<oological Journal of the Linnean Socieg (1992), 104: 89-96
Reviews of publications
Worms to Wasps, an Illustrated Guide to Australia’s Terrestrial Invertebrates, by Mark Harvey
and Alan Yen, illustrated by Graham Milledge. Oxford: Oxford University Press, 1990.
202 pp. Hardback E l 1.95. ISBN: 0-19-553081-0.
Potential purchasers of paperback books select them because of a colourful or welldesigned jacket; and they buy them after a flick through the pages, from back to front, to
assess the look of the contents. O n this basis, Harvey and Yen’s work will sell and sell.
The front cover bears striking colour portraits of four representatives, the intriguing
Peripatus, perhaps the most memorable of all, though no list of the species portrayed is
given. Nor will you find one inside, for the book is an identification guide only to order
level. Thus the entire order Lepidoptera is covered by a single study of a downy moth.
No butterfly is included in the book. There is a single beetle, too but no suggestion of its
size in real life.
It is clearly sensible to plan a work which takes one down to the order, and the
bibliographical references provided with each section lead the user to more detailed
sources. But the problem comes when plan is translated into practice. Thus the single
opionid shown is typical of the type; all harvestmen are reasonably similar. The same is
true of the neuropteran flies, so aptly named lacewings; and also of the dermapterans
known throughout the English-speaking world as earwigs. However, the familiarity
breaks down with other orders. The leeches of the Arhynochobdellae are a varied bunch
both in size and morphology, and so are the triclad turbellarians; and the single dipteran
fly is a singular specimen, unlike many others. The captions, ‘Diptera-Fly’ or
‘Phthiraptera-Louse’ seem vague, and it may be unfortunate that there is no specific
identification of the types selected as examples.
That said, this is a beautifully illustrated book. The drawings are all line-and-stipple,
a style I have used to illustrate published work, but which is here employed with such
sheer artistry and elegance as to discourage amateur imitators in the future. Milledge has
scaled his diagrams and figures so that they retain proportionality of line thickness in the
printed form. This degree of conscientious professionalism marks him out as a superb
draughtsman. One fails to find slips of the pen, too; clearly he knows his subjects well.
As a student one was regularly regaled with the need to refer to the printed word, to
taxonomic criteria that the brain should acquire before the eye was informed. In the real
world we recognize species and types because of the way they appear, and a clear
picture-book is by far the easiest way to carry out on-the-spot identifications.
Arthropods decorate salads in Australia, sometimes as ingredients selected by the
barbie-chef, at other times acquired through the exigencies of gravity operating on the
myriad of insects that investigate the lamp hanging over the dining-table on the
verandah. For those who wish to know more, this is a handy introduction to all the
orders known to occur in Australia. And the arbitrary selection of some of the examples
is a minor irritation when the task of production has been done so well.
BRIANJ. FORD
0024-4082/92/010089 +08 $03.00/0
89
0 1992 The Linnean Society of London
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Comparative Social Organization o f Wild Reindeer in Relation to Food, Mates and Predator
Avoidance, by T. Skogland. 1989. Berlin and Hamburg: Paul Parey. 74 pp. Price DM52
soft cover. ISBN 3-489-63236-2.
This monograph is presented as number 29 in the series: Advances in Ethology
[Monographic supplements to the journal Ethology]. As its title suggests it is a
comprehensive account of the social organization and behaviour of wild reindeer. Based
in essence on the results of Skogland’s own studies of the behaviour and ecology of six
wild and three semi-domesticated reindeer populations it treats these original data
within the established context of the published accounts of other authors. It thus offers
the reader a thorough review of the literature, with the added focus/flavour provided by
the input of Skogland’s own research material.
Petrie (1986) considers social organization as: “a description of two main aspects of
animal existence: i) the spacing pattern of the individuals (e.g. whether they form
groups) and ii) the form of the mating system (e.g. whether individuals are monogamous
or polygamous)”. This volume reviews group size, sexual segregation and mating
strategies among reindeer and seeks to explain the strategies adopted in relation to:
Group size: distribution pattern of food and competition for food; calving strategy,
including synchronization of calving and protection of neonates; general antipredator
adaptations.
Sexual segregation; male courtship strategy: predator avoidance, monopolization of
females and strategies of male choice.
