Book Reviews

AN ILLUSTRATED GUIDE TO THE PARASITIC WASPS

ASSOCIATED WITH CITRUS SCALE INSECTS AND

MEALYBUGS IN AUSTRALIA. By M. B. Malipatil, K. L.

Dunn and K. Smith. Agriculture Victoria, Knox®eld, Victoria,

Australia. 2000. Pp. iv + 152. A$40.00 (softcover). ISBN

07311 4458 9.

This guide is intended to serve as a user-friendly tool for

the integrated pest management (IPM) consultant, ®eld

worker and specialist entomologist to identify ®fty-three

common hymenopteran parasitoids (Hymenoptera:

Chalcidoidea: Platygastroidea) of citrus scale insects and

mealybugs in Australia. After a brief introduction to the

history of biological control of citrus scale insects and

mealybugs in Australia, the authors give some essential

details on collection, preservation and preparation of

chalcidoid wasps, which are pivotal steps toward correct

identi®cation. Technical terminology is reduced to a

minimum, as are the diagnoses of the ®ve families of

parasitoids included in the guide, viz. Aphelinidae,

Encyrtidae, Pteromalidae, Eulophidae and Platygastridae.

The three illustrated keys to genera and species of

parasitoids associated with citrus scale insects and mealy-

bugs form the bulk of the guide. Each deals with the

parasitoids of a single family of pests: Key A for Coccidae

(soft scale insects), Key B for Diaspididae (armoured scale

insects) and Key C for Pseudococcidae (mealybugs). Each

parasitoid name in the keys is referred to a brief account of

the species, with diagnosis, biology, distribution within

Australia and key references. Details of several uncommon

parasitoids (of Encyrtidae and Pteromalidae) not included in

the keys, but associated with soft scales and mealybugs, are

also given. A single comprehensive list of references and a

useful appendix on host-parasitoid association precede the

colour plates.

This guide certainly ful®ls its aim, being concise and very

easy to use. The colour plates, considering the minute size of

these parasitoids, are excellent. It also makes it clear that the

correct identi®cation of a natural enemy is essential in

biological and integrated control programmes.

From a user point of view, a couple of minor dif®culties

arise from the way the guide is organized. A single,

comprehensive key of parasitoids would have been easier to

use, by avoiding the stage of family pest identi®cation.

Moreover, illustrations that are used more than once through

the keys are always referred to with the same number, which

sometimes creates some confusion.

Overall, information and references given in this guide make

it a recommended purchase to anyone interested (not only in

Australia) in the biological control of citrus scale insects and

mealybugs.

EMILIO GUERRIERI

THE PRAYING MANTIDS. Edited by F. R. Prete, H. Wells,

P. H. Wells and L. E. Hurd. The John Hopkins University

Press, Baltimore, Maryland. 2000. Pp. xiv + 362. £58.00

(hardcover). ISBN 0-8018-6174-8.

Those who have spent years breeding and rearing mantids have

many questions to which answers have not been readily

obtainable until now, and for the novice this book will be a

treasure-house of information. The editors have happily

combined the expert contributions to present a logical and

lucid format, although the wealth of information provided

serves also to highlight the many unresolved questions and

puzzles still remaining.

Our changing attitude to mantids through the centuries is

discussed with much historical detail, from the symbolic role

of the mantis in ancient Egypt up to present-day misinterpret-

ations of mantis behaviour. The female praying mantis has had

a bad press for several centuries, as the cannibal who devours

her mate, before, during or after copulation. This misconcep-

tion has been complicated by the many reports of the half-

eaten male successfully copulating with the female who is in

the process of eating him. However, in recent years this has

been discovered to be an artefact of captivity in the majority of

cases, resulting from unnatural conditions for both male and

female, but the myth lingers on. The reluctance to accept that

males are occasionally eaten through hunger or by chance,

rather than by design, is apparently linked to the equally

erroneous attitude that mantids usually eat only small insects,

¯ies, etc., and that capturing large prey is unnatural. The `rare'

occurrence of mantids capturing and consuming larger insects

and even small vertebrates, lizards, etc., is shown to be part of

the normal behaviour pattern, and an area for further research.

