host’s tissues, eventually killing it, they obtain air from
the outside through a hole in the host’s body or by ‘plugging in’ to a trachea.
Tachinids are of immense importance in the control of natural populations
of pest insects. Many species have been used or are under investigation as bi-
ological control agents. For example, Triarthria setipennis was introduced to
North America from Europe to control its host, the European earwig (For-
ficula auricularia). Another well-known species is Cyzenis albicans, which
has been used against Winter Moth (Operophtera brumata), whose larvae
can cause serious defoliation of a wide range of trees. The host-finding and
egg-laying behaviour of some of these robust hairy flies is intriguing. Take,
for example, Ormia ochracea whose larvae are parasitic inside the bodies
of crickets. Mated female flies lay larvae on the outside of several species
of Gryllus hosts, and the larvae burrow through the intersegmental mem-
branes. Once inside, the young larvae feed on thoracic muscle tissue. The
larvae then move to the abdominal cavity, moult, and feed on the fat body as
well as other tissues. The host is killed when the fly pupariates and emerges.
But how do the female tachinids find their host? The same way the female
crickets find their mates—by the love songs of the male cricket. The flies
locate the crickets by listening to the songs they emit, and as a result, they
can parasitize male crickets and the females they attract. The story does not
end there. It seems that both female crickets and female flies are preferen-
tially attracted to the same qualities in the male song, and there is evidence
that parasitoid pressure has shaped some of the characteristics of the cricket’s
songs. In fact, Ormia ochracea exerts such strong selection pressure on its
hosts, that most individuals of a cricket species in Hawaii have lost their ca-
pacity to produce songs altogether, in order to escape the attention of the
flies. Instead, they gather around the few males that are still capable of pro-
ducing songs and steal the females that come by! The location of hosts by
acoustic orientation also occurs in other species of tachinid that parasitize
tettigoniids. Previously, it was thought that tachinid attack always resulted in
the host’s death, but recent work suggests that non-lethal parasitism might
be quite common and might even explain the evolution of some host plant
choices. In this study, the survival of a caterpillar (Platyprepia virginalis) to a
tachinid attack (Thelaira americana) depended on what it ate. If parasitized,
the caterpillars chose to eat poison hemlock, and if not parasitized, the cater-
pillars ate lupin. Parasitized caterpillars that ate poison hemlock survived
better than those ate lupin, and, furthermore, the flies that emerged from the
hemlock-fed caterpillars were heavier than those that emerged from lupin-fed
hosts.
DIPTERA 219
The range of feeding habits displayed by the larvae of hump-backed or

scuttle flies (Phoridae) is very large; some feed on fungi, some on decay-
ing matter or carrion, while others act as scavengers in the nest of ants and
termites. Many are parasites on species in a number of insect orders, snails,
and millipedes. There are even predatory adult phorids that eat on fig wasps,
and whose larvae develop inside the fig fruit. The males and females of some
ant-parasitizing species are very attracted to the alarm pheromones given off
when their hosts fight. In the leaf-cutting ant Atta cephalotes, workers use tiny
‘minim’ workers to ride shotgun on top of their leaf loads to ward off other
species of phorid flies that would otherwise find them easy targets. The minim
workers are recruited by vibrational signals produced by the larger workers
as they cut their leaf fragments. Other species of phorid fly who are specialist
parasites of ant species can exert considerable influence of the behaviour of
their host, causing them to forage at night, collect less food, and even defend
themselves from enemies less effectively.
Many fly larvae are herbivorous pests. The larvae of the Hessian fly
(Mayetiola destructor), a species of gall midge (Cecidomyiidae), burrow
through the stems of grass crops like wheat, barley, and rye. Oscinella frit
(Chloropidae) is another serious cereal pest in the northern hemisphere. The
larvae of soldier flies (Stratiomyiidae) and rust flies (Psilidae) bore into root
crops, and leaf mining species (Agromyzidae) attack a wide range of crops like
alfalfa, cucumbers, and tomatoes. However, the major impact on humans re-
lates to disease-carrying and blood-sucking dipterans (see Box). Many species
of flies, such as muscids and calliphorids, are attracted to carrion, dung,
and decaying matter, and can act as mechanical vectors of large range of
pathogenic organisms that cause intestinal complaints, such as dysentery,
and other diseases. The larvae of some flies invade human tissues, and oth-
ers can act as vectors for human pinworm, hookworm, and roundworm, but
the biggest threat by far comes from blood-feeding nematocerans that act as
vectors for major diseases like malaria and yellow fever.
Many plants make use of flies as pollinators. The huge flower of the rare,
parasitic tropical plant Rafflesia has evolved the scent of rotting meat to at-
tract carrion flies. Although the flower is large, the flies are guided down
tight channels around the central column of the male flower where they pick
up sticky pollen on the back of their thorax. The flies are similarly confined
to tight spaces in the female flower where they come into contact with the
stigma.
Horse flies (Tabanidae) comprise more than 4,500 known species world-
wide. Common names for these insects, which have large and often patterned
eyes, include horse flies, deer flies, clegs, or gad flies. Adult females feed on
mammalian blood while males feed exclusively on plant sugars. Tabanid lar-
vae typically inhabit wet muddy areas by the margins of ponds, streams, or
Phlebotomus papatasi is a rivers, and eat small insects, worms, and other invertebrates. Most species are
vector of the viral disease nuisance pests, and in the warmer parts of the world, they are vectors of sev-
sand fly fever in
eral animal diseases. Two species of Chrysops are vectors of Loiasis, a filarial
Mediterranean and Oriental
regions. disease that affects humans, in west and central Africa.
The fly vectors and causative organisms of some human diseases
Disease Vector Organism

Anthrax Horse flies. Tabanus spp. Bacterium. Bacillus anthracis
(Tabanidae)
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Dengue fever Mosquitoes. Aedes aegypti and Virus. (Flaviviridae)
Culex spp. (Culicidae)
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Elephantiasis Mosquitoes. Culex fatigans Nematode. Wuchereria
(Culicidae) bancrofti
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Encephalitis Mosquitoes. Aedes and Culex Virus. (Flaviviridae)
(several) spp. (Culicidae)
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Filariasis Biting midges. Culicoides spp. Nematode. Mansonella and
(several) (Ceratopogonidae) Dipetalonema spp.
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Leishmaniasis Sand flies. Phlebotomus and Protozoan. 12 species of
(several) Lutzomyia spp. (Psychodidae) Leishmania
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Loiasis Horse flies. Chrysops spp. Nematode. Loa loa
(Tabanidae)
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Malaria Mosquitoes. Anopheles spp. Protozoan. 4 species of
(several) (Culicidae) Plasmodium
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Onchocerciasis Black flies. Simulium spp. Nematode. Onchocerca
(Simuliidae) volvulus
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Sleeping Tsetse flies. Glossina spp. Protozoan. Trypanosoma
sickness (Glossinidae) brucei
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Tularaemia Horse flies. Chrysops spp. Bacterium. Francisella
(Tabanidae) tularensis
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Yellow fever Mosquitoes. Aedes aegypti and Virus. (Flaviviridae)
other spp. (Culicidae)
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Zika virus Mosquitoes. Aedes aegypti and Virus. (Flaviviridae)
A. albopictus
One of the most remarkable examples of insect engineering is the con-

struction of mud cylinders by African species such as Tabanus biguttatus. The
larvae live in pools, which eventually dry out. When the larva is fully grown, it
first descends through the mud following a tight spiral path. It then spirals up-
wards, cutting across the descending channels thereby making a near-isolated
cylinder of mud. Near the surface, it burrows straight up and enters the centre
of the cylinder from the top. Once inside, the larva forms a sealed pupation
chamber. As the mud dries out and bakes hard, cracks forming across the
surface bypass the cylinders, leaving the pupating fly safe inside.
Although not vectors of any human disease, biting midges (Cerato-
pogonidae), such as the Scottish Biting Midge (Culicoides impunctatus), are
well-known pests, and many parts of the northern hemisphere are rendered
virtually uninhabitable due to these and other species of biting fly, like black
flies and horse flies. Culicoides species of the imicola complex can, however,
be vectors of serious viral animal diseases like African horse sickness and
Bluetongue (Reoviridae).
DIPTERA 221
Fly larvae in the family Oestridae are parasitic and eat the flesh of various
animals. Species of calliphorid and sarcophagid flies lay their eggs or larvae in
carrion, but some will also lay eggs in live tissue—a condition called a myia-
sis. Flies from all these families have been known to cause myiasis in humans.
Although the principal hosts are rats, larvae of the Tumbu Fly (Cordylobia
anthropophaga), a calliphorid species from tropical Africa, can burrow into
human skin. Eggs can be laid on damp bedding or clothes, and, when hatched,
a young larva can burrow under skin in less than a minute. A large painful
swelling appears as the larva feeds and grows. The larva leaves the host to pu-
pate in just over one week. Another calliphorid, the New World Screw-worm
(Cochliomyia hominivorax), is an important pest of cattle, sheep, and horses
in the North and South America, and will lay eggs on wounds, scratches, and
even mucous membranes. The larvae burrow deeply into tissue and make a
large boil or lesion, which can lead to disfigurement or even death. Control
of this scourge has been achieved through the mass release of males, which
have been sterilized by irradiation.
The Human Bot Fly (Dermatobia hominis) lives in Central and South
America and uses a clever trick to provide their larvae with a suitable place to
live. The females catch blood-feeding flies, lay their eggs on them, and then
release them unharmed. When the eggs mature and the flies next feed on a
mammalian host (including humans), the eggs hatch and the larvae burrow
under the skin. In the cases where the larvae have been deposited on a human
scalp, the afflicted person can hear the bot fly larvae munching through their
flesh at night. Blow flies, such as Lucilia spp. (Calliphoridae), are responsible
for flystrike of sheep. Traps baited with sheep offal can reduce local popu-
lations of these flies, but they are very significant pests in many parts of the
world.
Warble flies or bot flies (Oestridae) live by laying live larvae in the noses
or in the hair of their mammalian hosts. The larvae either feed in the nasal
passage or burrow extensively under the skin. The cattle grubs, Hypoderma
lineatum and Hypoderma bovis, are serious pests of domestic cattle in the
northern hemisphere. Adapted to the migrations of their hosts, some species
have an incredible ability to fly for long periods. A mated female Reindeer
Warble Fly can cover up to 900 km during her life and may lose up to 40% of
her initial body mass.
Gruesome as it might seem, sterile calliphorid larvae of certain species are
sometimes used to clean infections which are difficult to treat with antibiotics.
The grubs eat infected flesh and leave healthy tissue alone.
Mosquitoes (Culicidae) probably have a greater harmful effect on hu-
mans than any other insect family. They are slender, delicate flies with small,
subspherical heads and elongate, slender, sucking mouthparts. The body is
covered with tiny scales and appears pale brown to reddish brown. The wings
are long and narrow with scales along the veins and margins. Adults belong-
ing to the two important subfamilies, Anophelinae and Culicinae, can be
recognized by the way they rest. Anopheline mosquitoes (Anopheles spp.) rest
with their heads down and the body inclined at 45 degrees, whereas culicine
(Aedes and Culex) species rest with their bodies held horizontally.
It is only the females that suck mammalian blood and therefore transmit
disease. Males feed on nectar or honeydew. Anopheles eggs have small lateral
air floats, and are laid singly on the surface of water, whereas Culex eggs are
laid in floating rafts of 40–300 eggs. The larvae can be found almost anywhere,
from rain-filled containers and tree holes to ponds and lakes, and are mostly
saprophagous, although some are predacious. In most species, air is obtained
from the surface. Anopheline larvae lie horizontally under the surface while
culicine larvae hang at an angle from the surface film by a posterior siphon.
(a) The life cycle takes between 15–20 days.
Many mosquitoes are very common vectors of human diseases such as
yellow fever, dengue fever, filariasis, and encephalitis, but malaria is by far
the most serious. Malaria has killed people throughout the course of human
history, and its effects are described in the Old Testament and even earlier
(b) texts. Causing more than 400,000 deaths every year, the majority of them pre-
school age African children, malaria’s impact continues to be immense. It is
estimated that one-third of all human beings live in regions of the world where
infection by the malarial parasite is likely, and 94% of all cases occur in Africa.
Malaria is only carried by mosquitoes belonging to the genus Anopheles.
The discovery of the malarial parasite (Plasmodium) by Charles Laveran
Typical resting position of
in 1880, and the elucidation of the role played by the mosquito in the life
(a) the culicine mosquito cycle of the parasite by Ronald Ross in 1898, made prevention a possibility.
and (b) the anopheline The disease had been treated with the bark of the cinchona tree from which
mosquito.
quinine was eventually purified, and, to complete the victory, the breeding
grounds of the mosquitoes were drained or covered with oil to suffocate
the larvae and huge amounts of insecticides were used to kill adults. The
development of DDT and its extensive use after the Second World War
did, indeed, rid many countries of this terrible scourge, but it was not long
before the mosquitoes developed resistance to insecticides, and the use
of some were banned because of persistence and the harmful effects they
had on non-target organisms. Today, there is clear evidence of a massive
resurgence of the disease, partly due to resistance of the four species of
Plasmodium parasites to drugs, such as chloroquine, and the resistance of
the vector mosquitoes to insecticide, but also to the drug industry’s lack
of interest in producing new anti-malarial compounds for a long period of
time. The hopes of a vaccine promised by molecular biology are diminishing,
but new ideas involving genetic engineering may still win the war. Certain
approaches include the release of millions of transgenically sterilized male
mosquito vectors, which would mate with most females and outcompete
fertile males, resulting in a huge drop in their overall numbers. Other
approaches include the introduction of transgenic males that possess a gene
that destroys X-chromosomes and results in male-only progeny. One thing
is certain: the fight against malaria is ongoing and will likely continue for a
long time. Recent evidence of global climate change suggests that the range
of malaria may increase. The disease occurred in many parts of southern
Europe (including England) in the past, and as the vector mosquitoes are
still present, there is a real risk that reintroduction could take place.
Tsetse flies belong to the family Glossinidae and are dull brown or grey
coloured, medium to large-sized flies (6–14 mm long). There are 21 known
DIPTERA 223
species, all found in subtropical and tropical parts of Africa, and their pres-
ence causes immense human suffering and renders large areas unfit for cattle
rearing. In total, some 10 million km2 of land are affected and 70 million hu-
mans are at risk from sleeping sickness—the disease carried by these flies.
Both males and females feed on blood, and their needle-like mouthparts can
penetrate the toughest skin or hide. Tsetse flies can be distinguished from
other sorts of flies, such as horse flies, because some of the wing veins mark
out a distinctive hatchet-shaped cell, and at rest, the wings are flat and crossed
scissor-like over the abdomen. Different tsetse species are found in different
types of habitat and prefer to bite different sorts of animals. Some are only
found in east African countries, while others are more widespread. Some
species breed in savannah woodlands, while others prefer coastal areas or
damp, riverside forests.
Female tsetse flies are unusual in that they produce only one egg at a
time, which hatches and stays inside the mother’s body. Safe inside a struc-
ture called a uterus, the larva feeds on secretions produced by special nurse
or ‘milk’ glands. When fully grown, the tsetse larva is large and may even
weigh a little more than its mother. Larvae are deposited on the ground, or in
the host’s nest or resting place, where they pupate almost immediately. Ade-
notrophic viviparity, as it is called, also occurs in the Hippoboscidae (a family
of more than 200 fly species, which are blood-sucking parasites of birds and
mammals), the Nycteribiidae, and the Streblidae (two similarly sized families
of blood-sucking parasites of bats). The advantages of being able to carry and
feed young until they are just about to pupate are considerable and do away
with the need for the larvae to fend for themselves. After a three- or four-
week pupation, the adult flies emerge from the soil and find a mate within
a day of two of emergence. Females store sperm inside a spermatheca, and
only need to mate once in their lives. The first larva will be produced in 2–3
weeks and, depending on the species, female tsetse flies will produce more
larvae every 8–10 days. Both sexes need blood meals every 2–3 days and dif-
ferent species exhibit preferences for cattle, horses, pigs, humans, and even
crocodiles. In the wild, males live for about six weeks as adults, but females
can live for more than three months.
One hundred years ago a Scottish microbiologist, David Bruce, serving
in the Royal Army Medical Service, and his wife Mary Elizabeth Bruce (née
Steele), went to Africa to investigate the cause of a disease of ruminant an-
imals called nagana. In three years, they found that the disease was caused
by protozoan organisms called trypanosomes (named Trypanosoma brucei,
in their honour) that lived as parasites in the blood and tissues of affected
animals. They also discovered that human sleeping sickness was caused by
the same parasite, and that it was transmitted by tsetse flies when they fed on
animal or human blood.
Once infected, a tsetse fly carries the parasite for the rest of its life and
passes it on to the animals on which it feeds. The parasites are injected into
the host animals along with the fly’s saliva. Only about one in a hundred flies
is infected with the parasite, so not every bite will result in the infection be-
ing passed on. People and animals get infected because they get bitten many
times. In west and central Africa, domestic pigs are the main animals carrying
the disease, whereas in east Africa it is mainly cattle and wild animals such as
antelopes that are infected. Animals like antelope and buffalo are unaffected
by the parasites, but they act as reservoirs from which the tsetse flies can pick
up the infection. Although, in theory, all species of tsetse fly can transmit
the trypanosome parasite to humans or domestic and wild animals, only a
few—Glossina palpalis, Glossina fuscipes, Glossina tachinoides, Glossina mor-
sitans, and Glossina pallidipes—are regular human-biting species associated
with outbreaks of sleeping sickness.
In humans, the first sign of the disease is a local inflammation and swelling
of the skin where the fly took a blood meal. Next, the victim will have
influenza-like symptoms with a fever and chills, headache, and slight swelling
of the hands and feet. Initially, only the bloodstream and the lymphatic sys-
tem are infected, but damage to the brain and nervous system will also occur.
This may take only a few weeks to happen, or it could take several years, but
when it does, the victim feels sleepy and tired all the time, and may have con-
vulsions and go into a coma. Untreated, the disease is almost always fatal,
but some types of drug, if given early enough, can help. On a positive note,
the number of cases is steadily plummeting, with about 1,000 new infections
annually (compared to many tens of thousands in 1990s). Production of a
vaccine is difficult as the parasite keeps changing slightly so that vaccine will
not work properly.
In tsetse fly areas, prevention mainly involves trying to avoid being bitten,
but this can be difficult because the flies are active during the hottest time of
the day, not easily discouraged, and can bite through clothing. Many methods
of control have been attempted, and these are aimed at trying to get rid of the
flies themselves. This is not an easy job as the areas they live in are vast, and
the flies can travel long distances. Tsetse fly control has included extensive
habitat clearance, and the use of insecticides, but due to the low densities at
which they exist, traps baited with animal dung and urine are more effective.
The biconical trap is a clever device for catching tsetse flies. It works on the
principle that tsetse flies are attracted to large dark shapes, which they mistake
for animals. The bottom section, which has four oval holes around the outside,
is made of dark blue cloth. The upper section is made of white cloth. Inside
the bottom section, there are two pieces of black cloth at right angles, which
makes the inside of the trap look very dark and inviting. Tsetse flies enter
the trap through the holes, and then fly up where they enter a collecting bag.
When the bag has a good catch, it is simply emptied of dead flies. The flies are
killed by the heat that builds up inside the trap, and no insecticide is needed.
It has been proposed that the patterning of some animals may be the result
of selection to avoid tsetse fly bites. For instance, the characteristic vertical
stripes of zebras may be important in protecting them, as tsetse flies may not
see the zebras as well as they see non-striped hosts, and it has been shown
that some blood-feeding flies have an aversion to landing on striped surfaces.
The origins of Diptera go very far back in time, with the oldest fossil flies
dating to the Triassic (about 230 mya). As stated in previous chapters, the
closest relatives of Diptera are the Mecoptera and the Siphonaptera.
DIPTERA 225
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227
Amphiesmenoptera
This superorder consists of caddisflies (Trichoptera) and butterflies and
moths (Lepidoptera). Amphiesmenoptera means ‘clothed-wing’ in ancient
Greek, referring to the hairs and scales that cover the wings of Trichoptera
and Lepidoptera, respectively.
228
Trichoptera
(caddisflies)
Common name Caddisflies Distribution Worldwide except

Derivation Gk. trichos–hair; pteron–a Antarctica
wing Number of families 52
Metamorphosis Complete (egg, larva,
pupa, adult)
Thread-like
antenna
Maxillary palp
Hairy wings
Large compound
Narrow pronotum eye
Large
mesothorax Venation largely
longitudinal
Slender legs
Stenophylax vibex — Trichoptera
TRICHOPTERA 229
Key features
• aquatic larvae, typically in self-constructed cases or shelters
• nocturnal and moth-like, with hairs and sometimes scales
• legs and antennae slender
© Matt Doogue
A specimen of Stenophylax
permistus from Leighton
Moss, UK.
Many caddisfly larvae

construct a portable case
that they carry with them
wherever they go. The vast
majority are aquatic,
although the land caddis
© Paul Brock
(Enoicyla pusilla) imaged

here is exceptional in that it
inhabits leaf litter. UK.
Caddisflies can be found almost everywhere that there is fresh water. Esti-
mates point to there being as many as 50,000 species, the majority of which
will be confined to the eastern hemisphere. The slender, elongate adults are
sombre-coloured, and rather moth-like in appearance with long, slender legs.
The body and wings, particularly the front wings, are covered with hairs. Cad-
disflies are most closely related to the Lepidoptera, but the latter are covered
with scales and not hairs, and caddisflies lack the unique proboscis of most
lepidopterans. The head has a pair of large compound eyes, and long, thin,
multi-segmented antennae, which may be shorter or longer than the length of
the body. Two or three ocelli may be present. The weakly developed mouth-
parts allow adults to lick up water and nectar, but many do not feed at all.
The head and basal part of the antennae may have scent-producing organs
of various kinds, and the upper surface of the head and parts of the thorax
may have small, hairy, wart-like bumps. The front and hind pairs of wings
are membranous, similar in size, and held over the body in a characteris-
tically tent-like manner. In flight, the hind wings are coupled to the front
wings by means of special curved hairs or hooks. Two suborders are recog-
nized: the Annulipalpia and the Integripalpia (see Box). In the Annulipalpia,
the larvae construct stationary retreats, which gives them the common name
‘fixed-retreat makers’. The Integripalpia, which is the most diverse caddisfly
suborder, is characterized by larvae that make portable, often highly elab-
orate cases, while some carnivorous species are entirely free living. A third
suborder is proposed from within the Integripalpia, the Spicipalpia, although
whether this is a valid classification has remained controversial to this day.
Both Annulipalpia and Integripalpia are split into several superfamilies (see
Box).
Mostly nocturnal in habit, the adults hide in vegetation during the hours
of daylight and are quite hard to find. Mating takes place in flight (some-
times in swarms), or on vegetation. In some species, when the females have
coupled their genitalia with a male, the male alone flies the joined pair to
the water margin where copulation occurs. Females are typically the bigger
sex in caddisflies, but, in these particular species, males are larger to enable
them to cope with the load imposed by the nuptial flight. Eggs, which are
produced in masses or strings enveloped in a jelly-like substance, are either
dropped onto or under water, or laid on overhanging plants from which they
Limnephilid caddisfly larval
will eventually drop. The order is best known because of the aquatic larvae
cases vary enormously. that construct impressive and elegant cases, inside which they live and finally
Here, a species has used pupate. A typical larva is caterpillar-like with a strong head and thorax and a
small snails shells as a
construction material. soft abdomen. The head has chewing mouthparts and short antennae. There
are three pairs of five-segmented thoracic legs, each of which the larvae use to
move around, and, at the end of the abdomen, each has a pair of hooked pro-
legs to anchor it in its case. Larvae are apneustic (they have no spiracles) and
oxygen is obtained from the water either by direct diffusion or through ex-
ternal gills located on the abdominal, and, sometimes, the thoracic segments.
Caddisfly larvae may be herbivores, carnivores, or detritivores. The larvae of
some species are free-living (Rhyacophiloidea), and some make silken nets
to filter minute organic particles of food from the water currents (Hydropsy-
choidea). The majority of species make portable cases (e.g., Limnephiloidea)
and can be found in flowing and still water. The cases, made from a great
variety of materials, are held together with silk secreted from glands in the
head and fed out through the labium. Case design and construction is very
variable, tends to be specific to particular families, and in many instances, is
a useful identification feature.
TRICHOPTERA 231
Trichopteran suborders and some superfamilies
Suborder Common Larval habits: food

families
Annulipalpia
·········································································································································································
Hydropsychoidea Hydropsychidae Net spinners: invertebrates,
(7 families) algae, particles
·········································································································································································
Philopotamoidea Philopotamidae Net spinners: algae, particles
(2 families)
·········································································································································································
Integripalpia
·········································································································································································
Limnephiloidea Goeridae Case makers: algae, detritus
(9 families)
·········································································································································································
Limnephilidae Case makers: algae, detritus
·········································································································································································
Phryganeoidea (3 Phryganeidae Case makers: invertebrates,
families) detritus
·········································································································································································
Lepidostomatidae Case makers: plant detritus
·········································································································································································
Leptoceroidea (7 Leptoceridae Case makers: invertebrates,
families) algae, sponges, detritus
·········································································································································································
Sericostomatoidea Sericostomatidae Case makers: detritus
(12 families)
·········································································································································································
‘Spicipalpia’
·········································································································································································
Hydroptiloidea (2 Glossosomatidae Saddle case makers: algae and
families) detritus
·········································································································································································
Hydroptilidae Free-living then purse case:
algae and diatoms
·········································································································································································
Rhyacophiloidea Rhyacophilidae Free-living: invertebrates
(2 families)
The first four larval stages of purse casemakers (Hydroptilidae) have a

short abdomen, and in addition to the thoracic legs, they have longish abdom-
inal prolegs. They are active and free-living in water, and suck the juices of
water plants, such as Spirogyra, through modified mouthparts. The last larval
stage, which has a swollen abdomen, makes an open-ended purse or barrel-
shaped case of silk, often with small stones attached. Larvae of the northern
caddisflies (Limnephilidae) can be found in a wider variety of habitats than
any other family of trichopterans, and exhibit variable preferences in terms
of water current, temperature, and substrate texture. Their cases vary in size
(6–65 mm in length) and have a circular, triangular, or irregular cross-section
depending on species. The cases are constructed from a diverse range of mate-
rials including sand grains, pebbles, vegetation, sticks, and shells, or mixtures
of more than one type of material.
In species that exclusively use twigs as construction materials, the cases are
appropriately known as ‘log cabins’. The larvae of some limnephilid species
incorporate the occasional large twig in the case, which often projects side-
ways and deters fish from trying to swallow them. In some species, the cases
made by the younger stages can be of a different design to those made by the
older stages. Most limnephilid larvae are omnivorous, and feed on detritus
or scrape microscopic algae and other organisms from the surface of stones
and rocks. The larvae of giant casemakers (Phryganeidae) feed on plant de-
bris or are predacious on small aquatic animals, and as a result, need a strong
but light case that can be easily moved around. The beautifully regular, silk-
lined, tapering cases are made of spirally arranged plant fragments with the
occasional bit of gravel incorporated. Interestingly, left- and right-handed
spirals occur within the same species, and the plant fragments used for the
cases, which need to get longer as the case grows, are measured and expanded
accordingly by the larva.
In some families, only bits of vegetation, such as cut leaves or sticks, are
used. In others, there may be a high mineral content with everything from
small sand grains to quite large pebbles being incorporated. Some species
even use empty snail shells exclusively as their case-building material.
Cases of limnephiloid families may be long and thin, squat, or spiral in
overall shape and several studies with various predators, such as fish and drag-
onfly nymphs, have shown that the design and construction of the case has
a significant bearing on survivorship. A study of survivors of many species
shows that they have stronger and sometimes wider cases than those that were
eaten. The behaviour of the caddisfly larvae is also important. When con-
fronted with a predatory fish, many caddisfly larvae will retreat inside their
case and play dead. The fish may wait to see if the occupant moves again, but
it is usually the insect that can wait the longest. If larvae are taken out of their
cases and thus forced to rebuild them, the first priority is rapid protection,
although some species will respond to the local conditions by making the
new cases heavier in faster-flowing water. Net-spinning caddisflies, such as
the Hydropsychidae, also alter their net construction, overall size, and mesh
size in response to the amount of food present in the water and the speed at
Caddisfly larva use many
which it is flowing. They are therefore able to maximize the prey capture rate,
types of material in the
construction of their cases. while not risking damage to the net due to drag. There is evidence of larvae of
Many designs are species- algal grazing species defending patches of algae on the surface of rocks from
or genera-specific.
each other, and even from other species of grazer, such as nymphal mayflies.
The size of the patch is related to the size of the territory holder.
After between five and seven larval stages, pupation takes place within the
silk-sealed larval case or in a specially made pupal case. The whole life cycle
can take one year but is longer if the species occurs in mountainous or cold
regions. When ready to emerge, the pharate adult (enclosed inside the pupal
exoskeleton) cuts its way out of the case and swims to the surface or crawls
out of water on plants where it emerges. The wasp Argiotypus armatus (Ar-
giotypidae) is an unusual parasitoid that swims below the water surface to lay
its eggs in pupal cases of the caddisfly Silo pallipes (Goeridae). The larvae are
ectoparasites and, after five instars, the wasp pupates. It remains inside the
pupal case of its host to overwinter and emerges in the following spring.
Many water courses become polluted by industrial effluent as well as by
pesticides, fertilizers, and herbicides from farm run-off. As different caddis-
fly species vary enormously in their ability to tolerate various factors, such
as acidity, oxygen content, and chemical composition of the water, they
are invaluable indicators of changes in water quality. The levels of specific
TRICHOPTERA 233
pollutants, such as pesticides and polychlorinated biphenyls (PCBs), can be