In general I found the exploration of this issue plausible and thought provoking and I
found the book a delight to read: authoritative, concise and well-crafted. The
presentation is excellent-and the volume itself is well-produced. The presentation is
rather that of a thesis offering separate sections of: Introduction, Methods, Results,
Discussion. As a specific monograph it is by design uncompromisingly focused on
reindeer and their behavioural ecology-and at times I found myself regretting that it
was quite so tightly ‘tuned’. Much of what Skogland has to say is every relevant to
general theories of population ecology and social organization in ungulates and I was
perhaps a little disappointed that he restricted himself so tightly to reindeer and caribou;
the standard of writing and discussion was such that, selfishly, I was sad not to read how
he might handle this more general material and how he reconciles his results within the
context of that wider literature on large ungulates in general. However, given the
constraints of the text-at 74 pages-the concentration on reindeer and caribou studies,
even in the more comparative sections of Introduction and Discussion, is perhaps
understandable.
As a monograph in the narrower area of the social behaviour and ecology of reindeer I
found the book definitive. The Introduction in particular clearly presents the theoretical
framework for the study, in general considers all the relevant published literature on
Rungifer [although I was surprised to see no reference to Nigel Leader-Williams’
extensive studies on South Georgia] and sets out very clear hypotheses to be examined.
Methods are unexceptional and Skogland’s detailed presentation of his results is logical
and clear. Neither I, nor any of my research students, to whom I showed the volume, felt
that we would challenge any of the conclusions drawn. Skogland’s own data are indeed
tremendously impressive and I found, throughout, his analysis of these original data
rigorous and searching.
The Discussion showed a broad familiarity with other literature-at least within the
confines of those publications specific to reindeer or caribou, and the interpretation of his
results on the responsiveness of grouping to food distribution, calving and antipredator
strategies seemed very reasonable. Skogland did not, however, offer any adequate
explanation for sexual segregation among reindeer, rehearsing instead some of the more
group-selectionist theories of, for example, Geist and Petocz (1977); nor did I find
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91
convincing the explanation offered for the dispersal of young males from maternal herds
to male-only groups as being “to facilitate the assessment of strength among the males”.
Otherwise I felt the material was in general well-handled. Overall, I found this an
excellent little monograph and one I would highly recommend to anyone generally
interested in sociobiology of large mammals.
RORYPUTMAN
REFERENCES
GEIST, V . & PETOCZ, R. G., 1977. Bighorn sheep in winter: Do rams maximise reproductive fitness by
,
1802-1810.
spatial and habitat segregation from ewes? Canadian Journal of ~ o o l o g y 55:
PETRIE, M., 1986. Reproductive strategies of male and female moorhens (Gallinula chloropus). In D. I.
Rubenstein & R. W. Wrangham (Eds), Ecological Aspects of Social Evolution: 43-63. Princeton: Princeton
University Press.
The Jackson Elk Herd: Intensive Wildlife Management in North America, by M. S. Boyce.
Cambridge: Cambridge University Press, 1989. 306 pp. ISBN 0-52 1-34147-7.
This volume is the latest in the Cambridge University. Press series of Studies in
Applied Ecology and Resource Management. I t offers a detailed analysis of the general
ecology of one population of North American ‘elk’ (in the U.S. elk is used for the wapiti
Cervus canadensis rather than Alces alces) in Jackson Hole, Wyoming and further considers
the effects of different management strategies applied to control and harvesting of this
population over the years. As such it potentially offers an interesting ‘case study’ or
worked example for general principles of wildlife management applied to populations of
large ungulates.
In practice, I felt that opportunity was lost. No attempt was made in the book to
establish any general principles, or to broaden consideration away from the elk of
Jackson Hole to consider the wider issues. Indeed, the whole reads perhaps as the sort of
research report and management plan I would expect to see in an internal end of
contract report to, let us say, the Wyoming Game Department, not a book aimed at a
general readership. I was frankly amazed that C U P should publish such a parochial
volume, which certainly makes no attempt to appeal to a wider audience.
Such a missed opportunity is a shame, for there is no doubt that there is a wealth of
scientific information now available on the elk of Jackson Hole and on the effects of
different management strategies (harvesting regimes, winter feeding practice etc.) upon
those populations, which could be used to exemplify more general principles of wildlife
management, rather than be related purely to the management problems of that one
population. Boyce, however, makes no attempt to do this; indeed by converse, he makes
little use of the wider published literature on red deer and wapiti in interpreting his own
data. The whole approach is somewhat isolationist.
After introductory chapters outlining the history of the population of elk at Jackson,
and changes in their numbers and management to the present day, Boyce presents basic
information on movement and migration patterns, habitat use and feeding. He then
considers the management implications of supplementary winter feeding and the
potential for habitat management, before moving on to address the effects of
management (by culling) of the elk population itself. (Curious that; I myself would have
to consider the impact of harvest upon the populations before considering implications of
habitat or food manipulation). He ends with a detailed chapter of conclusions and
management recommendations for the Jackson elk herd.