The appropriate psychological attitude towards mantids

having been established, their position in the scienti®c world is

detailed with an excellent introduction to their morphology and

to the current taxonomy of the group, but also pointing out that

`a great deal of work needs to be done before we will arrive at

an appropriately detailed and entirely satisfactory taxonomy'.

Aspects of mantis biology are discussed under three main

headings, commencing with ecology and mating behaviour,

although the ecological studies are based on only three species,

Mantis religiosa, Tenodera aridifolia sinensis and Tenodera

angustifolia. Experiments have shown that hungry mantid

nymphs move around more and that larger mantids eat larger

prey, and also demonstrate the ever-present problem of

experimental subjects being cannibalized. These and other

features of mantis biology will be recognized as true for the

majority of species reared in captivity. The chapter on mating

behaviour reviews the cannibalism of males by females within

Mantodea and the associated costs and bene®ts, while

indicating further avenues of research. The majority of results

are based on studies of copulation during the day, because of

# 2001 Blackwell Science Ltd 125

Systematic Entomology (2001) 26, 125±129Systematic Entomology (2001) 26, 125±129

the diurnal activity patterns of researchers, irrespective of

potential mantis behaviour. Further work is also called for on

the subject of female pheromone attraction of males. Although

documented only in Acanthops falcata, several researchers

suspect that sedentary females emit pheromones to attract

actively ¯ying males.

That praying mantids possess tympanal hearing organs may

still come as a surprise to many readers, although it is nearly

15 years since their discovery and published description. This

hearing organ, their DK (`Deep groove with Knobs') ear or

Cyclopean ear, is located in the ventral midline of the thorax

near its junction with the abdomen, and is possessed by ¯ying

mantids in all but the four most primitive families, although

not by brachypterous, non-¯ying females. Flying mantids use

this ultrasonic hearing to protect themselves from echolocat-

ing, insectivorous bats, and although predator evasion is the

prime function, further research on mantis acoustic ability is

continuing.

The studies of binocular vision and distance estimation, and

the complexities of prey recognition in mantids, involve both

mantis behaviour patterns and those of research workers with

`small ¯y prey' preconceptions. Discussion of prey capture

begins with facetious, but fascinating, descriptions of how not

to analyse behaviour, and concludes with the illuminating

suggestion that before even reading any scienti®c literature on

mantis predatory behaviour, ®rst acquire and carefully watch a

living mantis in natural conditions!

Mantids have also developed characteristic and complex

defence mechanisms, involving crypsis, `I am not here';

mimicry, `I am not palatable'; special resemblance, `I am not

edible'; and behavioural strategies. Different behaviour

patterns may be seen in the same species at different stages

of development, from assuming cryptic postures or making

escape runs, to displaying the threatening deimatic response

seen in many adults. Discussion of these mechanisms is

illustrated with a tantalizing selection of colour photographs.

Finally, rearing and breeding techniques (we all have

learned a little from our own mistakes, now all may bene®t

from the combined expertise available) and histological

procedures, complete a book which is written with a superb

combination of enthusiasm and expertise by the many

contributors. Although some of the experimental procedures

described may not appeal to the more general enthusiast, this

book is a must-have for all those involved with mantids.

JUDITH MARSHALL

BUTTERFLIES OF AUSTRALIA. THEIR IDENTIFICA-

TION, BIOLOGY AND DISTRIBUTION. By Michael F.

Braby. CSIRO Publishing, Collingwood, Victoria, Australia.

2000. Two volumes, 1008 pp., colour illustrations (hardcover).

$US195.00. ISBN 0643065911.

Michael Braby is to be congratulated for producing these two

lavish new volumes on the butter¯ies of Australia that are

smartly united by a single lycaenid on the spine. Unlike other

major faunal projects updated by modern biological informa-

tion, the entire fauna is covered in one publishing swoop, and

is not available separately. The project was achieved in under

seven years. The introduction, faunal checklist, and

Hesperiidae, Papilionidae and Pieridae (with all the colour

plates) are included in Volume 1; Nymphalidae and

Lycaenidae (including riodinids) and a series of useful

appendices are included in Volume 2.