analysed in adult body tissues.
Although the vast majority of species are freshwater dwellers, a few species
from several families can be found in brackish water, and four species in the
strange family Chathamiidae are marine (Chathamia and Philansius). Known
only from parts of New Zealand and New South Wales, the larvae of these cad-
disflies make their cases from grains of sand and fragments of algae on which
they feed. Bizarrely, the adult females of Philansius plebius, which have long
and fairly tough ovipositors, are known to lay their eggs inside the coelomic
cavity of two species of intertidal starfish. It is likely that females oviposit
through pores on the aboral surface of the starfish. Inside the coelom, the
eggs are protected from harmful waves and predation. After several weeks
of incubation, the eggs hatch and the first instar larvae leave the host, either
through the pores or, perhaps, through the stomach wall. They pass through
seven instars feeding on calcareous and non-calcareous algae.
Some molecular studies estimate that caddisflies first appeared during the
Permian (about 275 mya), which coincides with the earliest known fossils
of early Trichoptera (or close relatives). Phylogenomic studies suggest a con-
siderably younger date, between 215–154 mya. Additional data will further
disentangle the origins of caddisflies.
Key reading
Andersen, D. T. and Lawson-Kerr, C. (1977). The embryonic development of the marine
caddisfly, Philanisus plebius Walker (Trichoptera: Chathamiidae). Biological Bulletin 153:
98–105.
Bournaud, M. and Tachet, H. (eds) (1987). Proceedings of the Fifth International Symposium
on Trichoptera. Entomologica series 39. Junk Publishers, The Hague.
Ivanov, V. D. (2002). Contribution to the Trichoptera phylogeny: new family tree with
considerations of Trichoptera-Lepidoptera relations. Nova Supplementa Entomologica
(Proceedings of the 10th International Symposium on Trichoptera) 15: 277–92.
Society, London.
Mackay, R. J. and Wiggins, G. B. (1979). Ecological diversity in Trichoptera. Annual Review
of Entomology 24: 185–208.
Morse, J. C. (ed) (1983). Proceedings of the Fourth International Symposium on Trichoptera.
Entomologica series 30. Junk Publishers, The Hague.
Misof, B., Liu, S., Meusemann, K., Peters, R. S., Donath, A., Mayer, C., Frandsen, P. B., et al.
7639–67.
Morse, J. C. (1997) Phylogeny of Trichoptera. Annual Review of Entomology 42: 427–50.
Nimmo, A. P. (1996). Bibliographia Trichopterorum: a world bibliography of Trichoptera.
Pensoft, Sofia.
Otto, C. (1987). Behavioural adaptations by Agrypnia pagetana (Trichoptera) larvae to cases
of different value. Oikos 50: 191–6.
Petersson, E. (1995). Male load-lifting capacity and mating success in the swarming
caddisfly, Athripsodes cinereus. Physiological Entomology 20: 66–70.
Riek, E. F. (1976). The marine caddisfly family Chathamiidae (Trichoptera). Journal of the
Australian Entomological Society 15: 405–19.
Shields, O. (1988). Mesozoic history and neontology of Lepidoptera in relation to Tri-
choptera, Mecoptera and angiosperms. Journal of Palaeontology 62: 251–8.
St. Clair, R. M., Dean, J. C., and Flint, O. S. (2018). Description of adults and immature stages
of Antipodoecia Mosely from Australia and synonymy of the families Antipodoeciidae
and Anomalopsychidae (Insecta: Trichoptera). Zootaxa 4532: 125–36.
Thomas, J. A., Frandsen, P. B., Prendini, E., Zhou, X., and Holzenthal, R. W. (2020). A
multigene phylogeny and timeline for Trichoptera (Insecta). Systematic Entomology 45:
670–86.
Winterbourn, M. J. and Anderson N. H. (1980). The life history of Philanisus plebius Walker
(Trichoptera: Chathamiidae), a caddisfly whose eggs were found in starfish. Ecological
235
Lepidoptera
(butterflies and moths)
Common name Butterflies and moths Metamorphosis Complete (egg, larva,

Derivation Gk. lepidos–scale; pteron–a pupa, adult)
Size Wingspan 3–300 mm. Number of families About 141
Club-ended antenna
Compound eye
Wings covered
with minute, Large
overlapping scales mesothorax
Vanessa cardui — Lepidoptera
Key features
• entire body and both sides of wings usually covered with minute, overlap-
ping scales
• mouthparts usually in the form of a sucking proboscis (coiled at rest)
• some species are significant crop pests
• third-largest order
The Dusky Thorn moth,

Ennomos fuscantaria, feeds
© Anne Riley
on European ash, as well as
flowering plants and shrubs.
UK.
Many moths, such as

Speiredonia spectans from
Australia, possess distinctive
© Paul Brock
eye spots on their wings that

have evolved to startle
predators.
LEPIDOPTERA 237
Plume moths are
© Paul Brock
unmistakable due to the
unique shape of their wings.
Buckleria paludum. UK.
Members of this readily recognizable order occur everywhere that there is

vegetation. The first Lepidoptera appeared in the Jurassic, and then radiated
greatly with the rise of the flowering plants in the Cretaceous.
The higher classification of the Lepidoptera has as much a confused his-
tory as that of many other large taxa, and there have been many attempts to
divide the order into basic groups. A common theme among older classifi-
cations, and one that still pervades popular opinion, is that of a division of
the order into moths and butterflies. It is often said that moths are noctur-
nal, drably coloured, hold their wings outstretched, and lack antennal clubs,
whereas butterflies are diurnal, brightly coloured, hold their wings upright,
and have marked terminal, antennal clubs; there are many exceptions to this
rule of thumb. In fact, many moths hold their wings together in a roof-like
manner over the body at rest, and quite a few are brightly coloured and diur-
nal. Many butterflies are very drab, but at least they do all have a distinctive
antennal club. The skippers (Hesperiidae) with their odd, elongate anten-
nal club are very closely related to the true butterflies, and both belong to
the superfamily Papilionoidea, and together, they represent less than 15% of
all lepidopteran species. The Hedylidae, with a single genus (Macrosoma) of
35 neotropical species, have a range of moth-like and butterfly-like charac-
ters, and were long-thought to represent a distinct superfamily. However,
molecular data strongly support that they represent the sister-group to the
skippers plus the true butterflies. These three groups can be considered as
monophyletic.
Butterflies in tropical
climates frequently probe
the ground for nutrients
© George McGavin
such as salt and other
minerals, such as this
aggregation of male
Common Jays, Graphium
doson in Sabah, Borneo.
The beautiful colours of

butterflies, like those of this
Papilio machaon from
© Zestin Soh
Greece, have long attracted
the interest of entomologists,
amateurs, and artists alike.
‘Moths’ do not form a monophyletic group, but they are frequently referred
to as such for convenience. Lepidoptera is divided into four suborders, three
of them with only a handful of species; the fourth, the Glossata, contains the
huge majority of the species (see Box).
Lepidopteran suborders
Suborder Superfamilies Number of families:

larval habit
The odd, elongate antennal
club of hesperiid butterflies. Zeugloptera 1 1
Micropterigoidea Micropterigidae: litter, fungal grazers
·········································································································································································
Aglossata 1 1
Agathiphagoidea Agathiphagidae: seed borers
·········································································································································································
Heterobathmiina 1 1
Heterobathmioidea Heterobathmiidae: leaf miners
LEPIDOPTERA 239
Lepidopteran suborders continued
Suborder Superfamilies Number of families:

larval habit
Glossata 11 23
(non-ditrysians)* Acanthopteroctetoidea 1: leaf miners
Andesianoidea 1: borers?
Adeloidea 5: borers, gall formers, grazers, miners
Eriocranioidea 3: leaf miners
Lophocoronoidea 1: miners or borers?
Neopseustoidea 2: miners
Mnesarchaeoidea 1: grazers in litter, mosses, etc.
Hepialoidea 5: stem and root borers etc.
Nepticuloidea 2: leaf miners
Palaephatoidea 1: leaf miners and then leaf tiers
Tischerioidea 1: leaf miners
·········································································································································································
Glossata 30 114
(Ditrysia) Tineoidea 5: detritivores, fungivores herbivores, etc.
Yponomeutoidea 10: leaf miners and grazers, borers
Galacticoidea 1: leaf tiers
Gracillarioidea 3: leaf miners and grazers
Gelechioidea 21: leaf miners, gall formers, rollers,
detritivores, etc.
Simaethistoidea 1: ?
Cossoidea 7: wood borers, root feeders, stem borers
Carposinoidea 2: borers, leaf tiers, gall formers
Tortricoidea 1: borers, leaf tiers, and rollers
Sesioidea 3: wood, stem, and root borers, leaf miners,
and tiers
Zygaenoidea 12: external plant feeders, ectoparasites
Immoidea 1: external plant feeders
Schreckensteinioidea 1: external plant feeders, grazers
Urodoidea 2: external plant feeders
Choreutoidea 1: external plant feeders, grazers
Epermenioidea 1: external plant feeders
Alucitoidea 2: borers in buds, fruit, and seeds
Pterophoroidea 1: borers in buds, seeds, shoots, etc.
Hyblaeoidea 1: external plant feeders, borers
Thyridoidea 1: leaf tiers
Pyraloidea 2: leaf rollers, tiers, or gallers
Geometroidea 5: external-feeding herbivores, detritivores,
coprophages, predators
Drepanoidea 3: external plant feeders
Calliduloidea 1: external plant feeders
Papilionoidea 7: external plant feeders, leaf folders, tiers,
and borers
Lasiocampoidea 1: external plant feeders
Mimallonoidea 1: external plant feeders, saprophages,
predators
Bombycoidea 10: external plant feeders, communal web
makers, leaf tiers
Noctuoidea 6: external plant feeders, some predators
Whalleyanoidea 1: ?
* Here we have arranged all the non-ditrysian taxa together, although they are, strictly speaking,
regarded as six separate infraorders.
The Micropterigidae comprise about 180 species of small, caddisfly-like

moths whose adults have proper chewing mandibles, which they use to
feed on pollen and spores. The larvae feed on leaf litter, fungal hyphae,
or bryophytes. The two known species of the primitive Agathiphagidae oc-
cur in Queensland and Fiji, where the larvae feed on the seeds of Kauri
pines (Agathis spp.). As in the Micropterigidae, the adults have articulated
mandibles, but these are probably non-functional. About a ten species make
up the Heterobathmiidae. The larvae mine inside the leaves of Southern
Beech (Nothofagus spp.). The adults have functional mandibles and probably
feed on Nothofagus pollen.
The Glossata, species with a proboscis of varying length (sometimes very
reduced or vestigial), can be divided into two groups, the more advanced of
which forms a taxon known as the Ditrysia. In these lepidopterans, the fe-
males have two genital openings, one of which is used by the male during
copulation; the other is used for the passage of eggs during oviposition.
The most characteristic features of the order are the minute overlapping
scales or hairs that cover the entire body surface and wings, and, typically, the
possession of a long, coiled proboscis, which is used to suck liquids—usually
nectar fruit juices or honey dew. The proboscis is formed from the very long
maxillary galeae, which are deeply grooved and locked tightly together by
means of opposing rows of hooks and spines. Despite the number of people
who spend much of their lives studying these insects, the exact way in which
the proboscis is extended and coiled is not yet fully understood. Another
structure unique to most lepidopterans is the epiphysis—a spur-like structure
located on the inner face of the front tibia, used in cleaning the proboscis.
The extraordinarily rich variety of pattern and colour seen in lepidopteran
wings, must account, at least in part, for the enormous popular appeal of these
insects. The wing scales that give these insects their colours and patterns can
be pigmented or microsculptured with fine longitudinal ridges, such that they
refract the light falling on them. The brilliant and iridescent sheen seen on
the wings of many species are due to this physical colour production. It is
well known that wing colours and patterns, whether warning or cryptic, pro-
batesian mimicry
where an edible or palatable tect from attacks by predators, such as birds, and, in many species, palatable
species mimics the colours species mimic the colours and patterns of non-palatable species. As long as
and patterns of an inedible or
unpalatable species. The
the mimic is sufficiently rare, naive predators will always associate a particu-
mimics are rare relative to the lar colour pattern with non-palatability and the mimic will survive. There are
models two main types of mimicry: batesian and mullerian.
mullerian mimicry Wings are also of use in thermoregulation and courtship. Some butter-
where several inedible or
unpalatable species converge
fly wings have ultraviolet reflecting or absorbing patches, which are used in
in colour pattern. As all the signalling. In some species, males have specialized wing scales called andro-
species are non-palatable, conia that are responsible for the release of sexual odours. The head bears
mimicry rings, as they are
known, are enhanced by the well-developed compound eyes, the antennae, and, sometimes, two ocelli.
frequency of all ring members The antennae, which bear chemosensory receptors capable of responding to
and even distantly related
minute quantities of volatile compounds, such as sexual odours, are branched
species can be involved
or feathery in male moths, and simple in female moths. The thorax carries
three pairs of legs and two pairs of wings, which are mechanically coupled by
means of a frenulum and retinaculum or overlapped during flight.
LEPIDOPTERA 241
In males, the frenulum is a single stiff bristle, whereas in females it is