There is no question about the quality of Boyce’s data, nor of his scientific rigour.
However, I found the book disappointing in its lack of breadth and perspective. It
describes another autecological study not dissimilar to D. B. Houston’s The Northern
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Yellowstone Elk (1982-an
new.
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account of a single elk population!) and, I felt, added little
RORYPUTMAN
REFERENCE
HOUSTON, D. B., 1982. The Northern Yellowstone Elk: Ecology and Management. New York: Macmillan. 474 pp.
The Social Badger: Ecology and Behauiour of a Group Liuing Carnivore (Meles meles), by Hans
Kruuk. Oxford: Oxford University Press,
1989. 155pp. E15.95 HB.
ISBN 0-19-858703-1.
1989.
This book is a very welcome addition to the literature. It is not a general natural
history of the species but concentrates on giving an up-to-date picture of how badger
society works. I t is a well-written distillation of ten years’ research carried out by the
author and his helpers mainly in Scotland, and admirably reflects the enthusiasm,
ingenuity and endurance of those involved.
During the last two decades, field research has been revolutionized by a battery of new
techniques. Infrared or light intensifLing binoculars now enable detailed observations in
the dark, and with radio-telemetry badgers can be located and their movements followed
throughout the night. These and other devices, such as bait marking and the use of
radio-active isotopes have greatly helped the author in his research.
The book is an account of how the author set about the task of answering a number of
key questions such as: how do badgers organize their social life? How do they
communicate? Why do they live in groups? What benefits do they derive from being
territorial? What is the relationship between group size territory size, and food supply?
The reader is led through a logical sequence of question, investigation, deduction and
discussion.
This is a very personal account, and as the author points out, no attempt is made to
give detailed references to the work of other researchers, although useful suggestions for
further reading are made at the end of each chapter.
A very minor criticism is that the author does not conform to some terms in common
usage. ‘Social group’ is better than ‘clan’ which is apt for hyenas with up to 80 in the
group, but is inappropriate for badgers where group size is always small and in parts of
its range, often reduced to a single pair. Also when describing badger sounds, why invent
a new series of terms when adequate ones already exist? This can muddle.
However these points are relatively trivial and should not detract from the merits of
this admirable book which paints an excellent and comprehensive picture of badger
society.
ERNESTNEAL
Wildlge Production Systems: Economic Utilization of Wild Ungulates, edited by R. J. Hudson,
K. R. Drew and L. M. Baskin. Cambridge: Cambridge University Press, 1989. k55.00.
ISBN 0-52 1-34099-3.
The editors of this substantial volume based the book on papers presented at the 4th
International Theriological Congress, Edmonton, 1985, with some additional invited
contributions. As a reader, I had been hoping for something that I could recommend to
students as an illustration of how ecological principles, and especially principles of
population biology, could be used in the efficient exploitation of wildlife for food. At
present, students on our MSc course in Wildlife Management & Control at Reading
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93
University have to rely largely on Eltringham’s excellent book for a good, ecological
account of wildlife production, and I had been looking out for an additional text with
updated examples. I was therefore a little disappointed because this book emphasizes
other (though also important) aspects of wildlife exploitation. In particular,
socioeconomic aspects of subsistence hunting and sport are given extensive coverage in
the first half of the book, and much of the rest is case studies of various farming systems
with an emphasis on the structure of the industry (economics, scale and growth rates).
Population density is referred to in relation to ungulate stocking densities, moose
husbandry, body size allometry and fencing costs per stock unit, while regulation appears
many times but only in relation to legal rather than ecological constraints. This is not,
then, an ecological book though it is part of a series on applied ecology and resource
management.
Even so, applied ecologists will find plenty of material to interest them. Subsistence
hunting apart, economics must play a part in any exploitation system. The well-known
conflict that has to be resolved is between the emphasis on the annual balance sheet and
efficient use of capital (high return, rapidly) and the long-term conservation and hence
sustension of an exploitable resource. Ecologists who read this book will learn about
extensive ranching systems in north America and southern Africa and intensive
containment of game in other continents. They should then be able to think about the
possible effects of these large disturbances of terrestrial ecosystems (large either in scale or
intensity, or both), and some ecologists may even come up with useful research
proposals, or models that will help predict long-term consequences of wildlife
exploitation. If they read the chapters on subsistence hunting, conservation and
appropriate technology, they should also realize some of the limitations to the
application of ecological theory in the real world.