This two-book set, a major update of earlier works by

Common & Waterhouse (1981), has been critically reviewed

by a range of Australian butter¯y experts. Although one of its

stated aims is to provide a reliable identi®cation manual, this is

de®nitely one for the desk rather than the ®eld, although it will

possibly be carried by a few people in their cars.

Comprehensiveness is the second stated aim, and I will return

to this shortly, in relation particularly to biology, systematics

and distribution.

The introduction is fairly basic, regionally dedicated,

providing a very good introduction for the non-specialist

(who may, however, not be able to afford the volumes,

although the price is fairly reasonable for the two). It does not

add much new information. There is little information on

history and biogeography, classi®cation and the species

concept, but no section on systematics (instead, almost all

the useful systematic information is included under Family and

Subfamily sections). There is some general information on

morphology, a section on geographical distribution and

variation, including species concepts, and ®nally a good

coverage of conservation and protection.

The only (unresolved) overview of butter¯y relationships

appears on pages viii±ix and accords with the checklist on

pages 53±58. In fact, the classi®cation is identical to the highly

conservative butter¯y part of the checklist of Nielsen et al.

(1996), which itself follows Nielsen & Common (1991). Braby

does not attempt to justify such a conservative approach,

although more recent works, such as Ackery et al. (1995)

which review the evidence for more structured relationships,

are brie¯y discussed under higher taxa within the species

section. Anyway, there is nothing wrong with being con-

servative in systematics, especially considering that butter¯y

higher relationships are currently in a state of turmoil.

Nonetheless, I would argue here that it is disappointing that

more systematic structure based on more widely accepted

synapomorphies is not yet appearing in many modern

textbooks of this calibre. Of course, it will be a while before

molecular systematics revolutionizes our overview of butter-

¯ies and their allies in the way that arguably has already

happened for higher plants (as a matter of fact, Appendix V in

Volume 2 still uses a traditional plant classi®cation, although

the emerging modern structure would have been far more

useful for interpreting butter¯y foodplant relationships). A new

classi®cation is only better than an old one if it is more

explicit, especially in terms of its character, and there are now

some much better systematic frameworks. Good examples are

those proposed by Harvey for Fred Nijhout's (1991) now

classic book on butter¯y wing patterns, and the review by

Ackery et al. in Kristensen's (1999) monolithic new work (the

latter, however, did not appear in time for this work). For

example, some molecular data suggest that riodinids are better

# 2001 Blackwell Science Ltd, Systematic Entomology, 26, 125±129

126 Book Reviews126 Book Reviews

regarded as a separate family, probably the sister of lycaenids

(but there is the lingering problem of position of the Andean

Styx). The Old World riodinids tend to be Myrsinaceae-

feeders, and the one Australian representative of the family

Praetaxila is no exception to this pattern. Some other trends

are that morphological and molecular data tend to support a

clade including Acraeinae and Argynninae, as well as genus

Cethosia, that merit a more inclusive concept of Heliconiinae;

Amathusiinae are better placed in an expanded Morphinae; on

the other hand, Satyrinae and other monocot-feeders, as well as

the possibly paraphyletic Biblidinae (= Limenitinae), are in a

state of ¯ux at present and so these groups are indeed better left

unresolved at present.

In terms of biogeography, Australia is fascinating, although

surprisingly little has been made of this in the introduction on

pages 16±20. Much of Australia's butter¯y fauna has a very

derived feel to it, that would accord with (as far as we know)

distinctly post-Cretaceous/Tertiary radiation of butter¯ies

world-wide (Australia was only directly accessible to the Old

World about 30 million years ago, but to the New World via

Antarctica considerably closer to this famous geological

boundary). For example, the few mycalesine satyrines present

are probably quite derived with this endemic Old World

radiation, and are probably therefore relatively recent colonists

(however, the probable sister of other mycalesines,

Orsotriaena, whose Australian subspecies is protected, did

make it to the extreme north-eastern tip of the continent).