made up of more than one bristle. In butterflies, the base of the hind wings
is enlarged and serves to couple the wings in flight by supporting the front
wings.
Mate attraction and courtship in lepidopterans is highly variable, but typi-
cally involves displays, either singly or in groups, and the production of sexual
pheromones. In many moths, resting females release a pheromone from ab-
dominal glands. These scents are carried downwind and may be active over
very long distances. Chemosensory sensilla on the antennae of males pick
up the scent, often at very low concentrations. Males fly upwind to locate
the female, court her, and copulate. The males of some species may gather in
small groups or swarms near obvious features, such as a patch of sunlit fo-
liage, and dance to attract females. Males also produce pheromones to attract
females. In some arctiids and noctuids, males may have inflatable scent organs
called coremata, which can be everted during courtship. The scent produced
by these and other structures have several effects. They clearly act as a pow-
erful aphrodisiac, but also keep the females motionless. There are instances
where the production of sexual pheromones might be the downfall of a male
moth. The bolas spider is able to mimic the scent of female moths, and, there-
fore, able to lure males within range of its sticky snare. Acoustic signals are
used in the mating rituals of some Arctiidae, Pyralidae, and Sphingidae. In the
Lesser Wax Moth, Achroia grisella (Pyralidae), males produce ultrasonic calls
to attract mates over short distances. Louder ultrasound to attract females is
produced by males of an Australian noctuid moth (Hecatesia exultans). The
sounds carry up to 25 m, and males entering the territorial range of another
male will be chased off. Some sounds produced by adult lepidopterans are
defensive.
leading edge of front wing
retinaculum
(flap-life process)
♂
frenulum
(single composite spine)
leading edge of front wing
retinaculum
(group of stiff hairs)
♀
The wing coupling
mechanism of male and frenulum
female moths—viewed from (separate spines)
underneath where the wings
join the thorax. leading edge of hind wing
Butterflies that are diurnal rely more on visual courtship displays than
most moths. The males of many species are territorial and fly along a regular
patrol route looking for females to mate with or other males to chase away.
A good way to lure the brilliant blue males of Morpho butterflies is to whirl
a piece of metallic blue shiny paper on a weighted string around your head.
The territories held by males may be determined by the presence of the larval
food plant (and therefore oviposition site) or may be just easily recognizable
features of the landscape, such as hilltops, clearings in woods, or patches of
bare ground. The use of male pheromones, produced by hairs and scent scales
on the wings or from inflatable abdominal hair pencils, is widespread and a
variety of aerial manoeuvres performed by the males ensure that the females
are liberally covered with the ‘perfumes’.
After mating, the males of some species shower the female with a dif-
ferent scent, designed to put other males off. More commonly after mating,
especially in the Nymphalidae and Papilionidae, the male will insert a sticky
ditrysian
having two distinct sexual
secretion produced by his accessory glands into the genitalia of the female.
openings: one for mating, and This substance hardens to form a plug, which prevents other males from
the other for laying eggs mating. The eggs, of course, can still be laid because butterflies are ditrysians.
Eggs are scattered in flight, laid singly, or laid in batches near or on the
larval food. Leaf mining or boring species must burrow into the food plant.
Lepidopteran larvae, or caterpillars, are elongate and cylindrical in shape, and
may be smooth, spiny, or hairy. The toughened head has chewing mouth-
parts, very short, three-segmented antennae, and two laterally placed groups
of about six ocelli. There are three pairs of thoracic legs, each with a single
claw, and a variable number of abdominal prolegs. The abdomen typically has
ten segments, and prolegs usually occur on segments 3–6 and 10, but there
may be fewer than this number in certain families. The ends of the fleshy pro-
legs have many tiny hooks (crochets) arranged in circles or bands to help them
hold on to the foodplant. The vast majority of caterpillars are herbivorous and
often plant-specific, but there are species who eat fungi, dried organic mate-
rial, and lichens. The caterpillars of the moth Ceratophaga vastella (Tineidae)
burrow through the horns of dead animals, such as buffalo, and it is common
to find festoons of pupating moths, or their empty cocoons, adorning skulls
in the African savannahs. Nothing goes to waste in the natural world.
A few butterflies and moths have developed a carnivorous habit. The
caterpillars of certain species in moth families such as the Psychidae, Tinei-
dae, Pyralidae, and Noctuidae are predators of various types of scale insect.
The Australian moth family Cyclotornidae (Zygaenoidea) contains a strange
species, Cyclotorna monocentra, whose first instar caterpillars feed externally
on the tissues of leafhoppers. The first instar caterpillars then build them-
selves a shelter and moult to the second instar. The leafhopper colonies are
attended by ants, and interestingly, the second instar caterpillars are taken by
the ants into their nest. Here the caterpillars eat ant larvae but are not attacked
by their hosts on account of a sweet secretion they give in return. Also in the
superfamily Zygaenoidea, the Epipyropidae harbours some more odd species.
The caterpillars of small moths in the genera Epipyrops and Agamopsyche are
parasitic. About 40 species are confined to the tropical and warm temperate
LEPIDOPTERA 243
regions of the southern hemisphere. Large fulgorid bugs are the preferred
hosts, but some cicadellids and cicadas also serve as hosts for certain species.
Typically, the caterpillars are external parasites, feeding either singly or gre-
gariously in small numbers on the body fluids of their adult (and sometimes
nymphal) host bug species. Once a young caterpillar finds a suitable host, it
uses long, sharp mouthparts to penetrate the cuticle. When fully grown, the
fifth instar caterpillar detaches itself and pupates inside a cocoon. The effects
on the host are variable depending on size and the number of parasites. At
one end of the spectrum, the host may show hardly any ill effects, while at the
other end, the host may die soon after the caterpillars leave.
Even stranger are the ambush tactics employed by the caterpillars of some
Hawaiian moth species belonging to the genus Eupithecia (Geometridae).
These strange caterpillars strike in a snake-like manner at passing insect prey,
mainly small flies such as Drosophila.
Carnivorous caterpillars may come as something of a surprise, but among
the fruit-piercing moths that belong to the family Erebidae, there are species
of vampire moth. Calyptra eustrigata and some other species have made the
dietary switch from fruit juice to mammalian blood. It is interesting to note
that trainee medics use oranges to practice giving injections, as the resistance
offered by the peel is very similar to human skin.
Moths have other interesting relationships with mammals. In the family
Pyralidae, five species of moth in two closely related genera, Cryptoses and
Bradypodicola, have intimate associations with tree sloths. That the moths
lived in the fur of the sloths was known for some time, and it was assumed
that the caterpillars lived there as well, grazing on the algae growing on the
hosts’ fur. Sloths are celebrated for their lack of activity, but every week or
so they climb down to the ground to defecate. Why they do not simply drop
their dung from where they hang, high in the rain forest canopy, has more to
do with sloth social life than having an easy life. At any rate, it is at this point
that the sloth moths spring into action. Mated females leave the sloth while
it is defecating and lay their eggs on the dung. The caterpillars are, of course,
coprophagous. The female flutters back to her stately conveyance before it
disappears up the tree again, and, in time, presumably, the newly emerged
moths will find a sloth visiting the latrine.
Caterpillars may protect themselves by hiding in shelters or cases. The
caterpillars of Psychidae, which are commonly called bagworms, make tough
portable shelters from a variety of plant materials, including spines and leaf
fragments. Shelter building by some caterpillars can greatly increase their sur-
vival by allowing them access to a food source not easily utilized by other
insect herbivores. Such an approach is followed by a gelechiid moth in the
genus Polyhymno, whose larvae live inside a shelter they create by joining sep-
arate leaflets of Acacia cornigera together with silk. This plant has no chemical
defence, but instead relies entirely on patrolling ants (Pseudomyrmex ferrug-
inea) for herbivore control. Unfortunately for the tree, the ants are unable to
effectively rid it from this particular herbivore, as the latter constructs an ef-
fective and largely ant-proof shelter. In fact, Polyhymno infestations can be so
severe that they often result in the death of the tree. The case building habit is
also seen in other moth families, such as the Tineidae and Coleophoridae. A
leaf mining or boring habit might be seen as a common defence from preda-
tors, but many parasitic wasps are specialized to seek out feeding caterpillars
concealed in these ways. Parasitoids may be able to detect the movements of
leaf miners, but the probing and insertion of the wasp’s ovipositor can trigger
avoidance response in some species.
Externally feeding herbivorous caterpillars employ a range of physical and
chemical defences. They may possess a dense covering of poisonous or irritat-
ing hairs or spines, and some may be cryptically coloured, or resemble twigs
or bird droppings. Others may be brightly coloured to warn birds of the pres-
ence of toxic substances within their body, or the ability to release noxious
substances from eversible glands called osmeteria. Threat postures that dis-
play contrasting eyespots are fairly common, but some hawk moth caterpillars
have evolved patterns that are believed to resemble the heads of snakes.
After anything between four and nine larval stages (typically five), pu-
pation occurs. Lepidopteran pupae may be enveloped inside a silk cocoon
spun by the mature larva, inside an underground silk-lined cell, or suspended
in various ways from the foodplant. The term chrysalis is often used for
lepidopteran pupae. In primitive moths, the pupa has moveable mandibles
(decticous), and the appendages are not stuck to the body (exarate). In the
majority of species, however, the pupa does not have moveable mandibles
(adecticous), and the appendages are stuck to the body surface (obtect). The
pupae of the higher Lepidoptera are not totally defenceless as some have spe-
cial gin traps made of the opposable edges of adjacent abdominal segments.
These can be snapped shut by wriggling body movements to deter egg laying
by a pupal parasitoid.
In all insects air is distributed to the tissues through a system of spiracles,
tracheae, and tracheoles. Tracheoles ramify profusely throughout the tissues,
where oxygen and carbon dioxide are exchanged. However, not all tracheae
end up in tissue. The purpose of unusual tracheal tufts in the rear end of many
caterpillars’ bodies was unknown for almost a century. Now, incredibly, we
believe that they function pretty much in the same way as lungs. These thin-
walled tracheal tufts arise from the posterior pair of spiracles. While some of
these tuft branches are anchored in heart tissue, the majority of the tufts are
free to wave to and fro in the haemolymph as the heart contracts and relaxes.
In times of oxygen depletion, a certain type of haemocyte is attracted to these
tufts and becomes closely apposed to the surfaces of the thin-walled tracheae
in the tufts where they become oxygenated. Some of the tufts are concentrated
in a special compartment called the tokus at the very rear of the caterpillar’s
abdomen. When oxygenated, haemocytes leave this compartment, and they
are in exactly the right position to be picked up by the posterior ostioles of
the dorsal heart and pumped forward to the anterior end of the body. Further
studies may show that this system is common to the Lepidoptera and may
even occur in other orders.
The following section is solely concerned with some of the most
widespread ditrysian families and is not meant to imply that the early super-
families of the Glossata do not contain very interesting species. The case of
LEPIDOPTERA 245
Yucca plants and moths belonging to the non-ditrysian moth family Prodox-
idae (Adeloidea) is worthy of mention as it is a classic case of an insect–plant
mutualism. The female moths pollinate the Yucca flowers, but their larvae eat
some of the plant’s seeds.
The largest family in the Tineoidea is the Tineidae, and we have identified
about 3,000 species of these drab, usually dull-brown moths. Tineid cater-
pillars are best known for the destructive habits of some species that attack
fur, woollen, and other textiles, and dry foods, but the majority are scav-
engers and feed in fungi, decaying wood, or dry plant and animal matter.
Very few species feed on plant foliage. The caterpillars make a tunnel or web
of silk wherever they feed or construct a portable case from silk and debris.
Many species are pests, and several have a worldwide distribution due mainly
to commerce. The Webbing Clothes Moth (Tineola bisselliella), the Tapestry
Moth (Trichophaga tapetzella), and the Case Bearing Clothes Moth (Tinea pel-
lionella), are common household pest insects, whose caterpillars make holes
in clothes and carpets. Nemapogon granella is a widespread pest of stored
grain, and Niditinea fuscipunctella, the Poultry House Moth, can be found
in chicken houses, where the caterpillars feed on fragments of feathers.
Within the Yponomeutoidea, the Plutellidae is economically significant.
More than 100 species are found throughout the world, particularly in trop-
ical and subtropical regions. The family includes the diamondback moths of
the genus Plutella. Most species are cryptic, and many have bands or irreg-
ular markings. When the wings are folded, many species appear to have a
row of diamond-shaped marks along the back. The caterpillars mine, bore,
or feed externally on a range of trees, shrubs, and plants, and often make a
flimsy silk web. Many species are vegetable crop pests, especially on cabbage
species. Plutella xylostella is a very serious pest worldwide, especially since it
has developed resistance to Bacillus thuringiensis biopesticides.
More than 4,500 species of small or very small, grey or brown moths be-
long to the family Gelechiidae (Gelechioidea). These moths can be found
in a large range of habitats and the caterpillars feed externally, or inside
leaf rolls, stems, leaf mines, or galls. Many species are serious agricultural
pests and attack crops ranging from potato and peanuts to tomato and
strawberries. Coniferous and fruit trees can also suffer great damage. The An-
goumois Grain Moth (Sitotroga cerealella) is a serious, cosmopolitan pest of
corn and stored grain, and the caterpillars of the pantropical Pink Bollworm
(Pectinophora gossypiella) are major pests of cotton.
Carpenter moths (Cossidae) are medium or large, heavy-bodied moths,
which usually have spotted, irregularly patterned, or mottled wings. The
adults are nocturnal, and females lay their eggs in cracks in tree bark, or in the
emergence tunnels of adult moths. Although the larvae of a few species bore
into the pith of reeds, most attack trees. The caterpillars burrow into the wood
of the branch or trunk on which they were laid, and, as they grow, the diam-
eters of the tunnels inside the wood get bigger and bigger. Due to their large
size and poor diet, cossids can take several years to reach adulthood. The fully
grown larvae pupate within a cocoon made of silk and chewed wood fibres,
usually in their tunnels close to the surface or in the soil. Characteristically,
the empty pupal skin remains partly sticking out of the exit hole after the adult
moths have emerged. Many species are pests of commercially important trees,
such as hickory, maple, ash, oak, and pine, as well as fruit trees. The larvae of
leopard moths, Zeuzera pyrina and Zeuzera coffeae, damage and may kill fruit
and coffee trees, respectively. Cossus cossus, the European Goat Moth, whose
larvae may be up to 100 mm long when fully grown, damage fruit and the
softwood trees.
‘Witchety’ grubs, a favourite and highly nutritious delicacy of Australian
aboriginal people, are the larvae of some cossid species that bore inside the
roots and stems of certain Acacia (wijuti) species.
The Tortricoidea contains a single large family of smallish moths, the Tort-
ricidae, with more than 6,340 species worldwide. Many species are cryptically
coloured in brown, green, or grey, with patterns that resemble lichen, bark,
or even bird droppings, but some species are brightly coloured. Most species
have square-ended and broad front wings that are held roof-like over the body
at rest. Tortricid caterpillars feed externally or internally on leaves, shoots, and
buds, and many species tie or roll leaves together with silk. Some species bore
into fruit, seeds, or stems. These moths eat a huge range of plants, and some
species can be extremely polyphagous, having been recorded from almost 100
The caterpillar of the different host plants. Caterpillars of Cydia saltitans feed inside the seed of
European Goat Moth
(Cossus cossus) tunnels in Croton plants in parts of southern USA and Mexico. If too hot, fully grown
the wood of tree trunks and larvae wriggle violently inside the seed, causing it to jump, in an effort to
branches.
move into shade. These ‘Mexican jumping beans’ are sold as curiosities. About
one-third of all tortricid genera contain pest species that attack crops from av-
ocado and lucerne to all manner of trees and soft fruit crops. A well-known
and serious pest in apple orchards, the Codling Moth (Cydia pomonella), was
introduced to North America from Europe over 200 years ago.
There are more than 1,360 species of clearwings or clear-winged moths
belonging to the Sesiidae, the largest family in the Sesioidea. These day-flying
moths are brightly coloured, black, bluish, or dark brown with yellow and
orange markings, and many species mimic wasps or bees by having large ar-
eas of the wings clear of scales. The wings of many species are fully scaled
at first, but scales are shed during the adult’s first flight. Others may have no
scales from the start or may even have transparent scales. The Hymenoptera-
like appearance of these moths is accentuated by a banded abdomen and a
buzzing flight. Caterpillars bore inside the branches and roots of trees and
woody shrubs, and development of the caterpillars can take up to two years.
Sesia apiformis, the Hornet Clearwing Moth, damages the trunks and roots of
willows and poplars, and peach trees in North America suffer from damage
by caterpillars of Synanthedon excitosa, the Peach Tree Borer.
Pyralid, snout, or grass moths (Pyraloidea) make up a very large superfam-
ily of more than 15,600 species. They are very diverse varying enormously in
size, colour and shape. The wings may be broad or narrow and have closely
packed scales. In some species the front of the head has a small ‘snout’ formed
by elongate palps held outstretched. These moths are very widespread, and
the caterpillars can be herbivores or scavengers. Caterpillars of Galleria mel-
lonella, the Wax Moth (Pyralidae), even have special enzymes for digesting the
LEPIDOPTERA 247
honeycomb in bee colonies. The caterpillars of this and many other pyralid
species make silk tubes or webs on their food. A few species are predators
of scale insects or wood-boring larvae and a few feed in the roots of aquatic
plants. A very large number of Pyraloidea are pests and the list of crops and
stored products damaged by these small moths is very long indeed. The Cos-
mopolitan Meal Moth, Pyralis farinalis (Pyralidae), is a pest in stored grain
and caterpillars of the European Corn Borer, Ostrinia nubilalis (Crambidae),
damage young corn.
The Geometridae is an enormous moth family with more than 23,000
species. The wings of these moths often have patterns of fine lines and mark-
ings on a cryptic brown or green background. The bright green colour of
some of these moths is due to a unique pigment called geoverdin, which is
contained within the wing scales, and may be derived from chlorophyll. Ge-
ometrids can be found virtually everywhere there is vegetation, and the adults
are usually nocturnal. The name geometrid, literally ‘earth measurer’, is de-
rived from the characteristic looping motion of the caterpillars. Geometrid
caterpillars, which are called inchworms or loopers, have prolegs on the sixth
and tenth abdominal segments and move by drawing their hind end up to
meet the front end. They then hold on with the abdominal prolegs and extend
the head end forward and grip the substrate with the thoracic legs.
Caterpillars are protected by being cryptically coloured, and, at rest, many
will assume a very twig-like appearance, sticking out at an angle from the
host plant. The range of plants attacked by these moths is immense, and many
species are serious pests of commercial deciduous and coniferous trees, being
capable of causing severe damage and defoliation. The Winter Moth (Oper-
ophtera brumata) is a typical pest species, which was accidentally introduced
to North America from Europe. The caterpillars feed on a wide range of
deciduous trees, including apple and pear. Geometridae have a pair of well-
developed tympanal organs located at the base of the abdomen. These organs
give the moths bat-detecting ultrasonic hearing, which operates in the same
range as the echolocating signals sent out by most bats (25–45 kHz). The flight
behaviour of the moths depends on the intensity of the signal received. If the
bat is perceived as very close, the moth will perform a spiral dive towards the
ground; otherwise, the moth may just simply change direction.
A geometrid caterpillar
moves in a characteristic The Papilionoidea or true butterflies comprise more than 17,500 species in
looping manner using its seven families (including Hedylidae and Hesperiidae). The skippers (Hesperi-
thoracic legs and idae) are a group of about 3,500 diurnal species of early butterflies, which until
abdominal prolegs as
alternate anchors. recently, were considered as related to, but outside, Papilionoidea. In general,
they are moth-like and heavy-bodied with a wide head. Most species are drab
brown with white or orange markings, but a few are pale with a blue, green,
or purple sheen. The antennae terminate characteristically in an elongated
club, which comes to a point and is often curved. The name ‘skipper’ refers
to their rapid, darting flight pattern. The caterpillars, which are green, brown,
or white, often with longitudinal stripes, mainly eat grasses and sedges, and
the foliage of some herbaceous plants and trees. They usually feed at night
and live within a shelter of silk-tied or rolled leaves during the day. A few
grass-feeding species can be minor pests, as can Urbanus proteus, the Bean
Leaf Roller, which feeds on leguminous and cruciferous plants, and the Rice
Leaf Roller (Lerodea eufala), which can attack sorghum and sugar cane.
Papilionids are often large and can be spectacularly attractive. Most of the
580 or so world species are swallowtails (Papilioninae), with short or long
tails directed backwards from the ends of the hind wings. There are about 50
species of apollos (Parnassiinae), which do not have hind wing tails, and are
white or grey in colour often with two red or yellow spots on the hind wings.
Swallowtails are widespread, but apollos live in mountainous regions of the
northern hemisphere. A characteristic feature of the caterpillars is a forked
thoracic scent gland called an osmeterium. When threatened, the caterpillar
everts the brightly coloured osmeterium, which gives off a repellent odour.
When fully grown, swallowtail caterpillars pupate on their host plant and
the chrysalis is held upright by a silk belt around the thorax. Apollos pupate
in plant litter or under stones surrounded by a loosely woven cocoon. These
butterflies are of great aesthetic value, and the family contains the spectacu-
lar birdwing butterflies of south-east Asia. Ornithoptera alexandrae, from the
south-east of Papua New Guinea, is the biggest butterfly in the world with
A characteristic feature of females measuring 28 cm (11 in) across the wings.
the Swallowtail Butterfly The Pieridae are among the commonest butterflies anywhere in the world.
caterpillars is a forked
The common names, whites, sulphurs, and orange tips, refer to the wings of
thoracic scent gland called
an osmeterium—here these butterflies, which are usually white or yellow, with orange, black, or dark
everted in defensive grey markings. Pierids can often be seen feeding in groups at bird droppings,
posture.
puddles, or wet patches on the ground. It is thought that this ‘puddling’ be-
haviour, which is common in males, provides dietary salts. Caterpillars of
the best-known species feed on cabbage species and leguminous plants. The
chrysalis, which is angular with a single head projection, is fixed upright to
the host plant and held in place by a silk girdle around the abdomen. Many
species are serious agricultural pests. The Cabbage White or Large White
(Pieris brassicae) is a notorious pest of cabbage throughout Europe and the
Mediterranean, and the Small White (Pieris rapae), which also attacks wild
and cultivated cabbage, is perhaps the most destructive of all butterflies.
More than one-third of all butterflies belong to the family Nymphalidae,
which is composed of several distinctive subfamilies with a total of about
5,700 species. Nymphalids are often called brush-foot butterflies because the
front legs are very small and brush-like, and not used for walking. A good
identification feature, therefore, is the presence of only four walking legs. The
front legs have a sensory function, and, in females, at least, they are probably
useful in host plant recognition. Nymphalid caterpillars are generally spiny
and have many peculiar, branched projections, or have horns at either end of
the body. They feed externally on the foliage of a huge range of host plants.
The angular chrysalis is suspended, head down, from the host plant by a small
group of terminal hooks called the cremaster. The spectacular owl butterflies
(Brassolinae), the brilliantly metallic blue Morpho butterflies (Morphinae),
and the passion vine-eating heliconiids (Heliconiinae) live in Central and
South America. Browns and ringlets (Satyrinae) are pale yellowish-brown to
dark brown or greyish coloured butterflies, which often have numerous eye-
spots around the wing margins. These butterflies are characteristic of heaths
LEPIDOPTERA 249
and meadows, particularly in upland areas, and adults fly in an erratic and
bobbing manner. Satyrine caterpillars all feed on grasses or sedges and can be
easily recognized by their forked tail segment. The subfamily Nymphalinae is
the biggest in the family with more than 3,000 species. A well-known species,
the Painted Lady (Vanessa cardui), is migratory and can be found all over the
world. The caterpillars of this species usually feed on nettles and thistles, but
it can occasionally be a minor pest of soya bean. The milkweed butterflies be-
long to the Danainae, and the best-known species are conspicuously coloured
blackish-brown and reddish-orange with small white marks. The American
Monarch Butterfly (Danaus plexippus) is typical. Adults and caterpillars are
highly distasteful to predators on account of the toxic, cardiac glycoside com-
pounds they sequester from their Asclepias host plants (Asclepiadaceae). The
colouration of the adults and the yellow and black bands of the caterpillars
warn predators of their non-palatability. The Monarch is a famous migra-
tory species that has established itself all over the world. In North America
it migrates from Canada to California and Mexico, where it overwinters in
spectacular mass roosts.
More than 6,000 species of blue, copper, or hairstreak butterfly belong to
the family Lycaenidae. Most species are slender-bodied and have a tropical or
warm temperate distribution. Some species are very brilliantly coloured with
iridescent scales, and others have slender tails on the hind wings. The caterpil-
lars feed on the leaves, fruits, and flowers of a wide range of plants, and some
species are predacious on aphids, coccids, and other small soft-bodied insects.
Nearly one-third of the known larval life histories show complex, mutualistic
associations with ant species. Lycaenid caterpillars are squat, and from above
they look like a small slug or woodlouse. Caterpillars associated with ants
secrete a special fluid containing sugars and amino acids from abdominal
‘honey glands’. The ants allow the caterpillars to eat their larvae, or some-
times the aphids from which they harvest honeydew, in return for access to
the honey gland fluid. The ants will even guard these caterpillars, and there is
good evidence that the ants prevent parasitic wasps from attacking them as
well.
The Lasiocampoidea comprise more than 1,500 species of eggar moths
(Lasiocampidae), although lappet moths (Anthelidae) are sometimes in-
cluded in this superfamily. Lasiocampids are common in wooded areas,
hedgerows, and heathlands, where their host trees, grasses, or herbaceous
plants grow. The caterpillars are often very hairy or have dorsal and lateral
tufts and downward-pointing hair fringes. The caterpillars of some species
live and feed communally inside tents or webs, which they spin across the
foliage of their food plant. Pupation takes place inside a tough, papery, egg-
shaped cocoon. Many lasiocampids, such as the tent caterpillars of the genus
Malacosoma, are serious forest and orchard pests.
Moths belonging to the large superfamily Bombycoidea are stout-bodied
and have broad wings. Although the silk of lasiocampid cocoons has been
used by humans in the past, the main commercial silk-producing species be-
long to the Bombycidae. There are about 60 species of silk moths, the best
known of which is the Silkworm, Bombyx mori. Silk production originated in
China more than 4,500 years ago, and the Chinese fiercely guarded their valu-
able secret (silk was worth more than gold) until ad 555, when two Roman
monks smuggled some moth eggs and the seeds of the mulberry plant upon
which the caterpillars fed to Constantinople and other parts of the Eastern
Roman (Byzantine) Empire. The caterpillars are kept in large, airy, rearing
drawers, and eat huge amounts of leaves in the course of their development.
When fully grown, the caterpillars spin a cocoon in which to pupate. These
are collected and boiled, and the silk is wound off onto reels (the hard bit is to
find the end of the thread). A single cocoon may provide hundreds of yards of
silk, but the silk of several cocoons has to be twisted together to make a single
silk thread. Up to 2,000 cocoons may be needed to provide enough silk for a
dress. Nothing is wasted as the boiled pupae are canned and sold as food.
The Saturniidae (atlas, emperor, and royal moths) are very large, heavy-
bodied moths with very broad wings, which may span 200 mm or more. The
wings of most of the 1,500 species have a characteristic eyespot near the cen-
tre and transparent patches, and hind wing ‘tails’ also occur in some species.
The functional significance of the long hind wing tails of some species had
A caterpillar of the Adonis long been debated. They are clearly important, as they have evolved multiple
Blue Butterfly (Lysandra
times independently, but what do they actually do? Saturniids fly at night,
bellargus) is attended by
ants, which feed on the during the peak of bat activity, but they are at first glance defenceless against
sweet secretions from the their attacks. Research has shown that these tails act as acoustic decoys to
caterpillar’s ‘honey gland’.
echolocating bats. The tails do not contain vital organs, nor are they crucial
for flight. When in flight, these tails spin in a particular fashion, which lures
bats to attack the tails and not the main body of the moth. In this way, moths
survive more than half of bat attacks, and it is not infrequent to find saturniids
with one or both tails missing.
Most adult saturniids inhabit wooded areas, and the caterpillars feed on a
wide range of deciduous and coniferous trees, and shrubs. The body surface of
caterpillars has fleshy protuberances called scoli, which bear spines and long
hairs. Pupation takes place inside a very dense silk cocoon attached to the host
plant. The biggest moths in the world belong to the genus Attacus, species of
which can be found from South America to Mexico, and from Africa to the
Orient. Samia cynthia, the Ailanthus Silk Moth or Eri Silk Moth, has been
reared for the commercial production of silk. The green caterpillars of the
spectacular pale green, long-tailed American Luna Moth (Actias luna) feed
on deciduous trees, such as birch, hickory, and walnut.
Another charismatic family of the Bombycoidea are the hawk moths (Sph-
ingidae), with just over 1,200 species. Hawk moths are a medium to quite
large, heavy-bodied, powerful species with long, narrowish front wings. The
proboscis, which is long, and in some species, several times longer than the
whole of the body, is curled under the head when not in use. Some species
(Hemaris and Cephonodes spp.) resemble bees with large transparent areas
on the wings, and others (Macroglossum spp.) look like hummingbirds as
they hover at flowers. Adults feed on nectar, which is sucked from the corolla
tubes of flowers using a long proboscis. Hawk moth caterpillars, which can
be brightly coloured or cryptic, have a characteristic dorsal spine-like process
LEPIDOPTERA 251
at the end of the abdomen. Most species pupate and overwinter in weak co-
coons. The pupal stage of some species has a peculiar handle-shaped structure
at the anterior end, which contains the proboscis. The Death’s-head Hawk
Moth, Acherontia atropos, is native to Africa, but migrates to Europe dur-
ing the summer. The caterpillars eat potato and other related plants, and the
adults raid bee colonies for honey. They make high-pitched piping noises
similar to that made by queen bees, and it is thought that this calms the
bees. Xanthopan morgani has an incredibly long proboscis reaching more
than 25 cm (10 in) to feed from the very long nectaries of certain African
orchids. Some hawk moths are serious crop pests. Manduca quinquemac-
ulata, the Tomato Hornworm of North America, and Manduca sexta, the
Tobacco Hornworm, damage tobacco, tomato, potato, and other plants in
North America and elsewhere. The Striped Hawk Moth (Hyles lineata) is cos-
mopolitan, and its caterpillars attack a wide range of commercially important
plants.
The Noctuoidea is the largest lepidopteran superfamily with well over
70,000 species. The largest family are the Erebidae, which after major re-
classification based on molecular data now include 18 subfamilies which were
previously considered distinct families, such as: the Lymantriinae (tussock
moths), the Arctiinae (tiger and ermine moths), and the Herminiinae (lit-
ter moths), among others. Many of the erebid subfamilies were also part of
the Noctuidae, the family which gives Noctuoidea its name. It is likely that
The larva of a typical hawk the classification of this megadiverse lineage will change several times in the
moth, Sphinx ligustri, future, until multiple data types lead to a more stable phylogenetic system.
showing the characteristic
terminal horn.
Female tussock moths are weak fliers, and usually lay their eggs in quite
large clumps on the bark of a great variety of host trees and shrubs, and
often protect the mass by depositing irritant hairs from the end of their ab-
domens. The remarkably hairy and mostly brightly coloured caterpillars are
phytophagous and feed externally, and often gregariously, on the foliage of
their host plant. Most species have shaving brush-like tufts of hairs on the
dorsal and lateral body surfaces. As well as thoracic legs, prolegs can be seen
on five of the abdominal segments. The Gypsy Moth (Lymantria dispar) was
introduced to North America in 1869, with the idea that its cocoons would
yield silk. As has happened so many times with such introductions, it was a
terrible idea as the moths escaped, and since then, have spread to cover large
areas of the USA and Canada. In the early 1980s, this species defoliated an
estimated 10 million acres of trees in eastern United States. The caterpillars
of another pest, the Brown Tail Moth (Euproctis chrysorrhoea), feed commu-
nally inside silk webs on hawthorn and blackthorn trees. Humans can get
incredible skin rashes from coming in contact with lymantriids, especially
the caterpillars.
More than 11,000 species of tiger and ermine moths belong to the
Arctiinae (formerly Arctiidae). The adults and their hairy caterpillars are
poisonous, as many species feed on plants, such as potato, ragwort, and
laburnum, which contain cardiac glycosides and other toxic substances. In
addition to being distasteful, the adults have thoracic sound-producing or-
gans, which emit trains of ultrasonic clicks that jam bat echolocation systems,
enabling them to escape. The caterpillars, which are often as brightly coloured
as the adults, feed at night and hide during the hours of daylight. Many
species are polyphagous and normally feed on a wide range of low-growing
herbaceous plants, although some species eat the foliage of deciduous and
coniferous trees.
The Noctuidae are one of the largest lepidopteran families with more than
12,000 species, although this number may change with future reclassifica-
tions. Noctuid caterpillars, called cutworms, armyworms, and loopers, are
among the world’s most devastating agricultural pests, and affect a large range
of crops, vegetables, and other important plants. Cutworms (Noctuinae, es-
pecially Agrotis spp.) generally chew through the base of the stem and kill the
entire plant, and even a relatively small number of caterpillars can completely
destroy a whole stand of corn or cotton.
Many species are polyphagous, for example, the Old World Bollworm (He-
licoverpa armigera) damages cotton, maize, and tomatoes. The huge list of
Cutworms generally chew
names, which includes the Clover Cutworm, the Soybean Looper, the Corn
through the base of the Earworm, the Alfalfa Looper, the Celery Looper, the Tobacco Budworm, the
stem and kill the entire Wheat Armyworm, the Lawn Armyworm, the Cotton Bollworm, and the Flax
plant.
Bollworm, should convince anyone that nothing worth growing is going to be
immune from these moths.
Phylogenomic studies estimate that the origins of Lepidoptera go back to
the Late Carboniferous (about 300 mya), although the order started greatly
diversifying much later, in the Middle Triassic (241 mya), when the evolu-
tion of the proboscis allowed lepidopterans to exploit nectar during the rise
of the flowering plants (Angiosperms). The ancestor of true butterflies (Pa-
pilionoidea) was likely nocturnal (like most moths), but later descendants
switched and became diurnal, most likely to exploit the abundant nectar of
flowering plants that was available during daytime.
The oldest fossils of the order come in the form of lepidopteran wing scales,
and date to the Jurassic (about 200 mya). They are 100 million years younger
than the phylogenomic estimate of the origin of the order mentioned above,
but this is likely a consequence of the scarcity of early Lepidoptera in the fossil
record.
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255
Hymenoptera
(bees, wasps, and ants)
Common name Sawflies, wasps, bees, and Metamorphosis Complete (egg, larva,
ants pupa, adult)
Derivation Gk. hymen–membrane; Distribution Worldwide except
pteron–a wing Antarctica
Size Body length 0.25–70 mm Number of families About 102
Known world species about 150,000 (13.25%)
Antenna
Well developed
Ocelli compound eye
Wings joined
in flight by tiny
hooks
Hind wings Waist-like
smaller than constriction
front wings (not in Symphyta)
Vespula germanica — Hemiptera
Key features
• body usually with a constricted waist
• some species live in social colonies
• ovipositor may be modified as a sting
• second largest order
A female braconid wasp

(likely Diamblomera sp.)
probing a tree trunk with her
ovipositor. Once she detects
© Nicky Bay
an insect larva, she will use
her ovipositor to pierce it
and insert an egg. Singapore.
Many bees will rest on

© Zestin Soh
vegetation at night, as shown

by this aggregation of Nomia
strigata from Singapore.
HYMENOPTERA 257
Ants are the dominant

predators in many
ecosystems, both in terms of
numbers and biomass. Here,
a group of Oecophylla ants
are dismembering an
© Zestin Soh
unfortunate Tetragonula
atripes stingless bee.
Malaysia.
Species within the Hymenoptera exhibit an incredible diversity of lifestyles:

solitary or social; herbivorous, carnivorous, or parasitic. The number of un-
described species may eventually be many times that already described (up
to more than one million), and the order may be as big or bigger than the
Coleoptera. If Diptera represents the order that has the most harmful effects
on humans, then Hymenoptera must be regarded as the most beneficial. It
would be difficult to picture a world without the strong control of natural in-
sect populations exerted by parasitic and predatory wasp species, or without
the flowering plants pollinated mainly by bees.
A nest of Apoica pallens

© George McGavin
wasps from Costa Rica. This

species is highly unusual
among wasps in that it
forages almost exclusively at
night.
© Rupert Soskin
Closeup of the European
social wasp, Polistes nimpha,
drinking water. France.
The order is divided into two suborders: the Symphyta (sawflies) and the
Apocrita (wasps, ants, and bees) (see Box). While Apocrita are monophyletic,
the Symphyta are paraphyletic, and are therefore not a homogeneous ‘natural’
group. One group of sawflies, the Orussoidea, are parasitoids, and represent
the sister group of Apocrita. Consequently, they are ranked together as the
Vespina (Apocrita + Orussoidea) (see Box).
Although Apocrita are overall monophyletic, its megadiverse parasitoid
lineages were once lumped in a now-obsolete infraorder known as the Para-
sitica. However, molecular and morphological studies have shown that this
lineage is paraphyletic, and Parasitica are now split into many different
superfamilies that largely represent monophyletic natural groups (see Box).
Another rank in the Apocrita is the Aculeata, which includes a mono-
phyletic group of many related superfamilies that possess an ovipositor
modified into a venom-delivering stinger. Three superfamilies of aculeate
Apocrita—the Formicoidea (ants), the Vespoidea (social wasps), and the
Apoidea (bees)—are considered, at least in behavioural terms, to be the most
advanced of all insects.
Sawflies do not have a distinctly constricted waist, and females have a
saw-like ovipositor for laying eggs into plant tissue. In the Apocrita, the first
segment of the abdomen, called the propodeum, is fused to the thorax. The
combined structure is called a mesosoma (in bees) or sometimes alitrunk (in
ants). The second (and sometimes the third) abdominal segments are very
HYMENOPTERA 259
Symphyta narrow and form the petiole, which gives the distinctive wasp-waisted ap-
pearance. The swollen remainder of the abdomen behind the petiole is called
the gaster or metasoma. Parasitic apocritans have a slender, and sometimes
very elongate, ovipositor for penetrating and laying eggs inside other insects,
and aculeate apocritans have a modified ovipositor in the form of a sting with
an associated poison gland for defence. The presence of a waist in apocritans
greatly increases the manoeuvrability of the abdomen and allows the sting or
ovipositor to be brought into almost any position underneath the body, or
Apocrita even pointing forwards.
Hymenopteran superfamilies
Suborder Superfamilies Major larval habits

(no. of families)
‘Symphyta’ Xyeloidea (1) Phytophagous on coniferous and
(Sawflies) deciduous trees, some gall-formers
Xiphydrioidea (1) Borers on dead wood
Pamphilioidea (2) Phytophagous on coniferous and other
vegetation, leaf rollers
Cephoidea (1) Borers in grass and pithy stemmed
Parasitica plants
Siricoidea (2) Mostly wood-boring, some may be
parasitic
Tenthredinoidea (7) Phytophagous on trees, herbs, grasses
·········································································································································································
Vespina Orussoidea (1) Parasitoids of wood-boring beetles and
(Apocrita + sawflies
Orussoidea)
·········································································································································································
Apocrita (Wasps, Stephanoidea (1) Parasitoids of wood-boring beetles and
Aculeata ants and bees) sawflies
Trigonaloidea (1) Mostly hyperparasitoids of tachinids
The order Hymenoptera is and ichneumonids
·········································································································································································
divided into two suborders:
the Symphyta (sawflies) and ‘Parasitica’ Megalyroidea (1) Parasitoids of wood-boring beetles
the Apocrita (Parasitica— Ceraphronoidea (2) Parasitoids
parasitic wasps, and Evanioidea (3) Parasitoids
Aculeata—wasps, ants, and Ichneumonoidea (3) Parasitoids
bees). Although Symphyta Proctotrupoidea (7) Parasitoids
and Parasitica are
Diaprioidea (5) Parasitoids
paraphyletic, and therefore
unnatural groupings, they Platygastroidea (3) Parasitoids
are mentioned here due to Cynipoidea (5) Gall formers or inquilines in galls, some
their abundance in early parasitoids
classification schemes and Chalcidoidea (22) Parasitoids
studies of Hymenoptera. Chrysidoidea (7) Mostly parasitoids
Vespoidea (2) Parasitoids, or feed on provisioned
insect or spider prey
·········································································································································································
Aculeata* Pompiloidea (4) Parasitoids, kleptoparasites, or feed on
provisioned insect or spider prey
Thynnoidea (2) Parasitoids
Tiphioidea (2) Parasitoids, or feed on provisioned
insect prey
Continued
Hymenopteran superfamilies continued
Suborder Superfamilies Major larval habits