Australia is particularly interesting for the endemic presence of

the regent skipper Euschemon raf¯esia (a protected species in

one if not both senses of the word, despite the fact that it is

fairly widespread in tropical Australia, that even feeds on a

Gondwanan plant group, Monimiaceae). Whether this hesper-

iid is basal or highly derived and its af®nities with other

Pyrginae, particularly the widespread genus Celaenorrhinus,

may only be resolved by molecular analyses, interpreted by

better morphological information. For example, a recent paper

by Fanger (1999) points out that the morphology of the tergal

phragmata is very simple and plesiomorphic compared with

other skippers examined, adding more spice to the potential

interest of this (uniquely for a male butter¯y) frenulum/

retinaculum-equipped taxon. The presence in the Australasian

Region of Tellervo is also of great biogeographic, and maybe

even plate tectonic interest, but its true relationship with

Danainae and the exclusively Neotropical Ithomiinae has still

not been resolved once and for all. Doubtless, new systematic

work at a global level will provide fascinating insights into the

relationships of some other of the more peculiar Australian

skippers and browns, although again one gets only an inkling

of the possible excitement of future systematic research from

the book.

The layout is very concise throughout. Although there is no

typological reference (Nielsen et al., 1996, needs to be

consulted for that), there is a shadowed header section giving

a short diagnosis of each family, subfamily and genus, but with

no tribal level classi®cation. This is wise in consideration of

the vast number of nomenclatural changes we can anticipate

for butter¯ies in the next few years. There is a good quality

black and white photograph for each species (a nice touch,

especially with the arrows indicating distinguishing marks),

and a distribution range-®ll map that summarizes recognized

subspecies and also gives the distribution of adult seasonality

over the months. For simplicity and clarity, these maps are

®ne. However, in terms of their scienti®c content they are more

appropriate for a portable ®eld guide. Considering the range of

mapping programs now available, and the range of interpola-

tion and extrapolation techniques invented in Australia, it is

now straightforward, given a good database that should

have been available to this book project, to produce either

dot or grid distribution maps (compare the butter¯y distribu-

tion dot maps in Dunn & Dunn, 1991, the Flora of Australia

series, or those in Parsons, 1998), if not indeed some-

thing more sophisticated. If a research worker wanted to

estimate diversity patterns, for example, using such maps as

a basis cannot easily be justi®ed. The species treatment

has sections on vernacular names, short description, geo-

graphical variation, similar species, immature stages, larval

food-plants, life-cycle and behaviour, distribution and habitat,

and major references. Within the sections on food-plants

and ife-cycle resides the large bulk of completely new

information.

The book is remarkably free of typographic errors. The

plates are of very high quality, with excellent ®delity in

colour reproduction and the right subtlety of shadow, and it

is aesthetically good to view both halves of the adult

butter¯ies (in contrast for example to recent works by

Parsons), although the background does vary somewhat in

perceived texture. Critically for any identi®cation guide, the

plates are indexed back to the text pages, and one can use

the slight disjunction in numbers after 341 to recall where

Volume 2 starts. Commendably, the labels of each

organism are very well documented. Plates 65 and 66

show a handful of eggs and a few resting postures, and

there are only four plates of representative larval and pupal

photographs. With the number of biologies known and

photographed in Australia, it is surely a missed opportunity

that the early stage coverage is so reduced in this book.

Volume 2 has no colour, but contains a useful series of

appendices that include (otherwise untreated) peripheral and

non-Australasian Region island species, legally protected

species and larval food-plants. There is a glossary, a

comprehensive bibliography and a taxonomic (only) index.

In summary, the book is not to be viewed particularly as a

source of new systematic and biogeographic information. Its

strength lies in the wealth of new and updated biological

information in the species section. For this reason, and for the

smart presentation, these two volumes are recommended as a

purchase for libraries and all those interested (just as in the

title) in the identi®cation, biology and distribution of

butter¯ies, if not in the Australian fauna in general. This will

be an important shelf reference for many years to come for

anyone working on the Australasian fauna or interested in

comprehensive modern reviews of butter¯ies in general. The

price is a fair one for over 1000 beautifully printed and

virtually typo-free pages.