(no. of families)
Scolioidea (1) Feed on provisioned insect prey
Formicoidea (1) Larvae feed on plant, fungal and animal
material
Apoidea (17) Larvae feed on nectar and pollen,
provisioned insects and spiders,
* The Aculeata are the sting-bearing apocritans.
The head carries a pair of thread-like, multi-segmented antennae of various

designs, a pair of well-developed compound eyes, and, usually, three ocelli,
which are positioned in a triangular array on top of the head between the eyes.
The mouthparts are adapted for chewing and biting, but, in many species,
liquids are ingested. In the bees (superfamily Apoidea), the maxillae and the
labium are extended and modified to form a tongue of variable length through
which nectar is sucked. The mandibles are versatile tools used for a variety of
purposes, such as feeding, holding and moving prey, defence, and nest con-
struction. In parasitic species, they are used to cut the adult free of the body
of the host, and in other species, to leave the pupal cell.
Although many species lack wings, most members of the order are strong
fliers with an enlarged meso- and meta-thorax, and two pairs of membranous
wings, which are strongly joined in flight by a row of small hook-like structures
called hamuli, located on the front edge of the hind wing. The front wings typ-
ically have a dark pterostigma on the front edge and are slightly bigger than
the hind wings. Both pairs have a reduced pattern of venation marking out
a number of large cells, but, in many species, particularly in the superfami-
lies Chalcidoidea and Proctotrupoidea, venation is almost lacking. Legs are
generally not modified, except in Dryinidae (Chysidoidea), where the front
legs of females are raptorial to grasp hosts, and in worker bees where they
are modified for pollen gathering. Many male bees have secondary sexual
modifications of the legs, some of them bizarre.
Hymenopteran larvae are either caterpillar-like (Symphyta) or grub-like
(Apocrita). The larvae of sawflies have a well-defined head capsule, three pairs
of thoracic legs, and abdominal prolegs, although prolegs are lost in species
that bore into plant tissue. They can be separated from the caterpillars of
moths and butterflies by a number of features. The prolegs do not have cro-
chets, the eyes are simple with only one facet as opposed to several, and the
antenna are reduced to small bumps, not three segmented as in lepidopteran
caterpillars. Apocritan larvae tend to be simple and maggot- or grub-like with
no legs and a reduced head capsule. The reasons for this dichotomy are re-
lated to the lifestyle of the larvae. Sawfly larvae are active herbivores that
need to find food, whereas apocritan larvae develop from eggs laid inside the
body of a host insect, or inside a cell that is already stocked or provisioned
with food as the larva grows (only social hymenopterans practise progressive
provisioning).
HYMENOPTERA 261
parasitoid Parasitoids are insects with more than a certain amount of gruesome ap-
a species that develops by
peal. Most species of parasitoid are hymenopterans, the rest are tachinid,
consuming the body tissues of
a single host, eventually conopid, and some other flies. Parasitoids live and develop at the expense of
causing its death another organisms. They are not parasites in the usual meaning of the word,
in that the host never survives a parasitoid attack. It is an incredibly successful
lifestyle, and one that, unseen, regulates natural populations of other species
more effectively than any other single factor. Very many species are used in
biological control programmes against pest insects.
Parasitoids can feed inside the host or outside the host’s body, in which
case they are termed endoparasitoids and ectoparasitoids, respectively. An-
other difference in parasitoid habits is important to bear in mind. If the female
wasp kills or paralyses its host when laying an egg, and thus prevents the host
from developing any further, it is called an idiobiont parasitoid. The larvae of
idiobiont species are often ectoparasitoids. If, however, the female does not
kill the host when egg laying, thus allowing the host to develop further, it
is known as a koinobiont. Koinobionts tend to be endoparasitoids. In some
cases, the development of the parasitoid can be delayed until the host is nearly
fully grown. Species that are parasitoids of other parasitoids are called hyper-
parasitoids. Host-finding behaviour in parasitoids can involve a large number
of factors. Many parasitoids seek hosts in specific locations, such as inside leaf
mines or in rotten wood. The host itself may give out a chemical or vibrational
signal of some sort that is intercepted by the parasitoid, but, in some cases,
the signal may be produced by the parents of the host. For instance, Lep-
topilina heterotoma (Eucoilidae), a parasitoid of Drosophila larvae, uses the
pheromones produced by mating adults at oviposition sites as a host-finding
cue. Others, such as many members of the Dryinidae, locate their hemipteran
hosts by listening to their vibrational songs.
Most female parasitoids laying a single egg in a host will try to ensure that
the host is large enough to provide her offspring with sufficient resources to
complete its development. In other species, many eggs may be laid, or many
offspring will result from the repeated division of a single egg (polyembry-
ony). In some parasitoids, the eggs are laid outside the host and the first
instar larvae, which can move by wriggling, make contact themselves. The
differences between sawfly larvae and those of Apocrita extend to the inter-
nal organs as well. In sawflies, the gut is normal, whereas in most apocritans
the midgut is not joined to the hindgut until the last larval stage. Rather than
foul its cell or the host’s body, the larva stores its waste until just before it
pupates. Parasitoid wasps locate hosts and lay eggs in them wherever they
happen to be; inside wood, leaf mines, stems, or exposed. Many aculeate Hy-
menoptera (who have a sting) act as straightforward predators that catch and
kill prey items to feed to their larvae, but, in many species, prey items are
paralysed, and the developing larvae might be considered as parasitoids. The
difference here is that, while some spider-hunting wasps do look for hosts in
their own burrows, many hunt and transport the host either to a place where
it is interred and an egg laid on it, or to an already-prepared chamber. Con-
cealment is necessary to prevent the contents being raided or taken over by
another wasp.
Sex determination in Hymenoptera is by haplodiploidy. Males, produced

parthenogenetically from unfertilized eggs, are haploid, whereas females,
produced from fertilized eggs, are diploid. Inseminated females can control
the sex of their offspring by withholding sperm, which, in some species, can
result in very biased sex ratios. There are many species in which males are rare
or even unknown, and diploid females are produced parthenogenetically.
Herbivory is certainly the primitive and ancestral feeding habit of the or-
der, and parasitism and carnivory may have arisen via inquiline species that
may originally have been herbivores inside gall or other plant tissue but pro-
gressed to eating other small herbivores. With the exception of larvae of the
Orussidae, which are parasitoids of wood-boring beetles, sawfly larvae are
herbivorous on a wide range of plants. The Cephoidea contains a single fam-
ily, the Cephidae (stem sawflies), with about 160 species. Stem sawflies are
generally slim and cylindrical, and many species have yellow abdominal and
thoracic markings. The larvae of all cephids are stem or twig borers of grasses,
willows, raspberries, and other shrubs. Species in the genera Cephus and Tra-
chelus can be pests in cereal crops, such as wheat, oats, rye, and barley. Cephus
pygmaeus, the European Wheat Stem Sawfly was introduced to North Amer-
ica in the late 1800s and has become a pest in addition to the indigenous
species Cephus cinctus.
In the superfamily Siricoidea, about 150 species of horntails or wood
wasps make up the Siricidae. Siricids are large and stout and often strikingly
coloured—black or metallic blue, or with yellow, hornet-like markings. The
end of the abdomen has a distinctive terminal spine, which is short in males
and spear-like in females. Females have a long, very strong ovipositor, which
projects below the terminal spine. The ovipositor is used to drill through bark
into the wood of diseased or dead trees where a single egg is laid at a time.
There are two distinct subfamilies: the Siricinae, typified by species of the
genus Sirex that attack pines or firs, and Tremicinae, for example, Tremex
spp., that attack a range of deciduous trees. Siricid larvae bore into heart-
wood and eat the hyphae of wood-rotting fungus, which is introduced to the
timber along with the egg at the time of oviposition. The larvae pupate in
a cocoon made of silk and chewed wood just under the surface. Depending
on the species, there can be between five and eleven moults, which can take
two years or more. Sometimes, wood wasps have emerged from the timbers of
new houses or furniture. In one instance, the almost simultaneous emergence
of many large North American horntails from the wood behind the plaster of
a new house in Oxford caused such alarm that the owners nearly moved out.
The Tenthredinoidea is the biggest sawfly superfamily containing more
than 8,400 species, representing more than 85% of all sawfly diversity. Two
families—the common sawflies (Tenthredinidae) and the conifer sawflies
(Diprionidae)—contain significant numbers of pest species. Diprionids are
stout, mainly dull brown or black species. They are distinguished by feathery
or toothed antennae and a very broad abdomen. Larvae of the majority of
species attack pine trees, but a few species are found on cedars, hemlock, firs,
and spruce. The larvae feed externally on the needles and are often gregari-
ous and warningly coloured when young. When attacked, gregarious larvae
HYMENOPTERA 263
all simultaneously jerk their bodies upright, and have the ability to exude
distasteful resins, which they have sequestered from their food. When fully
grown, larvae pupate in a tough cocoon in the soil or glued to the host plant.
Species of Diprion and Neodiprion can cause serious damage and defoliation
in coniferous plantations. The Tenthredinidae, with more than 7,500 species,
are variable in appearance and can be found in terrestrial habitats all over the
world. Adult tenthredinids feed on pollen and nectar but many prey on other
insects. Females lay their eggs in the tissues of the appropriate host plant, and
the larvae usually chew or graze the foliage externally. Some species mine
leaves, while others, such as species of Pontania, induce galls on the leaves
of willow species. Many species are minor pests of garden plants, crops, and
trees, but some can cause significant economic damage.
The Stephanoidea contains a single family (Stephanidae) of about 350
species whose larvae are parasitoids of wood-boring beetles and horntails liv-
ing in dead wood. The adults locate their hosts by the sounds they make and
reach them using a very long ovipositor. The Megalyroidea also contains a
single family of about 50 species (Megalyridae), whose larvae, like stephanids,
are parasitoids of the larvae of wood-boring beetles.
The superfamily Trigonaloidea contains a single family (Trigonalidae or
Trigonalyidae) of around 90 species, in which life cycles can be very unusual.
The females lay a great number of small eggs into the edges of leaves where
they can be easily eaten by herbivorous insect larvae. A sawfly larva might be-
come the host to one of these parasitoids, but, if consumed by a lepidopteran
caterpillar, the egg hatches and the larva makes its way into the haemocoel
of the host. Here it will wait until the host is parasitized again, this time by
a tachinid fly or an ichneumonid wasp. The trigonalid larvae will then be-
come a hyperparasitoid. Even if an infected caterpillar is seized by a vespid
wasp, masticated, and then fed to the wasp’s larvae, the trigonalid egg will
still survive, hatch inside, and become a parasitoid of the wasp larva.
The Ichneumonoidea is dominated by two huge families, the Ichneu-
monidae and the Braconidae, with around 48,000 species collectively, and a
tiny family, Trachypetidae, with just seven species. The Ichneumonoidea rep-
resents 33% of all Hymenoptera and 3% of all complex animal life, and there
are likely as many, if not more, undescribed species. It is difficult to gener-
alize about taxa this large, but the families differ in a number of respects.
Most species in the family Ichneumonidae are ecto- and endoparasitoids
of the larvae and pupae of holometabolous insects, although a few attack
spiders and their eggs. The insects commonly used as hosts are sawflies,
moths, beetles, and flies. Many ichneumonids can be hyperparasitoids, ei-
ther of other ichneumonids or of tachinid flies. On the whole, the Braconidae
tend to be parasitoids of hemimetabolous insects, such as bugs (including
aphids), barklice, and termites, although a few do attack wasps and beetles.
Nothing is known about the biology of Trachypetidae. A fascinating aspect
of the life history of many Ichneumonoidea is their symbiosis with certain
viruses known as polydnaviruses (PDVs). These viruses are integrated into
the ichneumonoid genome, and they replicate in large numbers in the female
reproductive tract. Each ichneumonoid species has its own unique type of
PDV. When a female ichneumonoid oviposits inside an insect, she also in-
jects huge numbers of PDVs, which are essential for successful parasitization,
as they interfere with the host’s immune response, granting the parasitoid
larvae with a greater chance of survival. In this way, ichneumonoids have do-
mesticated PDVs, in an insect–virus symbiosis, which is so far unique among
arthropods (but may prove to be more common in the future).
Around 500 species belong to the seven families that make up the Proc-
totrupoidea. Most of the species in the Protoctrupidae are endoparasitoids
of beetle larvae. The related Diapriidae (superfamily Diaprioidea) are en-
doparasitoids of larval or pupal Diptera. About 4,000 species belong to the
A mummified aphid stuck to
foliage by a silk tent three families that make up the Platygastroidea. These wasps are usually
containing the cocoon of a endoparasitoids of insect and spider eggs.
pupating braconid wasp.
With the exception of some tenthredinid species, hymenopteran gall for-
mers overwhelmingly belong to the family Cynipidae, the biggest of five in
the Cynipoidea. Species in the other families are parasitoids of wood wasps
(a)
and flies. It is likely that the gall-forming habit has evolved from an ancestral
condition, perhaps a parasitoid of wood-boring larvae. Gall-forming species
belong to the Cynipinae and species of the vast majority (Andricus, Biorhiza,
Callirhytis, Cynips, Neuroterus, etc.) use oaks (Quercus) and related trees as
host plants. A few gall-forming species (Diplolepis spp.) use members of the
Rosaceae and Compositae as host plants. A few cynipids are not gall-formers
themselves but live as inquilines inside galls of other species. Plant galls are
(b)
a common sight and vary enormously in size, colour, texture, and location.
Many are highly species-specific, and it is easier to identify the gall-causing
species from the appearance of its gall than from its causer. Adult female gall
wasps lay their eggs inside plant tissues, and the host plant responds by devel-
oping a gall around it. The mechanism of gall induction in the Cynipidae has
remained elusive. The plant may isolate the gall wasp by growing a gall, but
its structure can be complex, and as it is protective and has a nutritive layer to
feed the developing larva or larvae, it is always beneficial to the gall former.
Gall formers can have complicated life cycles involving sexual and asexual
generations occurring either on different parts of the plant, or on two dif-
ferent host plants. When alternation of generations occurs, galls containing
the developing asexual generations (all females) are larger and occur during
the late summer and autumn. The females that emerge from these galls in the
following spring lay eggs that give rise to galls containing males and females.
These emerge, mate, and produce the next asexual generation. Gall wasps and
Gall-formers can have their galls often support a rich and varied community of interdependent or-
complicated life cycles
involving sexual and ganisms, many of which are parasitoid wasps that attack the gall formers, their
asexual generations. (a) The inquilines, and each other. Even when the galls are empty, they may provide
winged sexual form of
Biorhiza pallida having
shelter for other insects.
emerged from its oak apple The Chalcidoidea comprises 22 families and more than 22,500 described
gall. (b) The oak apple gall species of small or very small parasitic wasp. They are a megadiverse lineage,
caused by oviposition of
many eggs into a bud by the and estimates suggest that up to 500,000 remain to be described. Some chalci-
wingless asexual form that doids are herbivores and are seed feeders or gall formers, but the vast majority
emerges from root galls.
of species are parasitoids of the larval stages of insects, some spiders and
HYMENOPTERA 265
mites. The details of the life history of some of these species are truly remark-
able. The females of many species in the family Chalcidae have very enlarged,
strong hind legs, which they may use for fighting, but in a few they are used
to hold the jaws of antlion larvae apart while the female stabs her ovipositor
through the intersegmental membrane behind the head. The following is a
brief description of the activities of some of the major families.
More than 850 species of tiny wasps belong to the Trichogrammatidae.
These wasps parasitize the eggs of butterflies, beetles, bugs, thrips, flies, and
other wasps. Some even cling to the body of host females to ensure that they
Chrysidid wasps often roll are on hand when fresh eggs appear. The females of some species swim un-
into a ball as protection
derwater to parasitize the eggs of water beetles and dragonflies. The entire
against predators.
larval development may be as short as three days. Trichogrammatids are so
small (only 170 µm, or 0.17 mm—that is, smaller than many amoebae and
protozoans) that, to save space in their miniature bodies, most of their brain
neurons lose their nuclei. Even smaller than some trichogrammatids are the
so-called fairyflies, the Mymaridae, among whose member species can be
found the tiniest flying insects on Earth with wingspans of less than 0.25 mm.
These tiny wasps are also egg parasitoids of other insect orders. Nearly half of
all species use planthopper eggs and the eggs of related bug families, and the
amazing females of Caraphractus cinctus swim underwater to locate the eggs
of giant water beetles (Dytiscidae).
More than 650 species of fig wasp, Agaonidae, occur in subtropical and
tropical regions where fig trees grow, and as there is probably at least one
species of fig wasp specific to each of 900 or so fig species (and several species
might inhabit a single fig species), there might be as many as 1,000 agaonids
in total. Males and females are very different. The females are tiny, flattened,
winged wasps whereas the males hardly resemble wasps at all, being wing-
less with odd-shaped heads, very weak middle legs, and a curled abdomen.
Fig wasps and fig trees are totally dependent on each other for survival. The
trees rely on the wasps for pollination and the wasps, which have complex
lifecycles, can only reproduce inside figs. Both fig trees and Agaonidae play a
vital role in tropical ecosystems worldwide and provide invaluable ecosystem
services to traditional economies.
Nearly 3,500 species belong to the Pteromalidae. Most of these wasps
are parasitoids or hyperparasitoids inside or outside the larvae or pupae of
flies, beetles, wasps, butterflies, moths, and fleas. A few are gall-formers or
herbivores. Females may lay one egg, while others may lay hundreds of eggs
in a large host, which is paralysed and does not develop any further.
The Eulophidae, a family of more than 4,300 small species, are mainly par-
asitoids of leaf miner and gall-former larvae. The Torymidae comprise about
1,000 species that may be predacious or herbivorous in seeds as larvae. A large
number of species are parasitoids of gall-forming Diptera and Hymenoptera,
the females using their long ovipositors to drill through the tissue of the gall
to reach the host larvae inside. Other torymid species attack caterpillars, the
eggs of mantids, and the larvae of solitary bees and wasps that nest inside
plant stems.
The Encyrtidae, comprising more than 3,700 species of small wasp, are
mainly parasitoids of the immature stages and adults of scale insects, mealy
bugs, aphids, and whiteflies, although they also have hosts in many other in-
sect orders. Some species are hyperparasitoids of the larvae of pteromalids,
braconids, and even other encyrtids. Many encyrtids are polyembryonic. The
eggs divide repeatedly at an early stage of their development to produce any-
thing from ten larvae in small hosts, to a couple of thousand larvae in large
hosts.
Members of the seven families that make up the Chrysidoidea are para-
sitoids, with more than 6,000 species. Bethylids (Bethylidae) are a large group
of about 2,000 species that attack caterpillars and beetle larvae. The female
wasp locates a suitable host and stings it, causing paralysis or death. She then
lays a number of eggs on it, and, in some species, she may give a certain
amount of maternal care while her ectoparasitic larvae consume the host. An
even larger family, with more than 3,000 species, is that of the jewel wasp
(Chrysididae). The name of jewel or ruby-tailed wasp refers to the beautiful
and strongly metallic coloured body of these wasps. The body surface is ex-
tremely hard to protect them from bee and wasp stings and the underside of
the abdomen is flattened or concave, enabling the wasps to curl up into a ball.
Female chrysidids lay their eggs in the cells of nests of solitary bee or wasp
species. The chrysidid larva eats the host larva or nest provisions left by the
host bees or wasps. Some chrysidid species are parasitic on sawfly larvae or
pupae, some on lepidopteran pupae, and a few eat the eggs of mantids. Some-
times the life histories can be complex, involving an intermediate host. When
this host, carrying the jewel wasp’s egg, is paralysed by a digger wasp and
stored as food for the larvae, the egg hatches and the chrysidid larva feeds on
the digger wasp egg and food supply.
There is only one other large chrysidoid family: the Dryinidae. The larvae
of all 1,900 species are parasitoids of auchenorrhynchan nymphs or adults,
particularly those belonging to the families Cicadellidae, Delphacidae, Cixi-
idae, Membracidae, Cercopidae, and Fulgoridae. Female dryinids locate their
prey on vegetation and, when within close range, snatch their victim using
their legs and, sometimes, jaws. Many female dryinids have a strange grasp-
ing or pincer-like front tarsi that are used to hold the bug pressed against the
surface of the foliage or up in the air. The female then stings it and inserts an
egg. In most species, the egg is placed between abdominal segments, in others
the egg is inserted in the thorax or neck of the host bug. The effect of the sting
lasts a short time, and the host revives and continues its life cycle normally.
The hatched dryinid feeds, at first, on the host’s internal fluids. The first instar
is spent within the body of the host, but later instars develop inside a larval
sac, which bulges through the intersegmental membranes. The mature larva
will also eat the host’s body tissues and the sac may become as big as the bug’s
abdomen. When mature, the larva leaves the host, almost always killing it,
and moves towards the ground where it spins a cocoon.
The remaining superfamilies of the Aculeata contain a suite of fascinating
life histories. Tiphiid wasps (Tiphiidae, superfamily Tiphioidea) are slender,
shiny, yellowish-brown and black, or black wasps. Males of all species are fully
HYMENOPTERA 267
winged, but females can be fully winged or wingless. The underneath of the
middle thoracic segment has two lobes, which may cover the coxae of the
middle legs. Tiphiids are parasitic on the larvae of beetles, bees, and wasps. Fe-
males of the genus Tiphia dig through soil to find the larvae of scarab beetles,
and, when located, the female wasp will break open the cell, sting the beetle
grub, and may chew it with her mandibles before laying her egg. The wasp
then leaves and reseals the cell behind her. Females of the genus Methocha
are very ant-like and specialize in parasitizing tiger beetle larvae within their
burrows.
Velvet ants (Mutillidae, superfamily Pompiloidea), although velvety and
often ant-like, are not ants. Nearly 7,000 species are found, mainly in tropi-
cal regions. These wasps are black or reddish-brown with spots or bands of
red, yellow, or silver. They have short hairs and a head and thorax that has
a sculpturing of coarse dimples. Males are fully winged, but the females are
wingless and are most commonly seen running over the ground. Mutillid lar-
vae are ectoparasitic on the larvae and pupae of bees and wasps, but also some
dipterans and coleopterans. For example, species of the genus Mutilla special-
ize on bumble bees of the genus Bombus, while others attack a wider range of
hosts, including halictid bees, and pompilid and sphecid wasps. When female
mutillids find and bite open a host cell they check whether the larva inside is
suitable. If too young, the cell is resealed, but if the larva is fully developed,
an egg will be laid on it and the cell resealed. The mutillid larva hatches, eats
the contents, and pupates within a tough cocoon inside the host’s cell.
More than 5,000 species of spider-hunting wasps of the family Pompilidae
(Pompiloidea) are known. Many pompilids are moderately sized, but the fam-
ily includes some giants of the genus Pepsis measuring > 65 mm from head to
tail. Spider-hunting wasps are generally dark-coloured, blue or black, usually
with dark, yellowish, bluish, or black wings. Females are active hunters that fly
and run rapidly over the ground in search of spiders. Females may make mud
nests in crevices, dig branched burrows underground, or simply use the host
spider’s own burrow. After the spider is paralysed, the female makes a burrow
or drags it to a prepared burrow, lays an egg on it, and seals the burrow. Some
species are kleptoparasites, that is, they cheat by quickly laying their own eggs
on a spider paralysed by another wasp before it is buried. Some even locate
and break into an already-sealed nest.
More than 5,100 species in two families comprise the Vespoidea. Com-
mon or paper wasps belong the large family Vespidae. About 5,000 species are
known and most live in tropical or subtropical regions. The family is divided
into distinct subfamilies such as the Vespinae (yellow jackets and hornets),
Eumeninae (potter wasps), and Polistinae (paper wasps). Vespine and polis-
tine wasps are highly social insects living in colonies with a queen, males, and
sterile worker females. Eumenine wasps and species belonging to the other
subfamilies are solitary or very weakly social.
Yellow jackets and hornets make carton nests of paper made from chewed
wood fibres. The nest, which may be underground, in natural cavities, or
in bushes, may contain many thousands of workers. Developing larvae are
fed on a nutritious paste of butchered or chewed-up insect prey. This sort of
feeding system is known as progressive provisioning, as opposed to the mass

provisioning that occurs in spider-hunting wasps and others. Foraging may
be opportunistic but there is evidence that vespids react to aggregations of
suitable prey, such as mating swarms or leks, by picking up cues such as sex-
ual pheromones. They are able to change their foraging strategy to capitalize
on other prey items at different times of the day. The adults feed on nectar and
other sugar rich materials. The nests of Polistes may be covered with a paper
or mud carton, but many are uncovered in sheltered locations. Potter wasps
Species of Eumenes make
make small mud-lined nests in the ground, in stems, and a variety of natural
elegant vase-shaped nests
and stock them with a cavities. Species of the widespread genus Eumenes make elegant vase-shaped
number of paralysed nests of mud or clay attached to twigs or rocks. The nests are stocked with a
caterpillars or beetle grubs,
and an egg is suspended by number of paralysed caterpillars or beetle grubs, and an egg is suspended by
a slender thread from the a slender thread from the roof or walls of the nest before it is sealed.
roof or walls of the nest Ants all belong to one family, the Formicidae (superfamily Formicoidea),
before it is sealed.
which are the sister group to Apoidea (bees, sand wasps, cockroach wasps,
and aphid wasps). Just under 13,800 species are known and, although they are
morphologically quite similar, their biology is very diverse. Ants appeared in
the Early Cretaceous (145–65 mya) and radiated in the Late Cretaceous and
Early Tertiary. With the exception of Antarctica and a few oceanic islands,
ants are found everywhere on Earth, and have an immense impact on terres-
trial ecosystems. In most habitats, ants are the major predators, and in parts of
Central and South America, leaf-cutter ants (Atta spp.) are the major herbi-
vores and the most serious insect pests. In many terrestrial ecosystems, ants
move more earth than earthworms and are vital in nutrient recycling and
plant dispersal. Like the termites and some of the bees and wasps, ants are
eusocial and live in colonies ranging from a handful of individuals sharing
a tiny shelter, to tens or hundreds of millions inhabiting underground struc-
tures reaching 6 m below ground level. Being eusocial means that two or more
generations overlap at any one time, adults take care of the young, and there is
a division of labour between castes (reproductive kings and queens and non-
reproductive workers). Worker ants are wingless, sterile females, while the
queens and males, who form mating swarms at certain times of the year, are
winged. Mating may take place on the wing or on the ground; afterwards, the
males die, and the females lose their wings. Chemical messages in the form of
pheromones are important in colony regulation, trail making, raising alarms,
and defence.
Caste is determined by a complex and poorly understood interplay of
several factors such as diet, genetics, and epigenetics. The head is modified
according to caste and species and may be very large with massive or other-
wise specialized jaws. Some castes exhibit remarkable adaptations to the jaws
and head for crushing seeds, blocking nest entrances, dismembering enemies,
Section through a nest of a
mining bee (Halictus sp.) and catching fast-moving prey.
showing numerous cells Ants can be herbivorous seed gathers or fungus growers, carnivorous or
arranged around a central
omnivorous, and some rely exclusively on the honeydew produced by sap-
burrow.
sucking bugs. Many species of insect and plant have evolved symbiotic and
often highly complex relationships with ants. Many ants are associated with
species of plants, which may provide them with homes in the form of galls or
HYMENOPTERA 269
larger domatia (ant homes), often in return for protection of the plants from
herbivores, encroaching vines, or fire.
The family is divided in about 16 distinct subfamilies, of which the biggest
by far are the Myrmicinae and the Formicinae. Some myrmicine ants have
stings, while formicines defend themselves by spraying formic acid. It is esti-
mated that every year, the world’s formicine ants release one million tonnes
of formic acid into the atmosphere. The infamous driver and army ants of
tropical regions belong to the subfamily Dorylinae. Colonies move in mas-
sive columns, several million strong, raiding termite or ant colonies, and kill
and butcher anything that is unfortunate enough to get in the way (includ-
ing tethered vertebrates). Some species can be pests in buildings, and the Red
Imported Fire Ant (Solenopsis invicta) is an aggressive and abundant pest in
the southern states of USA that causes great damage to soya bean crops.
The Apoidea contains 17 families, with about 26,700 species. The common
Section through a nest of a
solitary bee (Andrena sp.). names of these insects include solitary hunting wasps, digger wasps, and sand
wasps, while the clade known as the Anthophila includes the honey bees,
carpenter bees, sweat bees, bumble bees, etc. Most of the non-anthophilan
Apoidea are solitary wasps that nest in soil, decaying wood, plant stems, in the
burrows of insects, or in self-made mud nests. The females catch prey, paral-
yse it with their sting, and transport it back to the nest, where they will seal
it in with an egg. Some species may use one single large prey item while oth-
ers may use several small ones. Prey varies widely across many insect orders,
as well as spiders, and many non-anthophilan Apoidea keep to a particular
sort of prey item or a particular size of prey item. In most cases, the nest cells
are mass-provisioned with enough food for the larva to complete its develop-
ment, but the females of some species practice progressive provisioning and
continue to supply their larvae with more prey as they grow. A few species
are kleptoparasite and lay their own eggs in already provisioned nests of other
sphecids. The European Beewolf, Philanthus triangulum (Philanthidae), pro-
visions its brood cells with paralysed honey bees. When there are few honey
bees around, the female chooses to lay more male eggs as sons are cheaper
to produce. The female will also provide each offspring with less food than
normal.
The Anthophila contains more than 16,000 species of bee, most of which
are solitary. There are seven distinct families of which the larger ones deserve
special mention. There are about 2,000 species of plasterer and yellow-faced
bees (Colletidae). The species in this family are solitary and make very sim-
ple nests. Plasterer bees excavate nest burrows in the ground and use a special
secretion from an abdominal gland to line the interior of their cells. This se-
cretion dries to form a waterproof, cellophane-like film, which protects the
interior. Yellow-faced bees nest in the pith of plant stems, the empty burrows
of wood-boring insects, or even empty plant galls. Colletids transport pollen
on their hind legs or carry it in their crops. The larval cells are provisioned
with a sloppy mixture of regurgitated pollen and nectar.
The Halictidae, a family of more than 4,400 species, are called sweat bees
because of the habit seen in some species of drinking perspiration. Many are
solitary, while others are more social. Sweat bees mostly nest on the ground,
and the nests usually take the form of a single vertical burrow with or without
lateral tunnels. Brood cells are arranged in dense clusters on the tunnels and
are lined with a waterproof secretion. Some species are parasitic in the nests
of other halictids. Many species are important crop plant pollinators.
More than 4,000 species of mining bees comprise the Andrenidae. Females
of most species are solitary and make nests in soil burrows, although these
often occur in large aggregations. Pollen from a few related plants is collected
and transported in a brush of hair on the hind legs called a scopa. The larval
cells are lined with a waxy substance secreted by an abdominal gland and are
provisioned with a mixture of pollen and honey moulded into a small pellet.
Leaf-cutter and mason bees (Megachilidae) are solitary, nesting in natural
cavities in the ground, dead wood, hollow stems, and snail shells. Megachilids
do not line their brood cells with glandular secretions, but instead collect ma-
terials such as mud, resin, and leaf material, or animal or plant hairs to do the
same job. Pollen, when collected, is carried in a brush of stiff hairs on the
underside of the abdomen. Leaf-cutter bees are named for their habit of cut-
ting neat circular pieces of leaves or petals to line their brood cells, which are
arranged in rows inside the nest cavity. Other species are called carder bees
because they use their jaws to strip the hairs from woolly leafed plants, which
they tease out to line brood cells. Some species are parasitic in the nests of
Corbiculum other bees, including other megachilids.
Just over 4,000 species of digger (Anthophorinae), cuckoo (Nomadinae),
and carpenter bees (Xylocopinae) belong to the Apidae, the family that in-
cludes honey bees and bumble bees. Digger bees burrow in the ground and
provision their brood cells with a mixture of pollen and honey. Cuckoo bees,
which can be very wasp-like, are all parasitic in the nests of bees of all other
families except the Megachilidae and Apidae. Carpenter bees make nests by
excavating tunnels parallel to the grain of solid wood, in which a series of
brood cells is made. Each cell is provisioned with a large mass of sticky pollen
on which the female lays a single large egg before sealing the cell with chewed
wood fibres. Female xylocopines will guard their nests against predators, par-
Location of the corbiculum
(pollen basket) on the hind asites, and even female conspecifics that might attempt to take over the nest
leg of a honey bee. and lay their own eggs.
There are about 250 Bombini (bumble bees) species and 8 Apini (honey
bees) species. Bumble bees are very hairy and stout-bodied, whereas honey
bees are smaller and slenderer. The females of most species have a special-
ized pollen-carrying apparatus, the corbiculum or pollen basket, on the outer
surface of each of the hind tibiae.
Members of these tribes are highly social and live in colonies with a queen,
males, and sterile worker females. Bumble bee nests may be under or on the
ground, and are made of grass with internal, wax-constructed brood cells.
Many species make their nests in disused burrows and nests of small mam-
mals. The larvae are fed on pollen and honey, which is stored in other cells.
The queen rears workers first to build up the colony, with males appearing
later in the summer. Some bumble bees behave like cuckoos, laying their eggs
in nests of other bumble bees.
HYMENOPTERA 271
Early humans collected honey from wild bees and bee-keeping is docu-
mented from at least 9,000 years ago. Honey bees belong to the genus Apis,
and the best known of the eight species is the Western Honey bee (Apis mel-
lifera), which has been spread worldwide. Honey bees may possess the most
complex social life of all eusocial insects. Each colony comprises a single egg-
laying queen, many tens of thousands of workers, and up to 2,000 males or
drones. The nest consists of a vertical array of double-sided wax combs di-
vided into thousands of hexagonal cells. The hexagonal shape of the cells is the
most economical shape and configuration, which will hold the most honey
for the least amount of wax. Despite the fact that the cell walls are less than a
tenth of a millimetre thick, a hexagonal array gives the comb great mechanical
strength. Individual cells are built at a slope of 13 degrees upwards to keep the
honey from running out. When honey bees start building a nest they gather
together in a ball, the inside of which soon heats up to 35◦ C, the temperature
needed for wax production. Wax is produced by glands on the underside of
the worker bees’ abdomens, and it is gathered and worked into shape with
the front legs and mandibles. Wild honey bees need to start building their
nest from scratch, but beekeepers provide their bees with a foundation sheet
pressed out from pure beeswax, reinforced with wire and mounted in a frame.
The modern type of hive known as the Langstroth hive was invented in 1851
in the USA and was a great advance over early bee skeps made out of straw.
The queen is kept in the lower brood chamber by means of a queen excluder, a
metal grille prevents her from laying eggs in the upper chambers. This means
that the upper chambers are only used for storage, which makes the collection
of honey and pollen very easy.
A colony has one queen, forty- to eighty-thousand workers, and a few hun-
dred male or drone bees, whose only job is to fertilize new queens. Worker
bees are sterile females, which are kept sterile by the queen who produces a
pheromone, called queen substance, from small glands in her head. This sub-
stance is transferred from bee to bee within the hive and stops the workers’
ovaries from developing. A queen can lay up to 1,500 eggs in a day, and in her
four-year life she may lay one million eggs.
During the summer, an adult worker honey bee only lives for six weeks
or so. When newly emerged, a young bee spends her first two days eating
pollen and honey. After this she spends three weeks working inside the hive
as a nurse feeding larvae with royal jelly, which she secretes from glands in
her head. Bee larvae, which are going to become workers or drones (males),
only get royal jelly for the first few days and then are fed on pollen and honey.
Larvae that are going to become queens are fed on royal jelly throughout their
development. When young, nurse bees produce wax, which is used to build
new cells and repair the comb. For the last weeks of her life the worker is a
forager. The exact age at which a worker becomes a forager varies according
to colony condition. It seems that a high number of active foragers will de-
lay the transition from nest bee to forager. Foragers gather nectar and pollen
from flowers. The nectar is sucked up and stored in an anterior region of the
gut called the honey crop. Enzymes in the gut act on the nectar and, when
back in the nest, the bees ‘pant’ to evaporate water from the nectar. They then
regurgitate all of the nectar into storage cells. Here the nectar is further thick-
ened by evaporation. Workers collect pollen and carry it in special baskets
on their hind legs. Pollen, rich in protein, is used to feed worker and drone
larvae. Foraging workers returning to the nest perform dances on the comb
to tell other workers the direction and distance of the food source she has
visited. A figure of eight waggle dance of varying speed is performed if the
food source is more than 100 yards away. The angle of the waggle run in the
middle of the dance relative to the vertical allows the bees to follow a course
The Round Dance means to the food source, relative to the position of the sun. If the food is less than
the food supply is nearby
(up to 100 yards).
100 yards away, a round dance is performed.
Honey bee colonies attract a number of vertebrate and invertebrate in-
truders whose aim is to steal honey. Workers attack en masse, and many will
die in the process. The sting of a honey bee is barbed, unlike that of a social
a wasp, which can sting repeatedly. An amazing thermal execution technique
is used by honey bees in response to enemies such as hornets. The workers
w crowd tightly around the hornet and shiver their thoracic muscles. This gen-
erates a large amount of heat inside the ball of bees and the hornet is literally
fried alive. Dying in defence of the colony has been pivotal for understanding
the biological basis of altruism in animals. Due to the hymenopteran method
The Waggle Dance shows in of sex determination, haplodiploidy, females that are diploid share 75% of
symbolic form the direction
their genes with their sisters (50% comes from their haploid father and 25%
and distance of the food
source. Direction in relation from their diploid mother). Worker bees are thus more closely related to each
to the sun’s position is other, than to their offspring, if they were able to produce any: if they had
indicated by the angle
(a) between the waggle run daughters themselves, they would only share 50% of their genes with their
and the vertical. Distance is offspring, which is less than the 75% that they share with their sisters. There-
indicated by the duration of fore, it makes evolutionary sense for them to help rear and defend their sisters
the waggle run (w).
and the queen, as the latter is essential in producing more worker bees.
When a queen ages, when the colony gets too crowded, or when there is
not enough food, the workers construct large queen cells. Just before the first
queen emerges, the old queen and about half the workers leave the colony as
a swarm. The first queen to emerge stings any other queens to death and then
leaves the hive for a mating flight. The drones compete with each other, and
the queen may mate with several males. After mating the males die, but their
sperm will be used by the queen to fertilize all the eggs she lays during her
life.
Apids are major crop pollinators and the value of crops pollinated by
honey bees alone greatly exceeds the value of the honey, wax, and other use-
ful products they provide. Leaving aside the incalculable benefits that aculeate
Hymenoptera bring, a significant number of people die every year from be-
ing stung. The Africanized ‘killer bees’ found in South America and in some
southern United States of America, resulted from accidental mating between
drones of Apis mellifera mellifera and queens of a very closely related African
subspecies Apis mellifera scutellata. This hybrid is very industrious, but, un-
fortunately, very aggressive, and will readily attack anything that comes too
near to the colony. The sting volume and the venom toxicity is the same as
that of a normal honey bee, but, as victims may receive many hundreds of
stings, the effect, particularly on the young and the elderly, can be fatal. Their
HYMENOPTERA 273
behaviour is sufficient to identify them, but if you want to be really sure, you
can measure the ratio of two wing veins, but it is best do it with a dead one.
The oldest hymenopteran fossils are known from the Triassic (around
224 mya). Phylogenomic work suggests that Hymenoptera arose considerably
earlier, sometime between the Carboniferous–Triassic boundary (280 mya),
rendering this order a very ancient lineage indeed. Hymenopterans are the
sister group to all other Endopterygota.
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SECTION 3
Fieldwork
279
You can learn a certain amount from books, but in order to really learn about
the secret lives of insects, you must study them at first hand. With organisms
as small, numerous, and diverse as insects, you will not be able to get very far
without collecting and killing them for identification. A few people profess to
be able to recognize many species on sight, but for small and cryptic species
their identifications will be questionable. In any case, an identification with-
out voucher material carries little weight. Many species are distinguishable
only by looking at their dissected genitalia or other minute details. Even with
perfect eyesight and a hand lens it is impossible to count the number of bris-
tles on the head or body of live flies, and most small moths rest with all their
useful identification features obscured. To quantify biodiversity in any habi-
tat, whether for a basic site description, environmental impact assessment, or
pest management survey, it is necessary to find out what species are present,
and in what numbers.
However, it is important to consider conservation issues. A growing ap-
preciation of the importance of insects resulted in the protection of many
species around the world, especially in the light of the growing destruction
of natural habitats and other environmental changes. Among the influen-
tial international organizations that exist, the IUCN (the World Conservation
Union, formerly the International Union of Conservation of Nature and Nat-
ural Resources) based in Gland (Switzerland) and the World Conservation
Monitoring Centre based in Cambridge (UK) are particularly relevant for
those interested in insects. Legal protection has been extended to particular
species (Collins, 1987 & 1988; Heywood, 1995), and a list of threatened inver-
tebrates is given by Wells et al. (1983), and of swallowtail butterflies by Collins
and Morris (1985). An IUCN Red List of Threatened Animals (1996) lists 73
insect species as extinct or extinct in the wild, and a further 537 species are
recorded as critically endangered, endangered, or vulnerable. In reality, this
is the tip of an iceberg and little data are available. Whenever you collect, re-
member that there is a code of practice for collectors of biological material
(Jermy et al.,1995, p. 57), and many countries have legislation that will affect
your activities (e.g. Wildlife and Countryside Act, 1981).
Collecting
The indispensable reference work is Ecological Methods (Southwood and
Henderson, 2000). Methods for collecting insects can be divided into two
sorts—relative and absolute. Relative sampling methods provide only pres-
ence/absence data and indicate how abundant one species is relative to
another, but do not allow you to measure the population sizes. Examples
of relative sampling methods are pitfall trapping, flight intercept traps, and
light trapping. In all these cases, it is unknown what area or unit of the habi-
tat has been sampled. Using absolute sampling methods, the density of the
species collected can be readily calculated (number per m2 is a standard mea-
sure). Insecticidal knockdown techniques, suction sampling, hand searching
(over a known area), and extraction from a known quantity of leaf litter are
280 SECTION 3 FIELDWORK
all examples of absolute sampling methods. No single method will collect all
the species present in a sample area, but by using absolute techniques you will
be able to make meaningful comparisons between different sites or studies.
Relative collecting techniques

There are many methods for collecting insects. The key to successful collect-
ing is using the right equipment for the job, as well as the right timing. This
section discusses relative collecting techniques.
Butterfly and aerial nets Aerial or hand nets are lightweight nets designed
for catching flying insects. They should be lightweight and strong with as large
a mouth as is practicable. Quick reactions and balletic movements are needed
to catch very fast-flying or highly manoeuvrable insects, such as dragonflies
and skippers, but gentle passes over the top of vegetation will collect large
amounts of small flies and wasps. It hardly needs saying that fine aerial nets
should never be used for sampling insects from dense or thorny vegetation.
Sweep nets Sweep nets are made from strong, close-woven material and
are designed for thrashing through vegetation. The bag should be deep
enough to allow you to trap the catch by gathering the material together in
one hand below the frame. Apart from suction sampling, it is the only way
to sample insects quickly from low-growing vegetation and grasses. A sweep
net is considerably lighter and easier to use than any motorized suction sam-
pler but there are, of course, drawbacks. Sweep sampling is a relative sampling
method and is (at best) only semi-quantifiable. Sweeping will not catch leaf-
mining and stem-boring species and insects that live close to the soil surface.
You may also miss the larger, faster species, which, alerted by your vigor-
ous progress through their habitat, will escape before you get to them. The
sweep net can also be used to collect insects in a rough manner from trees
and shrubs. It is impossible to use this technique if the foliage is wet as this
will result in much of your catch being reduced to a crumpled mush.
To make your results comparable between different sites you will need to
standardize your sampling effort in some way. You should use the same net
(the mouth diameter of the net is an important factor) and do a fixed number
of sweeps per sample (25, 50, or 100 sweeps are typical). How you sweep is
important and people vary a great deal in their technique, so it is best to get the
same people to do all the sampling. Another point worth remembering is that
you should always sweep into the wind. Describing the proper technique is
difficult, but bear in mind that the object is to sweep as evenly as possible from
a particular area or volume of vegetation. Stoop down and hold the handle of
the net firmly in both hands, and then sweep the mouth of the net to one side
as far as you can comfortably reach. The essential point is that the net should
be kept as low to the ground as possible at all times, and at the end of each
pass the net is turned swiftly to begin the next pass or sweep to the other side.
The movement of the net must be nearly horizontal and not parabolic. Repeat
the sweeping action the required number of times as you walk through the
vegetation, and at the end of the last sweep, turn the net mouth or grab the net
above the catch so that you do not lose anything. The bigger the number of
COLLECTING 281
sweeps the more difficult it will be to sort out the catch from the plant debris
that you will unavoidably collect as well. If you are sampling for a specific
insect species or group, you might simply search through the catch in situ,
collecting what you need with a pooter (see figure below), and returning the
rest to the wild. Otherwise you need to bag up the material and sort it out
later.
Transferring a sweep net catch to a strong polythene bag is easiest with
two people. One person holds the plastic bag so that the opening assumes the
shape of a slit. The other person grasps the net together above the catch. The
contents of the sweep net are everted into the plastic bag by gradually draw-
ing the material of the net gradually through and over the retaining hand.
Expel most of the air from the plastic bag and tie the top (after putting a label
inside).
There are two schools of thought as regards sorting sweep samples (and
suction samples): the live sorters and the dead sorters. Killing the catch (for
instance, with fumes of ethyl acetate) in the field is useful, as some of the
catch might start to eat the others, especially if there is going to be a long
delay between sampling and sorting. Also, heavier and stockier insects might
trample over and damage more delicate species. Live sorting is also preferable,
as dead insects, especially small ones, are ten times more difficult to sort out
from debris than live ones and anything not required can be returned to the
field. If you opt for live sorting, try to keep bagged catches out of the sun. Live
sorting of sweep samples is greatly facilitated by using a sorting hood. This
simple device, which can readily be made up in the field, employs the principle
that most insects will move towards a light source. The sides, top, and bottom
of the hood should be opaque with only the back or part of the back panel
transparent. With a light placed behind or the hood simply angled towards
the sun, your catch can be carefully tipped out and the live insects pootered
or brushed into alcohol as they move towards the light. A simple hood can be
made from a cardboard box lined with white paper with a cellophane sheet
or fine netting taped to a hole in the back.
Beating trays Victorian collectors were clever. You cannot easily collect
insects in the rain so why carry a beating tray as well as an umbrella? Al-
though better if made of a white material against which to see your catch, any
umbrella held upside-down could serve as a reasonable beating tray.
Various designs of beating tray are available or you can make you own.
The tray is held under the foliage to be sampled and the branch shaken or
given sharp, jarring blows with a stick. Do not thrash the vegetation as you
will damage the plants and end up with a large pile of leaves and debris on
your tray, which will take longer to search through. Some insects are very
good at holding on and you will not, of course, catch internal feeders, such as
leaf miners and stem borers. If you are specifically trying to catch very active
species, a modified beating tray in the form of a funnel-shaped net with a
collecting bottle might be preferable. It is best to keep spiders and insects
separate, as a spider silk quickly renders a pooter tube full of insects a tangled
mess.
A beating tray and butterfly

net.
Like sweep netting, beating is a very simple and quick technique, but
also suffers from being qualitative. It is also restricted to how high you can
comfortably reach. You can, however, try to compare the beaten insect com-
munities from different species of tree or shrub by replicating samples and
standardizing the effort (the number and vigour of blows, perhaps) and area
sampled as best you can. The beating of wet foliage is not recommended for
the same reasons as sweeping wet vegetation.
Light traps That some insects are attracted to light must have been noted
shortly after humankind discovered fire. Eighteenth- and nineteenth-century
entomologists used various forms of hurricane lamps and Tilley storm
lanterns to go ‘mothing’, but, in recent decades, the use of light sources that
produce large amounts of ultraviolet radiation has revolutionized light trap-
ping. There have been many designs of light trap produced in the last 50 years,
and all exploit the phenomenon that many insects (but by no means all) will
fly towards bright lights at night. Moths, notably, are attracted in this way,
but it should be remembered that it is mainly the males who respond, females
preferring to sit in concealment among vegetation. The reason why moths are
attracted to light is that in nature, they use the moon to navigate (transverse
orientation). If they keep the angle that the light rays subtend to the omma-
tidia constant, they should fly in a more or less straight line. When confronted
with a brighter light that is much nearer, the same behaviour will cause them
to take a decreasing spiral path into the light source.
In tropical areas, lights of almost any kind can pull in fantastic numbers
of beetles, bugs, and other insects, as well as moths. Around the world the
installation of vapour discharge lamps for street lighting has reduced popula-
tions of some moths. Many predators have learned to cash in on this bonanza.
COLLECTING 283
Bats will forage in the vicinity of bright lights, and birds will clean up the re-
mainder in the morning. In Papua New Guinea, one of us (George McGavin)
has often observed foraging columns of weaver ants (Oecophylla smaragdina)
queuing up along lit fluorescent tubes to snatch insects as they fly towards the
light.
The trouble with light traps is that you cannot be sure what will be at-
tracted to them, and what will not: indeed, some insects are repelled by light.
Many physiological, behavioural, and environmental factors are involved.
The biggest problem with light trap data is that it is impossible to know over
what area the trap is effective. In wooded habitats the range of the light may
be quite small due to obstructions, but even in open habitats you cannot be
sure. Experiments using a 125-watt mercury-vapour lamp have shown that
the effective trap radius, far from being 90 m as originally thought, is rather
less than 5 m.
Light traps are most useful to show the presence and perhaps relative
abundance of particular species. Quantitative analysis is tricky, but with
replications, using the same trap design, you might be able to get a rough com-
parison of different areas or an idea of how the populations of some species
fluctuate over time. They are thus useful for general survey work but be very
careful not to make more of the data collected than it warrants.
Large traps, such as the Robinson MV moth trap, which use mercury-
vapour discharge bulbs, require connection to a mains supply or a generator.
This type is not generally useful far from a field station or where you can-
not drive a vehicle. Smaller variations of the Robinson trap, such as the
Heath Trap that can be run from a 12-V car or motorcycle battery, can be
used in more remote locations. There are a number of portable designs that
use smaller batteries permitting their use in very remote or difficult terrain.
Various sorts of light trap, their operation, and limitations can be found in
Muirhead–Thomson (1991), Davies and Stork (1996), and Fry and Waring
(1996). A very convenient light trap for use in the tropics is described by
Robinson and Jones (1996). The heart of this cheap and simple trap is a 4-
watt actinic tube powered by four standard D-size batteries. Larger-wattage
tubes are available but need bulkier batteries.
Material that comes towards light can be collected by hand or by pooter
and should be killed and preserved in a manner appropriate to the taxon
in question. Most non-lepidopterans can be put into an ethyl acetate (avoid
inhalation and contact with plastics, especially styrenes) or cyanide (avoid
contact and inhalation) killing jar before being decanted into paper envelopes
and stored dried or fresh in a sealed plastic tub with chlorocresol. Small or
micro-moths should be taken from the killing jar and pinned using a minute
or micro-pin vertically through the thorax. They are then pinned into small
plastic boxes lined with Plastazote™ (or similar expanded polyethylene foam),
and the wings should be set forward and held in place with additional micro-
pins. In this manner a large number of fragile specimens can be kept and
transported safely. Butterflies and robust moths, such as hawk moths, can be
put in small envelopes or folded paper triangles inside a sealed plastic tub
with chlorocresol as an antifungal agent.
Like most skills, the art of setting butterflies and moths is best learned by
watching a professional, but instructions are given in a number of publica-
tions (Arnett, 1985; Dickson, 1992; Walker and Crosby, 1988).
Flight intercept traps Any sort of static trap that catches insects in flight
could reasonably be called a flight intercept trap (FIT). The term is used more
specifically to describe a single, rather simple type of trap. A piece of material
(usually black terylene) is stretched tightly between poles or trees, and plastic
containers are arranged on the ground along its length. The containers are
part-filled with water, preservative, and a little detergent to break the surface
tension. Insects flying into a FIT will crawl up and escape from the top, drop
downwards, or just fly off again. Some specialists use a roof over the FIT to
prevent insects from flying away from the trap. If the traps are going to stay on
site for more than a couple of days, placing crystals of chloral hydrate or pour-
ing propylene glycol in the water containers will prevent decomposition of the
catch. Although a remarkably effective technique to catch flying insects, one
problem with FITs is that you cannot be sure whether they are acting purely
passively or if some insects are attracted to them. Also, if you require direc-
tional data, such as whether or not insects are flying in a particular direction,
for example, in or out of a woodland or other habitat, you will need to make
modifications to available designs or make you own. Insects collected from
the tubs can be strained out using a tea strainer or small sieve and transferred
to alcohol in labelled tubes.
© George McGavin
A flight intercept trap, with

collecting trays beneath.
Malaise traps, FITs of a particular design, are named after entomologist

René Malaise, who noticed that insects that entered his tent always seemed
to accumulate ‘at the ceiling-corners in vain efforts to escape at that place
without paying any attention to the open tent door’. His design, which was
published in 1937, has spawned many variants over the years. Malaise com-
mented on the main advantages of his trap; it is ‘better than a man with a net’
COLLECTING 285
and ‘it can catch all the time, by night as well as by day, and never be forced
to quit catching when it was best because dinner-time was at hand’.
© George McGavin
A Malaise trap.
Malaise traps are essentially open-sided tents made of material, which

have a high point in the roof where the insects tend to collect and are
funnelled into a container. To increase the efficiency of the Malaise trap,
water-filled containers can be arranged on the ground under the central par-
tition. In this way insects that do not crawl upwards in response to flight
interruption will also be trapped. There is no doubt that they are effective in
catching flying insects and, lashed to a light frame, they can even be pulled up
into forest canopies. It is often said that Malaise-type traps are non-attractant,
but anyone who has used them will know that this is not true. The colour of
the material used to make them has a significant effect on the composition
of the catch. In some habitats black malaise traps are very attractive to horse
flies (Diptera; Tabanidae). This phenomenon has been put to good use in the
design of traps for blood-feeding flies.
For the majority of survey-type work, the collecting bottle of the Malaise
trap is filled one-third to one-half full with 70% alcohol, which serves to kill
and preserve the catch. If, however, you are specifically collecting Diptera,
most species are best collected dry with a killing agent, such as a piece
of Vapona Fly Killer™, or a similar domestic, slow-release strip of insect
killer containing dichlorvos wrapped loosely in tissue at the bottom of the
collecting bottle.
Sticky traps and window traps are variants of FITs. Sticky traps are sim-
ply pieces of Perspex or glass coated with sticky material (e.g., Hyvis™) and
fixed to a post. Window traps should be as transparent as possible and usu-
ally have a water-filled tray underneath to catch insects that hit the surface
and drop down. The major problem with sticky traps is that the glue used to
trap the catch is often difficult to remove and will almost certainly require the
use of dangerous and/or inflammable solvents. Even if you get specimens off
the traps in one piece, it is often very difficult to identify them. Sticky traps
can be useful for the regular monitoring of populations of pest insects where
identification does not pose a problem.
1 Dig hole slightly deeper
than pitfall trap Pitfall traps Pitfall traps are a very easy and cheap way to catch active
ground-living arthropods of all kinds. A hole is dug with a trowel and a plas-
tic container of some kind is sunk into the ground so that its rim is level or
slightly below that of the surrounding ground. It is no good at all if there is
even the slightest lip showing above ground, as small animals will simply walk
around the circumference. Bits of debris and foliage that lie across the top of
2 Place two plastic cups
inside hole the trap will act like little walkways allowing insects to crawl across. Once
properly set, pitfalls are left and animals will fall in, and the steep, slippery
sides should ensure that most of them will not escape.
Pitfall traps are usually one-third filled with water or alcohol to trap and
Outer cup kill the catch. Very small insects can sometimes escape from the surface of
has hole in water, so a few drops of detergent are added to lower the surface tension.
bottom
When using water, the catch will start to decay in a day or two, faster in warm
3 Backfill cup and hole and
tamp down
weather, so it is vital that the traps are emptied daily. If you are not able to ser-
vice the traps regularly, or if you wish to leave them undisturbed for a week,
you should use a mixture of 40% ethylene glycol (antifreeze) in the water as a
preservative. Some older texts recommend a saturate solution of chloral hy-
drate, but it should be generally avoided as it is very toxic and needs to be
handled with great care. Whichever technique you employ, the contents of the
4
traps should be preserved as soon as possible in 70–80% alcohol. Of course,
Discard
alcohol can be used in the traps in place of water, but it is expensive, will
evaporate faster than water, and a few studies have suggested that the smell
of alcohol can act as a repellent or attractant for some species. Do not use
formaldehyde solutions—they are dangerous and a potential environmental
hazard.
Using finger nail to hold A well-tried and tested protocol for pitfall trapping any given area employs
outer cup in place, take cheap, plastic drinks cups with a rim diameter of 7 cm (internal volume 200–
out inner cup and discard 250 ml). The standard unit of trapping effort could be, say 180 trap days; this
soil
could be 60 traps operated over a period of three days, or 30 traps over six
5 Carefully replace inner days. Traps are placed in lines of ten with a spacing of 4–5 m between each
cup and check rim
trap and between adjacent rows. Each trap unit consists of two drinks cups,
an outer one and inner one.
To set each trap, dig a hole larger than the drinks cups with a trowel and
make sure it is just deeper than the rim of the inner cup. Put two cups inside
each other into the hole and back-fill the space around the outer cup. Gently
Fill cup to 13 with pitfall
fluid (see text) press down and smooth the soil around the rim of the inner cup. Do not worry
if the inner cup is now partly filled with soil at this stage. The whole point of
Pitfall trap construction.
having two cups is that you can now take the inner one out and empty excess
soil, preferably off-site. The inner cup is now replaced with a clean one. The
outer cup protects the inner one and allows subsequent rapid emptying of the
inner cup. The outer cup should have a small hole punched in the bottom to
allow rain to drain away. If this is not done, rain can seep between the two
cups and the inner one will float up, spilling its contents. If you are going to
leave the traps for more than a day or two, or if there is the likelihood of very
COLLECTING 287
heavy rain, it is a good idea to make a neat hole in the inner cup with a paper
punch about 2 cm from the rim and glue a small piece of terylene or similar
material over the hole. In this way, if the cup fills up, the excess water will
drain away, and the catch will be retained.
When all the traps are dug in, the inner cups are one-third filled with wa-
ter containing a few drops of detergent. To save time and depending on the
nature of the habitat, the position of the grid can be marked with a cane.
Catches from all the traps from one site and one day are collected by pour-
ing them through a sieve and bulked together in 70% alcohol. If you really
want to keep them, any vertebrates collected should be stored separately from
the invertebrate material. As with keeping any biological material in tubes of
alcohol, the total volume of the catch should never be more than half the
volume of the storage tube. If you completely fill the tube there will be lit-
tle room for the alcohol, which will become diluted anyway, resulting in the
decomposition of the catch.
If you want to trap specific groups of insects, such as those attracted to
carrion and dung, it is a simple matter to incorporate appropriate bait. You
will, of course, still catch things that just blunder in but, with experience, these
can be ignored.
The use of subterranean pitfall traps has produced some very interesting
data on the occurrence, abundance, and distribution of species that spend
most or part of their lives underground. Owen (1995, 1997) describes the
construction and use of a simple pitfall trap for the repetitive sampling of
underground faunas.
Pitfall traps can be used dry for live collection of specimens, in which
case they need to be serviced much more regularly than those where the
catch is killed in fluid. A large ground beetle in a dry trap will eat anything
within it.
There are many drawbacks to using pitfalls. Sometimes, small mammals
and amphibians get trapped and die unnecessarily. When pitfalling in dry
areas, birds and other animals soon learn where to get a drink. Pitfall traps
with a large surface area might act as a water or pan trap and attract flying
insects as well. It would be impossible without watching it all the time to be
sure whether an insect stumbled and fell into the trap as intended or flew
into it. With large or long-term traps, it is often a good idea to arrange some
sort of cover to keep rain out or mesh of various grades to exclude certain
sizes of animal. Another problem is that you can never be sure whether even
ground-living species might be attracted to them in some way, but as with
light trapping, the biggest problem associated with pitfalls is that you cannot
be sure over what area they are trapping, and they are greatly affected by veg-
etation structure. Despite these very serious disadvantages, pitfall trapping
remains a standard qualitative sampling technique— it is cheap and produces
large amounts of material with relatively little effort.
Water pan traps (or yellow pan traps) A very simple trap can be made
from almost any sort of shallow tray or container. The pan is partly filled
with water and preservative to which a few drops of detergent are being
added to reduce the surface tension. A bright yellow background colour has
been found to be the most effective in attracting small flies, wasps, and bugs,
such as aphids, although variants with blue colour may work better in certain
areas. An approach that one of us (Leonidas-Romanos Davranoglou) and our
colleagues frequently use is to create two-coloured traps, where one half of the
trap is painted yellow, and the other half is blue—in this way, you can attract
species that prefer either colour. To prevent overflow due to heavy rain, very
small holes can be made on the sides of the trap. Pan traps have many ad-
vantages: you can put out scores of them very cheaply and you can use them
virtually anywhere that you can reach, from ground level to the tops of trees.
The main disadvantage, of course, is that the method is relative. Furthermore,
they might dry up in very hot conditions, they need to be emptied frequently,
and the catch preserved in alcohol. Lepidoptera caught will need careful
drying.
A yellow pan trap. © George McGavin
Lures and baited traps Insects will respond to all manner of environmen-
tal cues, and this can be used to locate food, mates, or egg-laying sites. Rather
than employing general collecting techniques, specific insects can be attracted
by various baits depending on their functional ecology. Indeed, many insects
will only be caught using this sort of approach. For example, scarabs of the
genera Deltochilum (subgenus Aganhyboma) and Canthon are highly unusual
in that they frequently prey on millipedes. One of the main ways to collect
them is by using pitfall traps baited with crushed or injured millipedes—the
defensive chemicals they exude are irresistible to the scarabs and can attract
them from far away. Baits can be divided into major groups: dung, carrion,
rotting or fermenting plant material, sexual odours (pheromones), and live
animals. There is plenty of scope for originality and ingenuity in the design
and refinement of baited traps.
COLLECTING 289
© George McGavin
A home-made bottle trap
baited with a fish head.
Animal dung and carrion will attract specialist flies and beetles, and the
baits can be incorporated into pitfall or FITs. The attraction of butterflies to
putrid fluids is well known. A small dustbin or bucket partly filled with lique-
fying rotting fruit (add yeast to accelerate process) and/or carrion will attract
certain nymphalid butterflies, particularly those of the genus Charaxes. The
Purple Emperor butterfly, a woodland species found across Europe and Asia
to Japan, was often caught by using ‘the juices of a dead cat, stoat, or rabbit’
or ‘a seething mass of pig dung’. Males (mainly) of many species around the
world also indulge in puddling. Large numbers of several species may gather
to drink around the edge of puddles, urine patches, salt runs, and other damp
areas. The physiological need for salt and other minerals has been put for-
ward as an explanation for this phenomenon. Butterflies of the neotropical
genus Morpho can be attracted to lures of bright flashing colour. Materials
that reflect ultraviolet will be attractive to many sorts of insects, including
water beetles.
The technique of ‘sugaring’, now not as fashionable as it once was, relied on
the attraction of moths to sweet and fermenting fluids. Old books had many
exotic recipes, but a good basic mixture would be a thick mush of rotting
bananas, brown sugar, and dark beer boiled for ten minutes, to which is added
a splash of rum or essence of pear drops (amyl acetate). The mixture is daubed
on posts and tree trunks after dark. The mixture can also be soaked on to thick
ropes, which are hung up.
Absolute collecting techniques