DAVID LEES

# 2001 Blackwell Science Ltd, Systematic Entomology, 26, 125±129

Book Reviews 127Book Reviews 127

References

Ackery, P.R., Smith, C.R. & Vane-Wright, R.E., eds (1995)

Carcasson's African Butter¯ies: an Annotated Catalogue of the

Papilionoidea and Hesperioidea of the Afrotropical Region.

CSIRO, Melbourne.

Common, I.B.F. & Waterhouse, D.F. (1981) Butter¯ies of Australia.

Angus and Robertson, London.

Dunn, K.L. & Dunn, L.E. (1991) Review of Australian Butter¯ies:

Distribution, Life History and Taxonomy. 4 Volumes. Privately

published by the authors.

Fanger, H. (1999) Comparative morphology of tergal phragmata

occurring in the dorsal thoraco-abdominal junction of ditrysian

Lepidoptera (Insecta). Zoomorphology, 119, 163±183.

Kristensen, N.P. (1999) The non-glossatan moths. Lepidoptera, Moths

and Butter¯ies, 1: Evolution, Systematics and Biogeography (ed. by

N. P. Kristensen), pp. 41±49. Walter de Gruyter, Berlin.

Nielsen, E.S. & Common, I.B.F. (1991) Lepidoptera. Insects of

Australia (ed. by CSIRO), pp. 817±915. Melbourne University

Press. Melbourne.

Nielsen, E.S., Edwards, E.D. & Rangsi, T.V., eds (1996) Checklist of

the Lepidoptera of Australia. Monographs on Australian

Lepidoptera 4. CSIRO, Melbourne.

Nijhout, F.H. (1991) The Development and Evolution of Butter¯y Wing

Patterns. Smithsonian Institution Press, Washington, DC.

Parsons, M.J. (1998) The Butter¯ies of Papua New Guinea. Their

Systematics and Biology. Academic Press, London.

HAWKMOTHS OF THE WORLD: AN ANNOTATED AND

ILLUSTRATED REVISIONARY CHECKLIST (LEPIDOP-

TERA: SPHINGIDAE). By Ian J. Kitching and Jean Marie

Cadiou. Cornell University Press, London. Pp. viii + 226, 8

colour plates. £72.50 (cloth). ISBN 0-8014-3734-2.

The name Sphingidae comes from Linnaeus's genus Sphinx,

named by the great man because the front of the larva is

frequently raised in the manner of the Egyptian ®gure. Not

only are sphingid larvae conspicuous (they also, typically,

sport a `horn' on the rear of the body), but so too are the adult

moths with their narrow wings and rapid ¯ight. Indeed, it is

from the appearance of the adult that the colloquial name of

`hawkmoths' is derived.

This is not a big book in terms of length, but its scope, style

and treatment are magisterial. And why do I describe,

unhesitatingly, this `checklist' as such? First, because the work

is comprehensive, dealing, as it does, with all species of

hawkmoths (the best known of moth families); second, because

it is soundly constructed and produced; and third, because

taxonomic decisions are explained succinctly and with elegance

and thoughtfulness. The work is truly comparative, dealing with

the family on a world basis. Such breadth, together with the

authors' evident depth of knowledge of their group, are features

that make this volume so authoritative. The book is timely, for

not since the massive and wondrous world revision by

Rothschild & Jordan (1903) has there been a world view of

Sphingidae. Moreover, because hawkmoths are popular, much

has been published that lacks the understanding gained only by

comprehensive knowledge. Such knowledge comes largely as a

result of access to all the relevant collections and taxonomic

literature. This work may be, at its core, a list, but the

annotations transform it into an in-depth treatment.

The book is divided into three sections: an Introduction

of thirty-four pages, the Checklist itself, which runs to

thirty-seven pages, and, lastly, well over 100 pages of

`Notes' cross-referenced to the list. There are also eight

colour plates: six of these are of cabinet-set moths; two are

of live moths photographed in resting postures. The

hawkmoths illustrated were selected chie¯y to give visual

references to cases where there were notable taxonomic

problems.