Hand searching Simply getting down on the ground and looking for in-
sects is a much under-rated technique. There are some habitats where no
other sampling method will yield results. The most valuable (and often ne-
glected) aspect of hand searching is that you might find out what the animals
are doing. Mass collection techniques such as sweeping, beating, or fogging
let you know what is present and in what numbers, but that is all they tell you.
Close observation will reveal much more.
Imagine you need to look at grass tussocks where a particular plant bug
lives. Sweeping and suction sampling may not provide much due to the dense
growth and the foliage might be permanently wet in any case. The bugs in
question live right on the surface of the soil and the only way to get them is
go after them with a pooter. A good pooter or aspirator is one of the most
useful bits of kit that you can have, and, although designs differ widely, they
mostly rely on the operator sucking. There are special blow-pooters for medi-
cal, agricultural, and forensic sampling, which, sensibly, rely on the Bernoulli
or venturi principle (you blow rather than suck). One of the simplest general
purpose sucking designs uses a Sterilin™ universal specimen tube (approxi-
mately 39 × 25 mm and 30-cm3 capacity). The tube is made from polystyrene
so do not use solvents such as ethyl acetate near them. A bung with two holes
drilled through it and short lengths of plastic tubing and pipe, and a small
piece of terylene net secured over the end of the ‘suck’ tube to prevent insects
from entering the operator’s mouth, are all you need to complete the pooter.
The diameter of the tubing is important. Too small and you will only be able
to collect small species. Too big and you will need a lot of suck to get anything
up the tube. Having collected what you want, the pooter tube can be tapped
to shake all the insects to the bottom and then the bung pulled out and re-
placed with a screw-top. Large insects can be grasped using a pair of forceps,
A pooter. while very delicate insects, if not pootered, can be picked up on the end of an
alcohol-moistened paint brush.
If pooters are used for long periods, there is a risk that minute particles,
micro-organisms, and fungal spores could be inhaled, which might lead to
allergies and chest infections.
Another situation where hand searching is useful is where arthropods have
to be sampled from rock surfaces, particularly in caves or lava tubes. In these
habitats, populations of animals are often sparse, and you can cover a lot of
ground with a pooter and a pair of forceps. Different sites can be compared
in a quantitative manner by introducing a time element.
Emergence traps and rearing Collecting insects by trapping them as they

emerge, usually as adults from pupae or cocoons following larval develop-
ment, diapause, aestivation, or overwintering, may be essential if you are
going to do anything more than compile a simple species list. Rearing in-
sects from leaf mines and stems, from under bark, from dead wood, from
parasitized insects, from corpses, from macro-fungi, from the soil, and so on,
may be the only practical way to collect many species. Different sorts of trap
will be needed for each habitat/micro-habitat. At its simplest, an emergence
trap or rearing container may be a cotton wool-plugged or muslin-covered
tube or tub. Collecting insects in this way will provide much more useful
biological information than general collecting techniques. For instance, beat-
ing the foliage of an oak tree may yield many species of gall wasp, but galls
placed inside simple, muslin-covered containers will yield the gall causers,
other gall occupants, and their parasites. It will also be possible to associate
and preserve larval and pupal exuviae (shed skins) with the emerging adults.
Deeming (1993) gives a good account of the simple techniques he uses to
COLLECTING 291
rear shoot flies belonging to several families. The same techniques could be
equally well applied to the rearing of other insect groups, such as parasitic
hymenopterans.
Many insect species emerge from soil or litter at the beginning of the grow-
ing season and are positively phototactic. Emergence traps are easily made
from any suitable upturned opaque, usually black, container dug into the
substrate and fitted with a transparent collecting bottle at the apex. Insects
emerging from aquatic habitats can also be collected in emergence traps.
Emergence traps can be considered an absolute collecting method pro-
vided you sample from a known area or volume of substrate or habitat.
Making simple and effective emergence traps to suit your particular purpose
may be preferable (and certainly cheaper) to buying a ready-made design.
A large, general-purpose model often used for processing large amounts of
wood, litter, flood debris, compost, etc., is the Owen Emergence Trap. Mea-
sured quantities of material are placed inside the trap and left. Emerging
insects should eventually find their way to the collecting bottle at the top.
Some traps containing rotting wood have continued to yield material for up
to two years.
Suction sampling From the 1950s onwards, the need for more quantita-
tive methods of sampling insects led to the development of a number of
suction devices. The commonest was the petrol engine-driven D-Vac, or,
more correctly, Dietrick suction insect sampler. Originally developed for use
in agricultural crop systems, the D-Vac became widely used in ecological
entomology, and virtually the regulation piece of equipment for sampling
invertebrates in grassland habitats. The advantages of using a standard ma-
chine with a known sampling area and sampling effort were that different
sites and different studies could be compared in a way that had been impos-
sible before. The disadvantages, however, were many and varied. There was
a tendency, among some users, to assume that everything was caught, but
the machines were very variable in their efficiency and few calibration stud-
ies were ever carried out. Good practice involved D-Vac-ing for a number
of timed sucks combined with careful hand searching afterwards to ensure
that nothing had been missed. D-Vacs were very expensive, never very easy
or comfortable to use, and needed constant maintenance. Despite modifica-
tions in the 1960s, these D-vacs are no longer available commercially. The
advantages of suction sampling remain the same and some devices are still
available. The high cost of commercial machines has resulted in a number of
people making very simple modifications to leaf-blowers and similar items
of garden machinery (Wilson et al., 1993; Stewart and Wright, 1995). Studies
have shown that these home-made samplers do a very good job indeed, but,
like all other designs, they will not sample everything with equal efficiency.
You must calibrate your sampling in some way to estimate what percentage
of organisms you are collecting. This percentage will vary greatly with insect
body size, the sort of habitat you are studying, and the sampling time. Work
out the cross-sectional area of the suction tube and you will be able to simply
calculate the area you have sampled. For low vegetation a sampling unit might
be 25 random timed sucks pooled from within a given area of the habitat. Al-
ternatively, you could use a cylinder of known area to isolate the vegetation
to be bug-vacuumed. Samples can be everted into plastic bags as described in
the section on sweep nets.
Extraction techniques Separating small invertebrates from the substrate
can be a difficult and time-consuming process, but scores heavily on being an
absolute method. Measured volumes, weights, or areas of sample will yield
quantitative data. Different techniques are needed for different substrates, but
the simplest form of extraction tool is a sieve. Here, gravity and agitation will
only effect a partial separation. Different gauges of sieve (e.g., 10-, 5-, and
2-mm meshes) might help, but animals may hold on to or hide inside large
debris and very small specimens will still need to be separated from fine de-
bris. It is generally better to persuade the animals to leave the substrate under
their own steam, and this principle is employed in a variety of extraction de-
vices. The obvious advantage is that the samples obtained are relatively clean
and require little further processing. A disadvantage is that the equipment can
sometimes be bulky and, in cases where a heat or light source is required, dif-
ficult to arrange in the field. It is worth bearing in mind that the smaller and
more delicate the animals of interest, the more difficult it will be to extract
them efficiently.
Extraction funnels such as Berlese/Tullgren designs are suitable for the
extraction of insects and other moderately robust specimens from leaf litter,
crumbled soil, decaying wood, or even fresh foliage. There are very many vari-
ations of the same theme. The measured sample is placed on a mesh or sieve
over a funnel, which leads to a collecting bottle containing alcohol (or dry
for live collection). A (preferably boxed-in) source of light and heat source
is placed directly above the sample causing the material to dry out. Animals
migrate downwards through the sample. Rapid drying should be avoided, as
this will kill smaller organisms before they escape from the sample.
An excellent piece of equipment for extracting invertebrates from moss,
dead wood, litter, and even things like bird nests in the field, is the Winkler
bag (also known as the Winkler extractor or Winkler apparatus). The bags are
commercially available, and Owen (1987) gives a full account of the use and
construction of this lightweight and highly effective apparatus. The Winkler
bag is relatively cheap to make and, unlike other devices, does not require an
artificial heat source to make it work. In essence, a wire-braced, box-shaped,
calico bag is used to enclose measured samples of debris contained inside
coarse netting. The bottom section of the bag tapers to a collecting bottle,
which may be used dry or with alcohol. The top of the bag is tied up and the
whole thing is hung up under cover to dry out gradually. Animals from sam-
ples that start fairly dry will be extracted in a few days, but wet samples may
take more than a fortnight. To speed up the process it is advisable to occasion-
ally take the samples from the inner bags and mix them to redistribute the
wet bits. Winkler bags are particularly good at collecting small beetles, ants,
hemipterans, parasitic wasps, flies, and cockroaches, and have resulted in the
discovery of countless new species that would have been hard or impossible
to collect with other methods.
COLLECTING 293
Canopy techniques All that is known to date about collecting arthropods

from forest canopies can be found in two books: Forest Canopies (Low-
man and Nadkarni, 1995) and Canopy Arthropods (Stork, Didham, and Adis,
1996). These two books, which have extensive bibliographies, provide up-to-
date information on the biology of canopy arthropods, and how to sample
and study them.
Essentially, you can either go up to collect material using a variety of ap-
proaches ranging from climbing, ladders, and cranes, to net rafts placed on
the canopy by means of a dirigible (airship, balloon), or you can get the mate-
rial to fall down using insecticides. Somewhere between these two extremes
are things like composite traps, which are hoisted into the canopy (Basset,
1988).
The variety of insecticidal knockdown techniques that have been used in
various parts of the world have a few things in common. Collecting trays of
a known area are set up on the ground below or suspended from vegetation
below the tree to be sampled. The trays, which can be conical or pyramidal, are
best made from smooth, close-woven material such as rip-stop nylon. They
should ideally have an integral collecting bottle at the bottom to speed up
collection and minimize the loss of material. Under small trees or bushes,
large plastic washing-up bowls can be used. A fast-acting insecticide, such as
a synthetic pyrethroid, is blown into the foliage in the form of a fine mist or
fog. Misting or fogging is commonly done early in the morning when there is
no wind. Obviously, the longer the drop the greater the risk of a gust of wind
blowing small insects clear of the trays. Wide treatment of the canopy around
the area delimited by the trays below will reduce this error. Early samples are
also thought to be more representative of the canopy fauna. Drop time is an
important factor. Most studies have shown that 60–90 minutes is more than
adequate. Samples should be preserved in alcohol as soon as is practicable.
Clearly, some techniques are better for some things than others. Hand
searching in canopy foliage tends to miss small things, whereas insecticidal
fogging or mist-blowing tends to under-record large things. Another prob-
lem with canopy sampling is that, although there have been a number of
well-documented studies, they have used a wide range of sampling meth-
ods. Fogging, mist blowing, fumigation, and gassing are not comparable, and
the use of different insecticides and even different formulations of the same
insecticide have added to the difficulties.
Aquatic and taxa-specific sampling techniques

Chapter 5 in Ecological Methods (Southwood and Henderson, 2000) and
chapter 10 in Aquatic Insects (Williams and Feltmate, 1992) are compul-
sory reading for anyone sampling freshwater habitats. In the main, insects
can be simply collected from freshwater habitats using any number of hand
net designs. Catches are simply tipped into white sorting trays or through
sieves of various mesh sizes (0.5 mm or greater is fine for insects and lar-
vae) and the species required removed for preservation in alcohol. Hand
nets are long-handled, and for deep rivers or lakes should be able to take
additional (screw-on) handle sections. The net bag should be made from
strong polyester or nylon 1-mm mesh. The most common hand nets have
circular or D-shaped mouths. Insects are caught by passing the net through
the water or vegetation. Hand nets can also be used downstream to collect
anything dislodged from vegetation or stones. Drift nets are rectangular-
mouthed, long, tapering nets, which are staked or weighted down to collect
organisms from shallow streams. A Surber sampler is a drift net with side
screens fitted to a quadrat of known area (usually 0.1 m2 ), which can be used
to sample from shallow streams where the flow rate is less than 10 cm per
second. Stones and plants within the quadrat are kicked, jarred, or brushed,
the organisms dislodged, and are carried into the net by the current.
While it is very easy to collect insects in a relative manner from aquatic
habitats, difficulties occur when trying to take reliable samples from a known
volume or area of habitat. When collecting from open water you can easily
calculate what volume of water has been sampled by your net. A number of
sampling cages, cylinders, and grabs have been made which enclose a known
volume of vegetation, bottom substrate, or sediment (Southwood and Hen-
derson, 2000). In deeper water, dredges and trawls can be used to sample a
known unit of substrate and open water habitat respectively.
Emergence traps, either fixed to the bottom or floating on the surface,
can be used to collect insects that emerge as adults from freshwater habi-
tats. Various sorts of artificial substrate (plastic mesh, discs, etc.) or natural
substrate (leaf packs, stones) have been used to encourage colonization by in-
vertebrates. These types of sampler are left in place for a specified time and
then retrieved.
There are a number of useful techniques, mainly specific to particular taxa,
which do not easily fit into the categories already mentioned.
Fleas Fleas (Siphonaptera) are specialized ectoparasites of mammals and
birds. Unlike parasitic lice (Phthiraptera), they do not spend their entire life
attached to the host fur or feathers. Flea eggs are shed in the nest or lair of
the host and the larvae are free-living and eat organic debris and the drop-
pings of the adult fleas. Fleas and other ectoparasitic insects, such as lice and
some Diptera, may be obtained by searching, combing, or fumigation (using
chloroform or ether) of live or dead hosts. Remember to find out about any
legislation in this area. You may need licenses to trap and handle live birds or
mammals and it is essential to get training, especially if the use of anaesthesia
is planned. Much larger numbers of fleas and other associated flies and beetles
can be found in nests and lairs, which may be located visually, but mammals
might only be active after dark or escape through vegetation after being re-
leased from live traps. Boonstra and Craine (1986) describe an inexpensive
spool-and-line technique for following small mammals to their nests. Essen-
tially, a thin thread is attached to a live trapped animals usually by gluing it to
the fur. When released, the animal pulls thread from a spool, which allows the
researcher to locate the nest. Early designs allowed tracking of up to 200 m,
but improvements have increased this to more than 1500 m, depending on
vegetation. Hawkins and Macdonald (1992) evaluate the technique for use
with larger mammals.
COLLECTING 295
Invertebrates living in nest material can be extracted using

Berlese/Tullgren funnels or Winkler bags. These methods will also yield
flea larvae, which would not be present on the body of the host. We are
grateful to R. S. (Bob) George for the following description of his technique
for collecting fleas from nest material. The equipment needed is a large,
steep-sided bowl, specimen tubes containing 70% alcohol, a dissecting
needle, and a large sheet of white paper. Before examination, nests must be
kept separately in sealed boxes, tins, or polythene bags. The collector should
wear a white shirt and work with rolled-up sleeves. A small amount of the
nest material is placed in the bowl, which should stand in the middle of the
white sheet. Adult fleas are picked up on the end of the alcohol-moistened
dissecting needle and transferred to a tube. The material will also contain
flea cocoons. These can be opened at the truncated end and, usually, adult
fleas will emerge with a rush only to be collected. When no more adults can
be found the material can be transferred to a sealed box either to breed out
any remaining larvae or to allow a re-examination. Any fleas jumping free
from the bowl are easily seen against the white paper, shirt, or bare arms
and collected. This technique will produce many times more specimens than
funnel extractions and is essential if you need quantitative data.
Butterflies If you can recognize butterfly species you encounter, then you
need not kill any. However, the butterfly transect walk techniques described
by Pollard and Yates (1993) does need accurate identifications. Mark-capture-
recapture techniques can be used to obtain absolute population estimates.
Bees Bees can be most easily collected at flowers but try looking for nest
sites in south or south-west facing exposures of bare earth or sand. In savan-
nahs and deserts, some species will often nest in the vertical faces of dried-up
riverbeds. These, and other species of bees that use the pith-filled cavities of
plant stems, abandoned wood-boring beetle burrows, and empty snail shells,
can be trapped in artificial nests. Trap nests can take the form of bundles of
bamboo, pithy plant stems, or even wide drinks straws tied together; simi-
larly, blocks of softwood drilled along the grain using a variety of hole sizes
from 2 mm to 2 cm in diameter can be used. If appropriate trap nest designs
are used, bees will start to colonize within hours of them being positioned in
the field, and occupancy rates can be as high as 50–70%.
Termites In tropical habitats, termites are vitally important as decomposers

and soil conditioners, and are significant pests. There are also many interest-
ing and largely unstudied species that live as inquilines in termite colonies.
Numerically abundant (up to 10,000 individuals per m2 ) the biomass of ter-
mites may outweigh all herbivorous vertebrate animal species put together.
Despite this, termites do not appear a great deal in samples collected us-
ing standard entomological techniques. Even insecticidal techniques will not
sample termites nesting in tree canopies. If sprayed or fogged, they will sim-
ply retreat inside their tunnels and nests. Sampling obvious structures, such
as mounds, will provide only one-tenth of the termite abundance in some
habitats. Eggleton and Bignell (1995) consider the various strengths and
weaknesses of a variety of termite sampling procedures and develop a stan-

dard, stratified sampling protocol designed to take into account the patchy
distribution and specialization of termite assemblages.
Collecting from live animals Blood feeding and other types of insect may
be collected by using live animals of various sorts, including humans, as bait.
There are a number of obvious hazards. Large animals, even domesticated
ones, can be dangerous at both ends. Tethered or not, approach from the front.
Flies can be caught directly as they feed, but only a very tame animal will
remain calm while an excited entomologist swipes at its hind quarters with a
net. Small animals, such as rodents and bats, can give painful bites and may
carry diseases such as rabies. Get training in handling live animals and wear
protective clothing if required.
Killing methods and data recording

Insects should not be killed needlessly. However, the correct identification
of many species may require the examination of minute body parts un-
der a stereobinocular microscope or the dissection and preservation of the
genitalia. This section discusses killing methods and data recording.
Killing methods
There are no clear guidelines about what constitutes the best or most humane
way to kill insects, but common sense should tell you that the quicker the
better is a good principle, as long as the specimens are intact and useable at the
end (see Martin, 1977; Arnett, 1985, and Davies and Stork, 1996). When using
chemicals, thoroughly familiarize yourself with the hazards and precautions.
Appropriate safety procedures must be observed at all times, and bottles and
jars should be labelled correctly.
A traditional method to kill insects is the cyanide killing jar. The technique
is still favoured by some professional lepidopterists and orthopterists because
it kills more quickly than other methods, keeps the specimens dry, and does
not cause colour changes (ethyl acetate fumes, for instance, can cause some
green pigments to fade). Glass or preferably high-density polyethylene jars
for field use are made in advance and with careful preparation may give
many months of service (Arnett, 1985). A typical jar uses a packed, 5-mm
layer of powdered potassium cyanide crystals under a 1-cm-thick layer of
sawdust and topped with a 1-cm layer of plaster of Paris. The plaster should
be allowed to dry out thoroughly before the jar is ready for use. Some peo-
ple cannot smell cyanide, and jars should be made under a fume hood or in
a very well-ventilated space. Contaminated equipment should be immersed
in dilute sodium hydroxide solution to which is added an excess of ferrous
sulphate solution. After an hour, rinse and flush away with an excess of wa-
ter. In its dry state, a cyanide jar will have a shelf life of one or two years. Once
in the field, a few drops of water soaked into the plaster layer will react with
the potassium cyanide to produce cyanide gas. Full production of the gas will
KILLING METHODS AND DATA RECORDING 297
build up over a day or two and should keep going for many weeks. An old-
fashioned, but very effective, technique still used by some researchers today is
the use of young, crushed laurel leaves wrapped in muslin inside a jar or bot-
tle. Small quantities of cyanide gas are released, and the humid atmosphere
keeps specimens soft and relaxed until they can be pinned.
Butterflies caught in a net can be quickly incapacitated by a gentle pinch
across the thorax using the nails of the thumb and forefinger. The technique
allows the transfer of specimens to a paper envelope with the risk of them
flapping around and losing wing scales very much reduced. The envelopes
can be stored flat in a plastic tub temporarily before being placed in an ethyl
acetate jar. Pinching does not work very well on moths.
When using a Malaise trap the catch is generally going to be killed and
stored in alcohol anyway. For many other mass collection techniques, such
as suction sampling or sweeping (the catch will be live), or mist blowing (the
catch will already be dead), the sheer volume of material means that simple
immersion in alcohol is the only practicable combined killing and storage
method.
Very large insects, such as dung beetles, can be killed very quickly by the
injection of small quantity of saturated oxalic acid solution into the thorax
or abdomen through the intersegmental membranes, but take care because,
apart from being poisonous, oxalic acid is a white, crystalline powder which
might arouse suspicion at customs. Dung beetles and other fairly robust
species can also be killed very quickly by dropping them into boiling water
(which also cleans them).
The vapours of various chemicals such as benzene, carbon tetrachloride,
ether, and ammonia have been used in the past to kill insects. Many of them
are dangerous and some highly inflammable, but probably the safest alter-
native to cyanide in the field is ethyl acetate. The fumes of ethyl acetate kill
more slowly than cyanide and are still flammable. Killing bottles or tubes can
be made with a layer of plaster of Paris at the bottom onto which a few drops
of ethyl acetate can be dropped. More simply, a layer of cotton wool with a
few pieces of filter paper on top can be arranged inside a polythene bottle.
Bulk killing bottles should have a capacity of at least 500 ml and a wide top to
allow you to get large specimens inside easily. Specimen tubes of strong glass
or polythene are best for killing small insects. A wick of filter paper with a
drop of ethyl acetate is held in place by the push fit top. Glass tubes are ac-
ceptable here and have the advantage of being transparent. It is best to have a
small wad of tissue paper inside the killing chamber to soak up excess mois-
ture. Ethyl acetate and acetone (sometimes used as a substitute and widely
available as nail varnish remover) will dissolve styrene plastics. When dead,
the insect should be removed and pinned or stored as required.
Data recording
Specimens collected must have adequate data associated with them. Informa-
tion on where, when, who, and how any particular insect was caught should
be as full as practicable. For instance, ‘Tanzania, August 1996’ is clearly inad-
equate as compared to ‘collected at UV light, 12 August 1996, Ibaya Camp,
Mkomazi Game Reserve, Tanzania, G. C. McGavin’. In the field the data

should be kept closely with the specimens (attached to it or inside the tube
or envelope). Access to GPS (global positioning systems) enables collecting
sites to be pinpointed as never before.
Do not write on the outside of tubes or on tube tops. Tube tops can get
accidentally switched and labels can fall or get rubbed off. Labels for putting
inside tubes of material stored in alcohol is best done on good-quality paper
with pencil or waterproof drawing ink.
Never think that you will recall where or when a particular specimen was
caught when you get back from the field. Specimens without proper data are
completely useless.
Specimen preservation
In the field, the simplest techniques are drying or storing in alcohol. Most ro-
bust insects, such as beetles and bugs, can be simply dried in air and packed
in layers of cellulose wadding or tissue. Delicate or soft-bodied species and
spiders should be preserved in 70–80% alcohol (ethanol). Some coleopter-
ists preserve small beetles in a 4% solution of glacial acetic acid until they are
dried and set. If kept cool, the material should be fine for many months. Moths
and butterflies should never be stored in alcohol and are best dried in small
paper envelopes or folded paper triangles, or set and pinned directly into
boxes. Set specimens take up a lot of room and are difficult to transport safely.
A problem can arise with large fat-bodied species, such as some grasshop-
pers and mantids. The large quantity of internal tissue can take a long time to
dry properly, and decay may take place. A good technique is to stuff the ab-
dominal and thoracic cavity with cotton wool. This is not as difficult as it may
sound. Using a pair of fine dissection scissors make a slit in the ventral surface
of the abdomen. You do not need to open the abdomen up from stem to stern.
Remove the abdominal contents with a pair of fine watchmakers forceps. Be
sure to reach up into the thoracic cavity to remove the front portion of the
gut. Do not rip the guts from the rear end of the animal as you may damage
or remove parts of the genitalia, which may be of value in identification at a
later date. Use fine scissors to cut through the rectum close to the end of the
abdomen. You do not have to remove every shred of soft material from the
inside. Starting at the thorax, use fine forceps to push little bits of cotton wool
into the cavity. When finished, the abdomen should be as close to the shape it
is in real life as possible. You do not need to sew up the incision, as the cuticle
will dry in position without being held in place.
The most useful field preservation technique for adult insects that we, and
many others, have successfully used for years involves the use of chlorocresol
(more correctly 4-chloro-m-cresol) (see Tindale, 1961). A small quantity (a
level teaspoonful) is sprinkled in the bottom of an airtight polythene food
storage box and covered with a layer of tissue or wadding. On top of this are
laid specimens in labelled, paper envelopes or layered in tissue. Some crys-
tals of chlorocresol should be sprinkled on top of every tenth layer or so. A
LONG-TERM PRESERVATION 299
crumpled tissue or two keeps the layers in place until the box is full, when
a final generous sprinkle of chlorocresol should be added. The chlorocresol
acts as a fungicide and bactericide, and keeps the insects moist and relaxed for
months, so that when you come to mount them the job is very easy. Air-dried
insects must be first relaxed in very warm humid conditions before they can
be pinned and have their appendages arranged. The chlorocresol technique
avoids all this, and even butterflies and moths treated in this way will remain
‘settable’ and in good condition for a long time.
Long-term preservation
For some studies involving large amounts of material, it might be simplest
to leave specimens in alcohol until such times when they really need to
be prepared in a more specific manner. However, spirit-preserved material
does require maintenance. For long-term storage, critical specimens should,
ideally, be kept in a freezer. This will reduce the need for topping up the al-
cohol and, as an extra bonus, keeps internal organs in absolutely first-class
condition for decades.
Usually, identification and systematic study of specimens will require them
to be mounted dry. Pinning insects enables them to be handled, examined,
and stored safely. Sometimes inappropriate mounting can destroy the very
features required for proper identification. General insect mounting, preser-
vation, and curatorial techniques can be gleaned from a variety of sources,
but among the best are Martin (1977) and Walker and Crosby (1988). Carter
and Walker (1999) deal with all aspect of the physical care of botanical and
zoological collections and give much practical guidance over the whole field
of natural history curation.
Publications giving specialist techniques for particular orders of in-
sects, such as the Dictyoptera (specifically, termites) (Krishna and Weesner,
1970), the Thysanoptera (Lewis, 1973), the Hemiptera (Dolling, 1991; Mc-
Gavin, 1993), the Coleoptera (Cooter, 1991), the Diptera (Stubbs and Chan-
dler, 1978), the Lepidoptera (Dickson, 1992), the Hymenoptera (Gauld and
Bolton, 1988), and the stoneflies, mayflies, and caddisflies (Macan, 1982),
should be consulted where appropriate. Many of these books also give details
of collecting methods.
Most robust-bodied specimens can be simply taken out of alcohol, dried,
and then mounted in the manner appropriate to the taxon concerned, but
small or delicate specimens will need some treatment—otherwise, they
will shrivel up and become useless. There are some simple techniques that
will allow these specimens to be taken out of alcohol, dried, and mounted
without bits or all of the body collapsing. We are grateful to Simon van Noort
of the South African Museum in Cape Town for the following recipe, which
can be used for a range of small or weakly sclerotized insects, especially
parasitic wasps.
The quickest and easiest approach, particularly where there are large
numbers of specimens, is to treat the whole sample in one go. Pipette the
specimens in alcohol onto a square piece of fairly stiff card, the edges of which
are folded up to make a miniature sort of tray. Arrange them in the position
required for subsequent mounting at this stage. Place the card in a petri dish
lid to keep the specimens saturated in alcohol while getting them into po-
sition. The specimens are best on their right-hand sides with wings folded
flat above the body, which takes a bit of time, but is worth the effort, so that
when it comes to mounting it is simply a matter of placing the specimen on a
blob of glue on a card point. Once the specimens are arranged, drain off the
excess alcohol by lifting up the card and letting it run off before placing the
card tray into an acetone-saturated environment. Acetone is soaked liberally
on to cotton wool at the bottom of a sealable acetone-proof container and
the cards, bearing their arranged specimens that are still wet through with
alcohol, should be placed horizontally or vertically, clear of the acetone, for a
minimum of six hours. To dry the material, lift out the card tray, which is now
saturated with acetone, and place it in a petri dish under a 60-watt desk lamp
(close to the bulb) for 20–30 minutes. Once the specimens are dry, be careful
not to breathe too hard over them, or they will end up all over the place. Once
dry, they do not need to be mounted straight away and can be kept quite safe
in a petri dish in a pest-free environment. The method generally gives good
results but sometimes specimens may still collapse, even though others in the
same sample are fine.
Another technique for the recovery of non-robust specimens from alcohol
storage is particularly useful for small flies or bugs. Start by pinning the speci-
mens direct from the alcohol. Stand the pinned specimens in a tube filled with
2-ethoxy-ethanol overnight (12 hours minimum). Transfer to a tube (glass)
filled with ethyl acetate (1.5 hours for small species, 3 hours for larger species).
When the specimen is removed, the wings, particularly, will stick together or
crease up. Dab off excess ethyl acetate with a tissue and blow rapidly on the
specimen to release the wings. If the wings do not immediately spring out,
tease them out with a fine pin or forceps (J. Ismay, personal communication).
Bees that have been stored in alcohol cannot be simply taken out and dried,
because the dense hair covering becomes matted and flat. A near life-like
dried specimen can be prepared in the following manner. Remove the bee
from alcohol, dab dry with tissue to get rid of most of the fluid, and immerse
in 2-ethoxy-ethanol for 24 hours. Remove excess by dabbing with a tissue. Pin
specimen in the normal manner, placing legs and wings in the desired posi-
tions. The male genitalia may be exposed at this stage using a small, hooked
pin. Soak the pinned specimen in ethyl acetate for 24 hours. On removal, dry
immediately with a small hair drier from behind. The hairs will fluff up and
remain erect (C. O’Toole, personal communication).
Should important spirit-preserved material ever dry out, it can be rescued
by soaking in a 2% solution of trisodium orthophosphate. The rehydration
rate depends a great deal on the size and condition of the specimens, but it
is best to check them every hour or so. A good indication that the process
is complete is when the specimens sink to the bottom of the container. The
LONG-TERM PRESERVATION 301
specimens should then be thoroughly washed in distilled water before being

put back into 70% alcohol.
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Subject index
Notes Tables and figures are indicated by an italic t or f following the figure.
Abedus sp., sexual Malaise traps 297 Xylocopinae 270

reproduction 146 pitfall traps 286 Anthophorinae (digger
absolute collecting specimen preservation 298 bee) 270
techniques 289–93 alderflies (Megaloptera) 156, anthrax, fly vectors 220t
Acanthoscelides obtectus 185 161–4, 162f, 163f Antipaluria caribbeana 118f
Acercaria Aleocharinae 182–3 ant-lions (Myrmeleonti-
(Paraneoptera) 122–55 Aleyrodidae, pests as 148 dae) 169–70
acetone, specimen Alienopteridae 103 Antliophora 196–226
preservation 300 alkaloid chemical defences 26 Diptera see Diptera (true flies)
Acherontia atropos (Death’s- Amblycera 128t Mecoptera 197–201, 197f,
Head Hawk animal hosts 128 198f, 199f
Moth) 251 mouthparts 128–9 Siphonaptera see
Achroia grisella (Lesser Wax ambrosia beetles (Platypodidae), Siphonaptera (fleas)
Moth) 241 fungal interactions 24–5 ant-nest beetle (Paussinae) 182
acoustic tracheae, American Cockroach see ants see Formicoidea;
Orthoptera 78 Periplaneta americana Hymenoptera
Acrididae (grasshoppers, (American Cockroach) Apicotermitinae 93
locusts) 78 Ametropis 47–8 Apini (honey bees) 270–3
model systems as 82 Amitermes meridionalis colonies 271–2
pests as 81 (Magnetic Termite) 91 pollination 272–3
swarms 81 ammonia vapour, killing Apis mellifera (Western Honey
Acrocinus longimanus methods 297 Bee) 271
(Harlequin Beetle) 179 Amphiesmenoptera 227–54 Apocrita 259f, 259t
Aculeata 258, 259f, 259t, 266–7 Lepidoptera see Lepidoptera Aculeata 258
Apoidea see Apoidea (bees) (butterflies and moths) Stephanoidea 259t, 263
Formicoidea see Formicoidea Trichoptera see Trichoptera Trigonaloidea 259t, 263–4
Scolioidea 259t (caddisflies) apodemes 14
Thynnoidea 259t Anabris simplex (Mormon apodous grubs 180
Tiphioidea 259t, 266–7 Cricket) 79 Apoica pallens 257f
Vespodea see Vespoidea Anacridium sp., pests as 81 Apoidea (bees) 27–8, 258, 259t,
Acyrthosiphon pisum (Pea Andrenidae (mining bees) 270 269–73
Aphid) 143 Annulipalpia 230, 231t Anthophila see Anthophila
Adephaga 178t Anobium punctatum (furniture collecting 295
adhesive hair pads, herbivorous or cabinet beetle) 186 long-term specimen
insects 25 Anopheles sp., human disease preservation 300
aedeagus (penis) 31 vectors as 220t, 221–2 aquatic sampling
Aedes sp., human disease 220t, Anoplura 128t techniques 293–6
221–2 Echinophthiriidae 131–2 Arachnids (spiders) 5
Aeolothripidae (banded mouthparts 129–30 Arachnocampa luminosa 217
thrips) 153–4 Antarctoperlaria 70 Archaeognatha (bristletails) 36,
Aeolothrips fasciatus 154 Anthonomus grandis (Cotton 38–40, 38f, 39f
aerial nets 280 Boll Weevil) 187 flight 19–20
Afrotrogla sp. 126–7 Anthophila 269–73 Archipsocopsis sp. 127
Agamopsyche 242–3 Andrenidae 270 Archostemata 178t
Agaonidae (fig wasps) 27, 265 Anthophorinae 270 Archotermopsidae 94t
Agathiphagidae 240 Apini 270–3 Arctiidae, acoustic signals 241
Agulla 159 Bombini 270 Arctiinae 251–2
alcohol Halictidae 269–70 Arctocephalus ogmorhini 131
long-term specimen Megachilidae 270 Arctoperlaria 70
preservation 299, 300 Nomadinae 270 Arixeniid earwigs 62
304 SUBJECT INDEX
army ants (Dorylinae) 269 Blattodea (cockroaches and bush hoppers

Asilidae (robber fly) 213 termites) 85–97, 85f, (Pyrgomorphidae) 78t
Asparagus Beetle (Criocerus 86f bush locust
asparagi) 185 cockroaches see cockroaches (Pyrgomorphidae) 78t
assassin bugs (Reduviidae) 136f termites see termites butterflies see Lepidoptera
Atelura formicaria 42–3 blood–brain barrier (butterflies and moths)
Atlantoraphidia (perineurium) 17–18, butterfly nets 280, 282f
maculicollis 158f 19f
Auchenorrhyncha 134, 137f, blow flies (Calliphoridae) 213
137t, 138f Body Louse (Pediculus humanus cabinet beetle (Anobium
digestive tract 140f humanus) 130, 130f punctatum) 186
morphology 137 body weight, species numbers caddisflies see Trichoptera
mouthparts 138 vs. 9f (caddisflies)
substrate-borne Bombadier beetles Caelifera 76
vibrations 145 (Brachininae) 181–2 Acrididae (grasshoppers,
Austroperlidae 70 Bombini (bumble bees) 270 locusts) 78t
avermectins 184 Bombycoidea 249–51 mating 79–81
Bombyx mori (silk principal families 78
moth) 249–50 Pyrgomorphidae (bush
bacteria, symbiosis 24 booklice see Psocodea (barklice, hoppers, bush
Baetidae 47, 47t booklice and true lice) locusts) 78t
baited traps 288–9 Boreidae (snow scorpi- Tetrigidae (pygmy
pitfall traps 287 onflies) 198, 199, locusts) 78t
banded thrips 200 Caenidae 47t
(Aeolothripidae) 153–4 Bot flies (Oestridae) 221 Calliphoridae (blow flies) 213
bark-boring beetles see Bovicola bovis (Cattle Biting Callistoleon illustris 170
Scolytidae (bark-boring Louse) 129 Callistoleon manselli 170
beetles) Bovicola equi (Horse Biting Callosobruchus sp. 185
barklice see Psocodea (barklice, Louse) 129 Calocoris quadripunctatus 134f
booklice and true lice) Brachininae (Bombadier Calyptra eustrigata 243
barnacles 6 beetles) 181–2 campaniform sensilla 29
basement membrane, Brachycera 213, 214t campodeiform larvae 180
cuticle 13, 15f Orthorrhapha 213, 214t canopy techniques 293–4
Batesian mimicry 240 sexual reproduction 216 Canthon, baited traps 288
beating trays 281–2, 282f Tabanidae (horse fly) 213, Carabidae (ground beetles) 181,
bed bugs (Cimicidae) 146–7 219–20 181f
bees see Apoidea (bees); see also Cyclorrhapha Carboniferous period
Hymenoptera Braconidae 263–4 Ephemeroptera (mayflies) 45
beeswax 271 Bradyporus sp. 80 herbivorous insects 26
beetles see Coleoptera (beetles) brain 16, 18, 20, 100, 192, termites 96
Belostoma sp., sexual 212–13, 224, 265 winged insects 44
reproduction 146 Branchiopoda (water-fleas, carbon tetrachloride
benzene vapour, killing fairy shrimps, and brine vapour 297
methods 297 shrimps) 5 carpels 27
Berlese funnels 292 Brassiolinae 248–9 carpenter bees
flea collection 295 brine shrimps (Xylocopinae) 270
Bethylidae 266 (Branchiopoda) 5 carpenter moths
biconical traps, tsetse flies 224 bristletails see Archaeognatha (Cossidae) 245–6
biodiversity crisis viii–ix, 32 (bristletails) Carrikerella sp. 101
Bittacidae (hangingflies) 198 brown lacewing Case-bearing Clothes Moth
Bittacus (hangingflies) 200 (Hemerobiidae) 169 (Tinea pellionella) 245
Blaberidae 88f Bruce, David 223–4 castes
Black Death (bubonic Bruchus pisorum 185 Formicoidea 268
plague) 206 bubonic plague (Black termites 91, 94
Blattaria see Blattodea Death) 206 Cat Flea see Ctenocephalides felis
(cockroaches and Buckleria paludum 237f (Cat Flea)
termites) bumble bees (Bombini) Cattle Biting Louse (Bovicola
Blattella germanica (German 270 bovis) 129
Cockroach) 90 Buprestidae (jewel cave crickets (Rhaphidophori-
Blattellidae 87, 88f beetle) 184–5, dae) 75t
Blattidae 88f 185f cell size, size constraints 11–12
SUBJECT INDEX 305
cellulose digestion, termites 93 Cimicidae (bed bugs) 146–7 Paussinae 182

centipedes (Chilopoda) 7 circulatory system 29 pests as 185
central nervous system claws, herbivorous insects 25 Polyphaga 178t, 182–3
(CNS) 18 Clematoscenea sp. 124f Pselaphinae 182–3
Cephalocarida 5 click beetles (Elateridae) 174f Scarabaeidae 183–5
Cephoidea 259t climate change, malaria Scolytidae 187
habitat 262 effects 222 sexual reproduction 179–80
Cerambycidae (longhorn Cloudsley-Thompson, John L. 8 species numbers 177–8
beetle) 179, 185, 185f coastal water habitats 31–2 Staphylinidae 182, 183
cerambycod beetles (Rhagium Coccinella septempuncta 177f suborders 178t
sycophanta) 177f Coccinellidae (ladybird) 185–6, Tenebrionidae 187
Ceraphronoidea 259t 185f, 186f Coleopterida 172–88
Ceratophyllidae 205 Cochliomyia hominivo- Coleoptera see Coleoptera
veterinary importance 207t rax (New World (beetles)
Ceratophyllus galli- Screw-worm) 221 Coleorrhyncha 134, 137t
nae, veterinary cockchafer (Melolontha) 175, substrate-borne
importance 207t 175f vibrations 145
Ceratovacuna lanigera (sugar cockroaches 87–91 collecting 279–96
vane woolly aphid) 143 Blaberidae 88f absolute collecting
cerci 8, 20–1 Blattellidae 87, 88f techniques 289–93
Cercopidae, gut Blattidae 88f aquatic sampling
modifications 139 Corydiidae 88f techniques 293–6
Cereal Leaf Beetle (Oulema Cryptocercidae 88f relative collecting
melanopa) 185 morphology 87 techniques 280–9
Chagas’ disease 147 nitrogen cycle 87 taxa-specific sampling
Chalcidoidea 259t, 264–5 Nocticolidae 88f techniques 293–6
characteristic features of pests as 88 Collembola (springtails) 6, 36
arthropods 4–7 principal families 88f Colorado Potato Bee-
Chathamia sp. 233 Pseudophyllodromidae 88f tle (Leptinotarsa
Chathamiidae 233 sexual reproduction 89–90 decemlineata) 185
Chelicerata 5 sound production 89 combing, flea collection 294
chemical defences, plants 26 vibration sensitivity 88–9 commensal relationships,
chemoreceptors 29 see also Blattodea Ephemeroptera
Chicken Body Louse (cockroaches and (mayflies) 48
(Menacanthus termites) Common Greenhouse
stramineus) 129 Coconut Rhinoceros Beetle Thrip (Heliothrips
Chilopoda (centipedes) 7 (Oryctes rhinoceros) 183 haemorrhoidalis) 154
chitin 4, 14, 16 Coleoptera (beetles) 173–88, common jays (Graphium
4-chloro-m-cresol, specimen 173f, 174f, 175f, 176f doson) 238f
preservation 298 Adephaga 178t common scorpionflies
Chloroperlidae 71 Aleocharinae 182–3 (Panorpidae) 198, 199
chordotonal organs 167 aquatic species 180–1 compass termite (Amitermes
Choristidae 198 Archostemata 178t meridionalis) 91
Chortoicetes sp., pests as 81 Brachininae (Bombadier compound eyes 28
Chrysidoidea 259t, 266 beetles) 181–2 conservation issues,
Chrysomeloidea (leaf Carabidae (ground fieldwork 279
beetles) 179, 185 beetles) 181 copepods 6
Chrysoperla carnea 165f Cerambycidae 185 Copris sp. 184
Chrysops sp. 219–20 Chrysomelidae 185 corbiculum 269f, 270f
human disease vectors 220t Cicindellinae (tiger Cordylobia anthropophaga
cicadas, sap-feeding 26 beetles) 181f (Tumbu fly) 221
Cicadellidae (leafhoppers) Curculionidae 186–7 Corn Weevil (Sitophilus
gut modifications 139 Dytiscidae 180 zeamais) 176f
pests as 148 Gyrinidae 180–1 Corydiidae 88f
Cicadidae, gut Haliplidae 180 Cossidae (carpenter
modifications 139 Hygrobiidae 180 moths) 245–6
Cicindellinae (tiger morphology 178 Cotton Boll Weevil
beetles) 181f multiple lifecycles 179 (Anthonomus
Cimex lectularius, blood- Myxophaga 178t grandis) 187
feeding 146f, Noteridae 180 Cottony Cushion Scale (Icerya
147 nutrient recycling 176f purchasi) 143–4
306 SUBJECT INDEX
crabs 6 Death’s-Head Hawk Moth driver ants (Dorylinae) 269

crane fly (Tipulidae), (Acherontia atropos) 251 Drosophilidae, courtship
mouthparts 215f Deathwatch Beetle (Xestobium songs 216
Crectomys 62 rufovillosum) 186 Drugstore Beetle (Stegobium
Cretaceous, flowering plants 27 Deltochilum, baited traps 288 paniceum) 186
crickets see Orthoptera dengue fever, fly vectors 220t, drying, specimen
(grasshoppers and 222 preservation 298
crickets) Dermaptera (earwigs) 61–4, Dryinidae 266
Criocerus asparagi (Asparagus 61f, 62f dry mounting 299–300
Beetle) 185 Hemimeridae 62 dung-burying beetles 183–4
crista acoustica, Orthoptera 78 Neodermaptera 62 dusky thorn moth (Ennomos
crustacea (Pancrustacea) 5–6 terminal forceps 62, 63 fuscantaria) 236f
crypsis, Phasmatodea 116 Dermatobia hominis (Human D-Vac (Dietrick suction insect
Cryptocercidae 88f, 90 Bot Fly) 221 sampling) 291–2
Ctenocephalides canis (Dog Desert Locust see Schistocerca Dytiscidae (diving beetles) 180
Flea) 202f, 205 gregaria (Desert Locust) Dytiscus marginalis 181f
veterinary importance 207t Devonian period, herbivorous
Ctenocephalides felis (Cat insects 26
Flea) 202f, 203f, 205 Diamblomera sp. 256f earwigs see Dermaptera
veterinary importance 207t Diamphipnoidae 70 (earwigs)
cuckoo bee (Nomadinae) 270 Diapriidae 264 ecdysis 16
Culex sp. 220t, 221–2 Diaprioidea 259t Echidniophaga sp. 206
Culicidae (mosquito) Dietrick suction insect sampling Elateridae (click beetles) 174f
human disease 221–2 (D-Vac) 291–2 Elenchus japonicus, pest
mouthparts 215, 215f digestive system 29–30 control 193
Culicoides impunctatus (Scottish digger bee elephantiasis, fly vectors 220t
biting midge) 220 (Anthophorinae) 270 Embia sp. 119f
Culicoides sp., human disease Diplopoda (millipedes) 7 Embiidina see Embioptera
vectors 220t Diplura (diplurans) 6, 36 (webspinners)
Curculionidae (weevils) 186–7, Diprionidae 262 Embioptera (webspin-
187f Diptera (true flies) 196, 210–26, ners) 118–21, 118f,
Curculionoidea (weevils) 179 212f 119f
cuticle 13–17 Brachycera 213, 214t emergence traps 290–1, 294
structure of 14, 15f, 16 see also Cyclorrhapha Emesinae 141
cuticulin 14, 16 flight 212–13 Empididae (dance flies) 216
cyanide killing jars 296 human disease vectors 220t Empusa pennata 100f
Cyclorrhapha 213, 214t morphology 210f, 212–14 encephalitis, fly vectors 220t,
see also Calliphoridae (blow mouthparts 215f 222
flies); Muscidae (house Nematocera 213, 214t Encyrtidae 26
flies); Syrphidae (hover Orthorrhapha 213 endite–exite theory of flight 20f
flies) parasites as 217, 221 endites 19
Cyclotornidae, carnivorous principal taxa 214t endocuticle 15f
habits 242 sexual reproduction 215–16 Endopterygota 59, 156–276
Cynipoidea 259t, 264 warning colouration 215 Neuropterida see
Cyphoderris sp. 79 dispersal, flight 20–1 Neuropterida
Cyzenis albicans 218 diving beetles (Dytiscidae) 180 Ennomos fuscantaria (dusky
DNA sequencing 8 thorn moth) 236f
dobsonflies (Megaloptera) 156, Enoicyla pusilla 229f
damselflies see Odonata 161–4, 162f, 163f entognathous species 8
(dragonflies and Dog Flea see Ctenocephalides Entomophthora muscae 25
damselflies) canis (Dog Flea) Ephemera danica 45f
dance flies (Empididae) 216 Dog Louse (Trichodectes Ephemerellidae 47t
dancing, honey bees 272, 272f canis) 129 Ephemeridae 47t
darkling beetle Dolichopodidae (long-legged Ephemeroptera
(Tenebrionidae) 187 flies) 213 (mayflies) 45–50,
data recording, fieldwork 297–8 Dolichopus ungulatus 211f 45f, 46f
dead sorting, sweep net Dorylinae (driver/army body morphology 46–7
samples 281 ants) 269 families 47t
death feigning, Phasmatodea dragonflies see Odonata fossil records 49
(stick insects and leaf (dragonflies and Ephydridae (shore flies) 217
insects) 116 damselflies) epicuticle 14, 15f, 16
SUBJECT INDEX 307
epidermis, cuticle 15f fleas see Siphonaptera (fleas) grasshoppers see Acrididae
Epilamprinae 86f flight 19–22 (grasshoppers,
Epipyrops 242–3 dispersal 20–1 locusts); Orthoptera
Eriosoma lanigerum (Woolly endite–exite theory of (grasshoppers and
Apple Aphid) 143 flight 20f crickets)
ether vapour, killing mechanics of 22–3 grids, pitfall traps 287
methods 297 muscles 22 Gripopterygidae 70
2-ethoxy-ethanol, long- origin of 19–21 Gromphadorhina portentosa
term specimen paranotal theory of 20f (Madagascar hissing
preservation 300 secondary flightlessness 22–3 cockroach) 89
Euarthropoda 5 flight intercept traps ground beetles (Carabidae) 181,
Eukinolabia 112–21 (FIT) 284–6, 284f 181f
Embioptera (webspin- flowering plants, Cretaceous Gryllidae (true crickets) 75t
ners) 118–21, 118f, period 27 Grylloblatella 106
119f foregut 29–30 Grylloblatta sp. 105f, 106
see also Phasmatodea (stick Forficula sp. 61f, 62f Grylloblattina 106
insects and leaf insects) Formicinae 269 Grylloblattodea (ice
Eulophidae 265 Formicoidea 258, 259t, 268–9 crawlers) 105–7, 106f
Eumeninae (potter diet 268–9 Gryllotalpidae (mole
wasps) 267–8 eusociality 268 crickets) 75t
Eupaea fraseri 52f Formicinae 269 singing 77
Eupithecia 243 Hemiptera mutualistic Gyrinidae (whirligig
European Rabbit Flea (Spi- relationships 142–3 beetles) 180–1
lopysllus cuniculi), Myrmicinae 269
myxomatosis 207 fossil records 7
European Rat Flea (Nosopsyllus wing development 19–20
habitats, size effects 10
fasciatus) 205 Frankliniella occidentalis
Hedylidae 237
European social wasp (Polistes (Western Flower
haemolymph 12, 29
nimpha) 258f Thrip) 151f, 154
Haldane, J. B. S. 9, 177
Eustheniidae 70 fumigation, flea collection 294
Halictidae (sweat bees) 269–70
Evanioidea 259t fungal interactions 24–5
Haliplidae 180
evolution 3–4 furniture beetle (Anobium
Halobates 31–2
exites 19 punctatum) 186
hand collection, light traps 283
exocuticle 15f fused-back mayflies
hand nets 293–4
Exopterygota 59, 60–276 (Pannota) 47
hand searching 289–90
exoskeleton 14
hangingflies (Bittacidae) 198
external genitalia 31
hangingflies (Bittacus) 200
extracellular symbionts 24
hangingflies
extraction funnels 292
Galloisiana sp. 105, 106, 106f (Harpobittacus) 200
extraction techniques 292
ganglion 18 Haplocercata 60–8
extremophiles, Grylloblatta 107
gaseous exchange 10 Harlequin Beetle (Acrocinus
Gelechiidae 245 longimanus) 179
Geometridae 246–7 Harpobittacus
fairy shrimps (Branchiopoda) 5 German Cockroach (Blattella (hangingflies) 200
fat body 24 germanica) 90 hawk moths see Sphingidae
fieldwork 278–302 giraffe weevil (Trachelophorus (hawk moths)
data recording 297–8 giraffa) 175f Head Louse (Pediculus humanus
killing methods 296–7 glacial acetic acid, specimen capitis) 125f, 130, 130f,
long-term specimen preservation 298 131f
preservation 299–301 Glossata 240 head morphology 28
specimen preservation 298–9 Glossina sp. 224 Heath Trap 283
fig wasps (Agaonidae) 27, 265 human disease vectors 220t heel walkers (Mantophasma-
filariasis, fly vectors 220t, 222 Glossinidae 222–3 todea) 108–11, 108f,
firebrats (Zygentoma) 36, 41–3, grain thrip (Limothrips 109f
41f, 42f cerealium) 154 Heliconiinae 248–9
fish flies (Megaloptera) 156, Granary Weevil (Sitophilus Heliothrips haemorrhoidalis
161–4, 162f, 163f granarius) 187 (Common Greenhouse
fish lice 6 Graphium doson (common Thrip) 154
flavonoids, chemical jays) 238f Hemerobiidae (brown
defences 26 Graphopsocus cruciatus 123f lacewing) 169
308 SUBJECT INDEX
Hemimeridae 62 see also Apocrita; Vespina leaf beetles