The Introduction is composed largely of a thoughtful essay,

which introduces the hawkmoths and provides a description of

the adult and immature stages. General aspects of the biology

of these insects are described, including a critical comparison

of their life-styles compared with those of their closest

relatives, the emperor moths (Saturniidae). This comparison

was based on Dan Janzen's intriguing contrast of sphingid and

saturniid modes of life in Costa Rica (Janzen, 1984). Kitching

and Cadiou add an additional dimension to these observations

by showing that those hawkmoths belonging to tribe

Smerinthini share many features of saturniid biology, thus

suggesting that the life-styles of the two families are not so

sharply de®ned as might be thought. The story deserves a

wider airing both because it provides a richer picture of the

moths and also because it demonstrates the explanatory value

of phylogenetic systematics in evolutionary biology. Other

topics addressed in the Introduction include the role of

hawkmoths as pollinators and their economic impact. There

are summaries of rarity issues, hawkmoth conservation,

classi®cation, distribution and faunal works. These sections

are followed by a critical examination of previous catalogues

and checklists and an explanation and justi®cation of the way

in which taxonomic decisions were adopted in the work under

review. The authors provide an explicit description of their

criteria for treating species-level taxa as species, subspecies or

infrasubspeci®c variants. All lepidopterists would bene®t from

a careful reading of this excellent and thoughtful text. There

are no ®gures in the Introduction, so readers with a reasonable

knowledge of lepidopteran anatomy will be better placed to

appreciate the ®ner points in the family description. But the

authors provide key references to sources of illustrations.

The detailed taxonomic argumentation is collated in the third

section of the book under the understated heading of `Notes'.

Each one of the 627 notes refers to a numbered entry in the

checklist. Most function to explain the taxonomic actions taken

by the authors. The issues addressed are often complex, and it is

to the great credit of Kitching and Cadiou that they provide

consistently clear and detailed justi®cations for their decisions.

You cannot write rich taxonomy without respect for what has

gone before, and the extent of taxonomic and nomenclatural

matters resolved in the notes is telling, revealing the authors'

intimate knowledge of the species and their taxonomic history.

All previously designated lectotypes are cited and new lectotype

designations are made in the work where they are required to

clarify identities. As a result of resolving particular taxonomic

problems, the authors describe two new species and two new

subspecies in the Appendix.

# 2001 Blackwell Science Ltd, Systematic Entomology, 26, 125±129

128 Book Reviews128 Book Reviews

Anyone valuing a scholarly work with the most reasoned

classi®cation of Sphingidae will want this book. Those

interested in the diversity of sphingid moths will need it if

they are to understand how, why and to what sphingid

names are applied. Apart from those interested in

hawkmoths, lepidopterists in general will welcome a

comprehensive list of the genus, and species-group names

to the most widely known of the moth families. By

studying this book, students and, indeed, those with

experience, will learn much about how to apply the rules

of biological nomenclature to real taxonomic situations.

Although not a stated aim of the authors, the content of

this book will add to a growing, but as yet somewhat

scattered, databank of Lepidoptera (we are not far from

having most available species-group names of the order

computerized). The book will surely help more widely the

documentation of insect biodiversity at the species level.

This is the kind of study best done in great museums ± the ®rst

author works in one, the second has a close association with it

and access to its material. Indeed, the book shows us just what

can be achieved if taxonomists use to such effect the collections

and libraries that are unique to such institutions. Kitching and

Cadiou have managed to document a slice of biodiversity (and a

conspicuous one at that), an activity persistently demanded by

prominent bio-environmental politicians across the globe. They

have done so with care and attention to context and detail in a

book nicely produced by Cornell University Press. It is dif®cult

to imagine a more appropriate and socially useful activity for

the modern day systematist than producing a practical and

scholarly book of this kind.

MALCOLM J. SCOBLE

References

Janzen, D.H. (1984) Two ways to be a tropical big moth. Santa Rosa

saturniids and sphingids. Annals and Magazine of Natural History

(8), 20, 230±231.

Rothschild, L.W. & Jordan, K. (1903) A revision of the lepidopterous

family Sphingidae. Novitates Zoologicae, 9(Suppl.), 1±972.

# 2001 Blackwell Science Ltd, Systematic Entomology, 26, 125±129

Book Reviews 129Book Reviews 129