Hemimetabola see Exopterygota flight 260 (Chrysomeloidea) 179,
Hemiptera (true bugs) 134–50, herbivory 262 185
134f, 135f, 135f, 136f larval stages 260 leaf-chewing insects 26
ant mutualistic lifestyle diversity 257–8 leaf-cutter bees
relationships 142–3 morphology 255f (Megachilidae) 270
disease-carrying 147 sex determination 262 leafhoppers see Cicadellidae
head morphology 139f superfamilies 259–60t (leafhoppers)
hunters as 141 hyperparasitoids 261 leaf insects see Phasmatodea
morphology 137–8 (stick insects and leaf
nutrition in 148 insects)
pests as 146–7, 148 Ibidoecus sp. 125f leishmaniasis, fly vectors 220t
sexual reproduction 145–6 ice crawlers (Grylloblat- Lepidophthirus macrorhini 131
sound production 144–5, todea) 105–7, Lepidoptera (butterflies and
145f 106f moths) 235–54, 236f
suborders 137t Icerya purchasi (Cottony acoustic signalling 241
see also Auchenorrhyncha; Cushion Scale) 143–4 baited traps 289
Coleorrhyncha; Ichneumonidae 263–4 carnivorous habits 242–3
Heteroptera; Ichneumonoidea 259t chemosensory sensilla 241
Sternorrhyncha idiobiont parasitoids 261 classifications 237
Heptagenidae 47t insect-borne illnesses vii collecting 295
herbivorous insects vii, 25–6 insecticide knockdown courtship 241
hermit crabs 6 techniques 293 crop pests 245
Hesperiidae 237 Integripalpia 230, 231t Hadylidae 237
Hessian Fly (Mayetiola integument 13 Hesperiidae 237
destructor) 219 interactions with larva behaviour 243–4
Heterobathmiidae 240 organisms 23–8 morphology 235f, 240
Heteroptera 134, 137f, 137t, intertidal zones 32 Noctuoidea 251–2
138f Ischnocera 128t reproduction 242
defence systems 142 animal hosts 128 subfamilies 238–9t
scent glands 141f mouthparts 128–9 Tineoidea 245
substrate-borne IUCN 279 visual courtship 242
vibrations 145 IUCN Red List of Threatened wings and flight 240–1
symbioses 140 Animals 27 Lepinotis sp. 127
Hexapoda 6, 36 Lepinotus patruelis 126
phylogeny 36f Lepisma saccharina 41f
hexapods (Pancrustacea) 5–6 jewel beetle Leptinotarsa decemlineata
hindgut 29–30 (Buprestidae) 184–5, (Colorado Potato
Hippoboscidae 223 185f Beetle) 185
Hodotermitidae 94t Leptoceroidea 231t
homeotic genes 3 Leptophlebidae 47t
honey bee see Apini (honey Kalotermitidae 94t Lesser Wax Moth (Achroia
bees) katydids (Tettigonidae) 75t grisella) 241
hornets (Vespinae) 267–8 Keroplatidae 217 Libelloides coccajus 166f
horse biting louse (Bovicola Kerria lacca (Lac Insect) 143 lift generation, flight 22
equi) 129 Kheper sp. 184 lights, insect repelled by 283
horseshoe crabs killing methods, light traps 282–4
(Merostomata) 5 fieldwork 296–7 Limnephilidae 231
house flies see Muscidae (house kings, termites 91–2 Limnephiloidea 231t
flies) koinobionts 261 Limothrips cerealium (grain
hover flies see Syrphidae (hover krill 6 thrip) 154
flies) Liothrips vaneeckei 154
Human Flea see Pulex irritans Liposcelis, sp. 127
(Human Flea) Lac Insect (Kerria lacca) 143 Liturgusa sp. 99f
Hydropsychoidea 231t ladybirds (Coccinelli- live animals, collecting
Hydroptilidae 231 dae) 185–6, 185f, from 296
Hydroptiloidea 231t 186f live sorting, sweep net
Hygrobiidae 180 Lasiocampidae 249 samples 281
Hymenoptera 256f, 258f, Lasiocampoidea 249 lobopods 3
265–75 Latinozoros sp. 66f lobsters 6
Apocrita 258 Laveran, Charles 222 Locustana sp., pests as 81
SUBJECT INDEX 309
Locusta sp., pests as 81 mark-capture-recapture Mormon Cricket (Anabris

locusts see Acrididae techniques, butterflies simplex) 79
(grasshoppers, locusts) collecting 295 mosquito see Culicidae
loiasis 219–20 Markia hystrix 76 (mosquito)
fly vectors 220t mason bees (Megachilidae) 270 moths see Lepidoptera
Lomamyia latipennis 168 Mastotermitidae 94t (butterflies and moths)
Lonchodes sp. 113f material colour, flight intercept moulting 16–17, 17f
longhorn beetle (Ceram- traps 285 termites 94
bycidae) 179, 185, matting, Galloisiana sp. 106 mouthparts 28
185f Maxillopoda (copepods, fish lice mudflats 32
long-legged flies and barnacles) 6 Mullerian mimicry 240
(Dolichipodidae) 213 Mayetiola destructor (Hessian Muscidae (house flies) 213
long-term specimen Fly) 219 mouthparts 215, 215f
preservation 299–301 mayflies see Ephemeroptera muscle structure 12
Lucanus cervus (stag (mayflies) mussel shrimps (Oligostraca) 6
beetle) 174f Mecoptera (scorpionflies) Mutillidae (velvet ants) 267
lures 288–9 197–201, 197f, 198f, 199, Mymaridae 265
Lycaenidae 249 199f mycetocytes 24
mecuronium 168 Mycetophilidae 217
Megachilidae (leaf-cutter and myrmecophilous species 183
mason bees) 270 Myrmeleontidae
Machilidae 39–40 Megaloprepus coerulatus 10 (ant-lions) 169–70
Macrosoma 237 Megaloptera (alderflies, Myrmicinae 269
Macrotermes (Termitidae) 91 dobsonflies, fish- myxomatosis 207
Macrotermitinae 93 flies) 156, 161–4, 162f, Myxophaga 178t
Madagascar hissing cockroach 163f
(Gromphadorhina Megalurothrips sjostedti 152f
portentosa) 89 Megalyroidea 259t, 263 Namkungia 106
Magicicada, synchronous mass Meganeura 10 Nannochoristidae 198
emergence 144–5 Megenillida 190 Nematocera 213, 214t
Magnetic Termite (Amitermes Megoura viciae 137f Nemoura sp. 70f
meridionalis) 91 Meinertellidae 39–40 Neodermaptera 62
Malacostraca (mantis shrimps, Melolontha (cockchafer) 175, Neopseustoidea 239t
crabs, shrimps, krill, 175f Neoptera 36, 59–276
lobsters and hermit Melolontha melolontha 173f neoteny 191
crabs) 6 Menacanthus stramineus Neotrogla sp. 126–7
Malaise traps 284–5, 285f (Chicken Body nervous system 17–18
killing methods 297 Louse) 129 neural lamella 18
malaria vii Mengenillidae 190 Neuroptera 165–77, 165f, 166f,
fly vectors 220t, 222 Menopon gallinae (shaft 167f
Malpighian tubules 30 louse) 129 courtship and mating 167
mangrove swamps 32 Meponidae 129 evolutionary history 170
Mantispa uhleri 168 Meropeidae 198 Hemerobiidae (brown
Mantispidae, parasitic Merostomata (horseshoe lacewing) 169
nature 168 crabs) 5 larval morphology 167–8
Mantis religiosa 101f methane production, morphology 166–7
mantis shrimps 6 termites 95–6 Myrmeleontidae 169–70
Mantoidea (praying man- microclimates, temperature pest control 168–9
tis) 98–103, 98f, 99f, regulation 13f Neuropterida 156–71
100f Microcoryphia see Ar- Megaloptera 156, 161–4,
cannibalistic chaeognatha 161f, 162f, 163f
behaviour 101–2 (bristletails) Neuroptera see Neuroptera
morphology 100–1, 101f Micropterigidae 240 Raphidioptera 156, 157–60,
predators as 100, 101 midgut epithelium 30 157f, 158f, 159f
reproduction 101–2 milkweed bug (Oncopeltus Nezara viridula (Southern
threat postures 102 fasciatus) 142 Green Stink Bug) 148
ultrasonic hearing 102 millipedes (Diplopoda) 7 Nicrophorus 183
Mantophasmatodea (heel mining bees (Andrenidae) 270 nitrogen cycle, cockroaches 87
walkers) 108–11, 108f, mole crickets see Gryllotalpidae nitrogen-fixing bacteria,
109f (mole crickets) termites 93
marine environments 31–2 monophyletic groups 3–4 Nocticolidae 88f
310 SUBJECT INDEX
Noctuidae, carnivorous Orthorrhapha 213, 214t Pea Aphid (Acyrthosiphon

habits 242 Tabanidae (horse fly) 213, pisum) 143
Noctuoidea 251–2 219–20 Pediculus humanus capitis
Nomadacris sp., pests as 81 Orussoidea 259t (Head Louse) 125f, 130,
Nomadinae (cuckoo bee) 270 Oryctes rhinoceros (Coconut 130f, 131f
Nomia strigata 256f Rhinoceros Beetle) 183 Pediculus humanus humanus
Nosopsyllus fasciatus (European Oulema melanopa (Cereal Leaf (Body Louse) 130, 130f
Rat Flea) 205 Beetle) 185 pelagic habitats 31–2
Noteridae 180 ovaries 31 Pemphigus spirothecae 143
Notiothaumatidae 198 ovoviviparity, Pentatomidae 141
Notoptera see Grylloblattodea cockroaches 89–90 perineurium (blood–brain
(ice crawlers) oxalic acid injection 297 barrier) 17–18, 19f
Nycteribiidae 223 oxygen supply, size Perlodes microcephala 69f
Nymphalidae 242, 248–9 constraints 11 peripheral nerves 18
Nymphalinae 249 Oxyphaloinae 89 Periplaneta americana (Amer-
ican Cockroach) 85f,
90
ocelli 28 research model viii
Odonata (dragonflies and Palaeoptera 36, 45–58 Perisphaerus sp. 86f, 90
damselflies) 51–5, 51f, see also Ephemeroptera peritrophic membrane 30
52f (mayflies); Odonata Perlidae 71
aquatic nymphs 56–7, 56f (dragonflies and Perlodidae 71
colouration 54 damselflies) Permian period, Ephemeroptera
flight 21f, 53–4 Pamphilioidea 259t (mayflies) 45
fossil records 57–8 Panarthropoda 4 Petrobius maritimus 38f, 39–40
morphology 52–3 Pancrustacea (crustaceans and Phaeostigma notata 157f
reproduction 55–6 hexapods) 5–6 Phasmatodea (stick insects and
Oecanthus sp., singing 77 Pannota (fused-back leaf insects) 113–17,
Oecophylla sp. 257f mayflies) 47 113f, 114f, 115f
Oestridae (warble flies/bot Panorpa communis 198f behaviour 115
flies) 221 Panorpa germanica 197f defensive strategies 116
Oligostraca (mussel shrimps Panorpidae (common morphology 115
and seed shrimps) 6 scorpionflies) 198, 199 pests as 116
ommatida 28 Panstrongylus sp., reproduction 115–16
onchocerciasis, fly vectors 220t Chagas’ disease sexual dimorphism 114f
Oncopeltus fasciatus (milkweed transmission 147 Phasmothrips, 154
bug) 142 paper wasps (Polistinae) 267–8 Philansius sp. 233
Oncothrips habrus 153 Papilio machaon 238f Philopotamoidea 231t
Oncothrips tepperi 153 Papiliomoidea 247–8 Phlaeopthripidae 153
Onomarchus sp. 74f Papilionidae 242 Phlebotomus sp., human disease
Onychophora (velvet worms) 4 Paraneoptera vectors 220t
origins of insects 7–8 (Acercaria) 122–55 Phoridae, feeding habits 219
Ormia ochracea 218 paranotal theory of flight 20f Phorina regina viii
Orthoptera (grasshoppers and parasites vii Phryganeoidea 231t
crickets) 73–83, 73f, 78f Parasitica 259t, 264–6 Phthirus gorillae 131
Caelifera see Caelifera Ceraphronoidea 259t Phthirus pubis (Pubic
families 75t Chalcidoidea 259t, 264–5 Louse) 130f, 131
Gryllidae (true crickets) 75t Chrysidoidea 259t, 266 phyla v
Gryllotalpidae (mole Cynipoidea 259t, 264 Phyllium sp. 114f
crickets) 75t Diaprioidea 259t physogastric queens,
hearing organs 78–9 Evanioidea 259t termites 94
jumping ability 76 Ichneumonoidea 259t Pieridae 248
morphology 75–6 Megalyroidea 259t, 263 pitfall traps 286–7, 286f
nymphal development 76 Platygastroidea 259t Plague Flea see Xenopsylla
reproduction 74f, 79–81 Proctotrupoidea 259t, 264 cheopis (Plague Flea)
Rhaphidophoridae (cave Vespoidea 257–8, 259t Plasmodium sp. 222
crickets) 75t parasitic diseases, secondary see also malaria
singing 76–8 flightlessness 23 PlastazoteTM 283
Tetrigidae (pygmy parasitoids 261 plastic drink cup, pitfall
locusts) 78 Pauropoda 7 traps 286
Tettigonidae (katydids) 75t Paussinae (ant-nest beetle) 182 Platygastroidea 259t
SUBJECT INDEX 311
Platypodidae (ambrosia beetles), carnivorous habits 242 Rhyniognatha hirsti, fossil

fungal interactions 24–5 larval behaviour 243 records 7
Plecoptera (stoneflies) 69–72, Pteromalidae 265 Rice Weevil (Sitophilus
69f, 70f Pteronarcyidae 71–2 oryzae) 187
Antarctoperlaria 70 Pteronarcys dorsata 71–2 Rickettsia prowazeki (typhus),
Arctoperlaria 70 Pterygota 44 Pediculus humanus
Chloroperlidae 71 Pubic Louse (Phthirus humanus (body
Perlidae 71 pubis) 130f, 131 louse) 130
Perlodidae 71 Pulex irritans (Human robber fly (Asilidae) 213
Plutellidae 245 Flea) 202f, 205 Robinson Trap 283
Polistes nimpha (European veterinary importance 207t Romaleidae 76
social wasp) 258f Pulicidae, veterinary Ross, Ronald vii
Polistinae (paper wasps) 267–8 importance 207t round dancing, honey
pollination v, vif, 27–8 pupariate 217 bees 272f
Apini (honey bees) 272–3 Pycnogonida (sea spiders) 5 royal jelly 271–2
pollution, Ephemeroptera pygmy locusts (Tetrigidae) 78t
(mayflies) 49 Pyralidae 243
polymorphism, termites 91 acoustic signals 241 salt marshes 32
Polyneoptera 60, 69–276 carnivorous habits 242 sampling methods 279
Polyphaga 178t, 182–3 Pyraloidea 246–7 sap-feeding insects 26
polyphyletic groups 3–4 Pyrgomorphidae (bush hoppers, Sarcophaga carnaria 210f
Pompiloidea 259t, 267 bush locusts) 76, 78t Saturniidae 250
pooters 290, 290f Satyrinae 248–9
light trap collection 283 Scarabaeidae (scarab
potter wasps beetles) 183–5
quantitative analysis, light
(Eumeninae) 267–8 Scarabeus sp. 184
traps 283
power, size vs. 9–10 scarabiform grubs 180
queens, termites 91–2
praying mantis see Mantoidea Schistocerca gregaria (Desert
(praying mantis) Locust) 73f
predators vii pest as 81
prey, insects as v, vii radiodonts 3 research model viii
Prionoglarididae 126 Ramsdelepedion schusteri 10 Schistocerca sp., pests as 81
Proctotrupoidea 259t, 264 Raphidia 159 Schistonota (split-back
procuticle 14, 15f Raphidioptera (snakeflies) 156, mayflies) 47
Protohermes motuoensis 162f 157–60, 157f, 158f, 159f Schizodactylus sp. 75f
protozoa, symbiosis 24 rearing 290–1 sclerites 14
Protura (proturans) 6, 36 Redi, Franscesco 217 Sclerophasma sp. 109f
Pselaphinae 182–3 Reduviidae (assassin bugs) 136f Scolioidea 259t
pseudergate caste, termites 94 relative collecting Scolytidae (bark-boring
Pseudophyllodromidae 88f techniques 280–9 beetles) 187
Psocodea (barklice, booklice relative sampling methods, fungal interactions 24–5
and true lice) 123–33, sweep nets 280–1 scorpionflies (Mecoptera) 197–
123f, 124f, 125f Remipeda (remipedians) 6 201, 197f, 198f, 199,
Anoplura 129–30, 130f reproduction 23, 31 199f
ectoparasites as 127–8 research models viii Scottish biting midge (Culicoides
habitats 124–5 resilin 204 impunctatus) 220
Menoponidae 129 respiration, size and 10 searching, flea collection 294
morphology 125–6, 128 respiratory system 30 sea spiders (Pycnogonida) 5
pests as 127 Rhagium sycophanta secondary flightlessness 22–3
Phthiraptera suborders 128t (Cerambycid seed shrimps (Oligostraca) 6
Psocomorpha 125 beetles) 177f sense organs 28–9
reproduction 128 Rhaphidophoridae (cave Sericostomatoidea 231t
Rhynchophthirina 132 crickets) 75t Serritermitidae 94t
secondary flightlessness 23 Rhinotermitidae 94t Sesiidae 246
sexual reproduction 126–7 Rhodinus sp., Chagas’ disease shaft louse (Menopon
Trichodectidae 129 transmission 147 gallinae) 129
Troctomorpha 125 Rhodnius prolixus viii shore flies (Ephydridae) 217
Trogiomorpha 124 Rhyacophiloidea 231t shrimps 6
wing sexual dimorphism 126 Rhynchophthirina 128t, 132 Sialis lutaria 161f
Psocomorpha 125 Rhyniella praecursor, fossil silk glands, Embioptera
Psychidae records 7 (webspinners) 119–20
312 SUBJECT INDEX
silk moth (Bombyx Spilopysllus cuniculi (Euro- surface area to volume ratio 9
mori) 249–50 pean Rabbit Flea), swarms
silverfish (Zygentoma) 36, myxomatosis 207 Ephemeroptera
41–3, 41f, 42f Spinopeplus sp. 114f (mayflies) 48–9
Simulium sp., human disease spiracles 10, 30 honey bees 272
vectors 220t split-back mayflies sweat bee (Halictidae) 269–70
Siphonaptera (fleas) 196, 202–9, (Schistonota) 47 sweep nets 280–1
202f, 203f spool-and-line techniques, flea symbiotic bacteria 24
Ceratophyllidae 205 collection 294 symbiotic protozoa 24
collecting 294–5 stag beetle (Lucanus Symphyla 7
disease carrying 207 cervus) 174f morphology 258–9
disease vectors as 206 standardization, sweep Symphyta 258, 259f, 259t
host specificity 207 nets 280–1 Cephoidea 259t
insecticidal control 207 Staphylinidae 182, 183 Orussoidea 258
Ischnopysllidae 205 Stegobium paniceum (Drugstore see also Vespina
life cycle 205 Beetle) 186 Pamphilioidea 259t
lifestyle 204–5 Stenolemus sp. 141 Siricoidea 259t
morphology 204 Stenophylax permistus 229f Tenthredinoidea 259t, 262–3
parasitic lifestyle 203–4 Stenophylax vibex 228f Xiphydrioidea 259t
secondary flightlessness 23 Stephanoidea 259t, 263 Xyeloidea 259t
veterinary importance 207t Sternorrhyncha 134, 137t, 138f synonymy 32
Siricidae 262 mouthparts 138 Syrphidae (hover flies) 213
Siricoidea 259t secretions 143 warning colouration 215
sister groups 8 Stichotrema dallatorreanum 193
Sitophilus granarius (Granary pest control 193–4
Weevil) 187 stick insects see Phasmatodea
Sitophilus oryzae (Rice (stick insects and leaf
Tabanidae (horse fly) 213,
Weevil) 187 insects)
219–20
Sitophilus zeamais (Corn sticky traps 285–6
Tabanus sp., human disease
Weevil) 176f Stolotermitidae 94t
vectors 220t
size 8–13 stoneflies see Plecoptera
Tachinidae 217–18
constraints of 11–13 (stoneflies)
Tapestry Moth (Trichophaga
power vs. 9–10 Streblidae 223
tapetzella) 245
sleeping sickness 222–4 street lamps, moth population
Tardigrada (water bears) 4
fly vectors 220t reduction 282–3
taxa-specific sampling
snakeflies (Raphidioptera) 156, Strepsiptera 36, 189–95, 189f,
techniques 293–6
157–60, 157f, 158f, 159f 190f, 191f, 192f
temperature regulation, size
snow scorpionflies (Bor- endoparasites as 190, 191
an 12–13
eidae) 198, 199, Mengenillida 190
Tenebrionidae (darkling
200 Mengenillidae 190
beetle) 187
social wasp see Vespoidea morphology 191
Tenthredinidae 262
soldiers, termites 92, 92f pest control in 193–4
Tenthredinoidea 259t,
sorting, sweep net samples 281 sexual reproduction 192–3
262–3
sound production, Stylopidia 190
pest species 262–3
cockroaches 89 structure of insects 28–31
Terebrantia 153
Southern Green Stink Bug stuffing, specimen
terminal forceps, Dermaptera
(Nezara viridula) 148 preservation 298
(earwigs) 62, 63
species numbers, body weight Stylogaster sp. 212f
termites 91–7
vs. 9f Stylopidia 190
Apicotermitinae 93
species proportions vif Stylops ater 190f, 191f
Archotermopsidae 94t
specimen preservation 298–9 Stylops melittae 189f
castes 91, 94
Speiredonia spectans 236f Stylotermitidae 94t
cellulose digestion 93
Sphingidae (hawk suction sampling 291–2
collecting 295–6
moths) 250–1 sugaring, baited traps 289
ecosystems in 94–5
acoustic signals 241 sugar vane woolly aphid
families 94t
Sphodromantis centralis 98f (Ceratovacuna
food as 94–5
Spicipalpia 230, 231t lanigera) 143
Hodotermitidae 94t
spiders (Arachnids) 5 Surber sampler 294
Kalotermitidae 94t
SUBJECT INDEX 313
Macrotermitinae 93 Trachelophorus giraffa (giraffe Tunga penetrans, veterinary

Mastotermitidae 94t weevils) 175f importance 207t
morphology 91–3 transfer, sweep nets 281 Tungidae, veterinary
moulting 94 trap collecting, bees importance 207t
nitrogen-fixing bacteria 93 collecting 295 typhus (Rickettsia prowazeki),
pests as 95–6 Triarthria setipennis 218 Pediculus humanus
polymorphic types 91 Triatoma infestans, humanus (body
range of 91 blood-feeding 146f louse) 130
Rhinotermitidae 94t Triatoma sp., Chagas’ disease Tyrannophasma gladiator 108f
Serritermitidae 94t transmission 147
soldiers 92, 92f Trichodectes canis (Dog
Stolotermitidae 94t Louse) 129
ultrasonic hearing, Mantoidea
Stylotermitidae 94t Trichodectidae, mammal (praying mantis) 102
Termitidae 94t biting 129
ultraviolet radiation, light
Termitinae, 93 Trichogrammatidae 265 traps 282
workers 92–3, 93f Trichophaga tapetzella (Tapestry
see also Blattodea Moth) 245
(cockroaches and Trichoptera (caddis-
termites) flies) 228–34, Vanessa cardui 235f
Termitidae 94t 229f velvet ants (Mutillidae) 267
Termitinae, 93 Annulipalpia 230, 231t velvet worms (Onychophora) 4
Termitomyces sp. 93 aquatic larvae 230–3, 230f, ventral nerve cords 18
terpenoids, chemical 232f Vespina 258, 259t
defences 26 habits 230 Orussoidea 259t
terrestrial ecosystems v Integripalpia 230, 231t Vespinae (yellow jackets and
testes 31 morphology 228f, 229–30 hornets) 267–8
Tetrigidae (pygmy locusts) 78t pollution response 232–3 Vespoidea 257–8, 259t
Tettigonidae (katydids) 75t pupation 232 vibration sensitivity,
Thephritidae, courtship sexual reproduction 230–3 cockroaches 88–9
songs 216 Spicipalpia 230, 231t viscosity, size and 21
Thermobia domestica 42f, 43 suborders 231t visual acuity 29
thorax morphology 28 Trigonaloidea 259t, 263–4 volatile chemicals, plants 26–7
Thorax porcellana 90 Truljalia hibionis 79
Thripidae 153 Trilobita (trilobites) 5
thrips see Thysanoptera (thrips) trisodium orthophosphate, waggle dancing, honey
Thynnoidea 259t long-term specimen bees 272f
Thysanoptera (thrips) 151–5, preservation 300–1 warble flies (Oestridae) 221
151f, 152f Troctomorpha 125 wasps see Hymenoptera
mouthparts 152 Trogiomorpha 125 water-bears (Tardigrada) 4
pests as 154 Trogium sp. 127 water-fleas (Branchiopoda) 5
pupa resting stages 152–3 trophallaxis 93 water pan traps 287–8, 288f
reproduction 153, 154 trophic level v water, pitfall traps 286
sub-social behaviour 153 tropical areas, light traps 282–3 Webbing Clothes Moth (Tineola
Terebrantia 153 tropical forests ix bisselliella) 245
Tubulifera 153 Tropidacris crisata 78f webspinners (Em-
Thysanura see Zygentoma true bugs see Hemiptera (true bioptera) 118–21, 118f,
(silverfish and bugs) 119f
firebrats) true crickets (Gryllidae) 75t weevils (Curculionidae) 179,
tiger beetles true flies see Diptera (true flies) 186–7, 187f
(Cicindellinae) 181f true lice see Psocodea (barklice, Western Flower Thrip (Franklin-
Tinea pellionella (Case-bearing booklice and true lice) iella occidentalis) 151f,
Clothes Moth) 245 Trypanosoma brucei 223–4 154
Tineidae 245 Trypanosoma cruzi 147 Western Honey Bee (Apis
carnivorous habits 242 tsetse flies 222–3 mellifera) 271
Tineola bisselliella (Webbing control of 224 whirligig beetles
Clothes Moth) 245 Tubulifera 153 (Gyrinidae) 180–1
Tiphioidea 259t tularaemia, fly vectors 220t window traps 285–6
Tipulidae (crane fly), Tullgren funnels 292 winged insects 44–276
mouthparts 215f flea collection 295 Winkler bags 292
Tortricoidea 246 Tumbu fly (Cordylobia flea collection 295
Torymidae 265 anthropophaga) 221 witchety grubs 246
314 SUBJECT INDEX
Wolbachia, insect Xenopsylla cheopis (Plague yellow pan traps 287–8

interactions 24 Flea) 206 Yersinia pestis infection 206
Woolly Apple Aphid (Eriosoma veterinary importance 207t Yponomeutoidea
lanigerum) 143 Xestobium rufovillosum economically important
workers, termites 92–3 (Deathwatch species 245
World Conservation Monitoring Beetle) 186
Centre 279 Xiphydrioidea 259t
Xyeloidea 259t Zika virus, fly vectors 220t
Xylocopinae (carpenter Zoraptera (angel insects) 65–9,
Xanthostigma 159 bees) 270 65f, 66f
Xenonomia 104–11 Zorotypus hubbardi 65f
see also Grylloblattodea Zygaenoidea 242
(ice crawlers); Man- yellow fever, fly vectors 220t, Zygentoma (silverfish and
tophasmatodea (heel 222 firebrats) 36, 41–3, 41f,
walkers) yellow jackets (Vespinae) 267–8 42f

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ESSENTIAL ENTOMOLOGY

The image shown here is a

impossible for most plants,

Section 1 Introduction to insect evolution

Section 2 The insect orders 35

The winged insects 44

Orthoptera (grasshoppers and crickets) 73

Division Endopterygota (= Holometabola) 156

Section 3 Fieldwork 277

Subject Index 303

George McGavin studied Zoology at Edinburgh University, followed by a

Leonidas-Romanos Davranoglou studied Zoology at Imperial College

how insects communicate using sounds by examining the morphology and

Aedeagus The organ of copulation in male insects.

Gall An abnormal plant growth caused by a virus, bacterium, fungus, mite,

Monophagous Restricted to eating a single species plant or type of

Synanthropic Associated with humans and their habitations.

The evolution of the arthropods

Front legs of the mole and

originate from a single worm-like ancestor, which gained a chitinous, seg-

Features that characterize arthropods

The arthropod-like groups (Panarthropoda)

Onycophora (velvet-worms) > 200 species

Tardigrades (water-bears) > 1,300 species

NB: Illustrations not to scale.

The true arthropods (Euarthropoda)

Mygalomorph spider Arachnida (spiders, scorpions, harvestmen, ticks, mites, psuedoscorpions,

Pancrustacea (crustaceans and hexapods)

Oligostraca (mussel shrimps and seed shrimps) > 13,000 species

Maxillopoda (copepods, ﬁsh lice, and barnacles) > 14,000 species

Remipedia (remipedians) 28 species

Collembola (springtails) > 9,000 species

Protura (proturans) > 800 species

Diplura (diplurans) 800 species

Insecta (insects) > 1,000,000 species

Pauropoda (pauropods) > 800 species

Diplopoda (millipedes) > 12,000 species

Chilopoda (centipedes) 8,000 species

Symphyla (symphylans) 200 species

The origins of insects and other

Nevertheless, Cloudsley–Thompson concluded that they evolved from

Five factors in a winning formula

Five factors contributing to the success of insects

cell at around 0.3 µm (micrometres) long to a blue whale at 30 m long. In the

Log species richness

A chrysomelid beetle hanging on to a leaf in a gale may resist a force ten or

Temperature regulation The large surface area-to-volume ratio of insects

Microclimate matters. The 27.5°C

fungal pathogens. In many cases, it is also able to mount an immune response

Cuticle structure We need to look closely at the structure of cuticle to un-

Wax filament Inner epicuticle

(a) Mature cuticle (b) Apolysis

(c) New epicuticle produced (d) Endocuticle digested Endocuticle–

(e) Molting fluid resorbed (f) New cuticle after ecdysis

The stages of moulting.

The nervous system and the blood–brain barrier

in osmotic pressure and ionic concentrations, and normal homeostatic pro-

Fibre Nerve cell body