Unfortunately, real-world economics may limit the availability of this book to
ecologists. At g55 it is rather expensive for individuals, and certainly few students will
buy it. Though not an ecological book it is of considerable interest to ecologists and on
the whole the varied contributions deserve to be read.
ROBERT
H . SMITH
Sessile Animals of the Sea Shore, by Vernon A. Harris, London: Chapman and Hall, 1990.
ISBN 0-412-33760-6.
The marine littoral is a complex habitat with a rich and varied fauna, and a
consideration of the ways in which prevailing geophysical variables have shaped the
biology of the different animal species is no mean undertaking. By attending only to its
sessile members, Vernon Harris reduces his task to more manageable proportions, and
his terms of reference are precisely defined in the title of the book. Hydrozoans and
anemones for example, are included while related reef building corals are not. In
contrast the book’s objectives are less clearly stated, and must be inferred retrospectively.
The conceptual unity of the group derives from the sessile life style of its members, and
they are of interest collectively for the ways in which different phylogenetic archetypes
have responded to the challenges presented by such an existence. However, much of the
coherence inherent in a comparative approach is lost in the present work, in which the
biology of representatives of the more common groups of sessile animals are considered in
a systematic manner. Any underlying thesis thus falls second to the wealth of structural,
biochemical and physiological detail regarding methods of attachment, protective
exoskeletons, climatic tolerance, feeding mechanisms and reproductive strategies, which
forms the main body of the text. This apart, the book is well researched, focusing on
developments within the field over the last three decades. It is lucidly written, with many
original line drawings, and provides a most useful reference work for any serious student
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of marine biology. Should it run to a second edition, it is to be hoped that the annelid
giant axon will be described as such, not as a nerve, and Pembrokeshire restored to its
rightful location on the Welsh coast.
ELFEDMORGAN
The Natural History of Weasels
Stoats, by Carolyn King. Bromley: Christopher Helm,
1989. 253 pages. E15.95. ISBN-9-7470-1800-6.
&
I
The weasel, so they say, is the Nemesis of Nature’s little people. That is probably the
opinion held by the hapless moorhen depicted in Carolyn King’s book, flapping along in
laboured flight with a determinedly predatory weasel swinging below, its teeth clenched
in the bird’s chest. However, whatever plans the weasel might have had in mind for
landing its catch had probably not included the moorhen’s next move: to dive, weasel
and all, into deep water. Stoats, too, are notoriously predatory, reputedly flinging
themselves into gymnastic convulsions which are said to mesmerise simple-minded
rabbits. It seems that the victims, so called ‘stoated rabbits’, often die from fright rather
than from the coup de grice at the finale of the stoat’s dance. But does the dancing stoat
really have rabbits in mind during the performance? Perhaps not. Stoats and weasels
both suffer the attention of a parasitic worm which burrows in their skulls and produces
a condition called skrjabingylosis. Skrjabingylosis is probably even more painful to
endure than it is to pronounce, so the convulsing stoat may be demented by pain, rather
than playing to the crowd. This unromantic proposal remains to be tested, but what is
certain is that larvae of the parasite lurk, encysted, in the head muscles and salivary
glands of mice and voles: a neat adaptation to the fact that weasels begin their meals at
the head, and may leave the viscera uneaten.
These intriguing snippets are characteristic of the diversity of information in this book.
However, this is not merely a book in the ‘fascinating facts’ genre. Rather, Carolyn King
seeks to organize the facts into a synthesis that exposes the principles that make weasels
tick. What, by the way, are weasels? For the purposes of this book they are the stoat, the
long-tailed weasel (of North America), and the two subspecies of the smallest member of
the genus Mustelu, namely the least and common weasels. Collectively, all four are
termed weasels.
Stoats, of course, are widely known by their stage name, ermine, due to their rBle on
royal shoulders. It turns out that there is much more to the business of turning white in
winter than meets the eye. Whether or not stoats blanch in winter is determined by
temperature. It just so happens that the crucial temperature is such that white stoats are
common in Scotland and Wales, but not in England, due to the warming influence of the
Gulf Stream. On the borders, a piebald compromise reflects the climatic uncertainty. For
weasels the story is rather different. British weasels almost never turn white, indeed nor
do any south of Stockholm. There, however, things change in a 100 km wide zone where
some turn white and others remain stolidly brown. This zone is the meeting of two
subspecies: the northerly least weasel, and the southerly common weasel.
But what of Dr King’s synthesis? Weasels are small, long and thin. This is because they
evolved as specialist hunters of rodents, able to follow their prey beneath the snowy
blanket of northern winters and down their burrows too. Designed like an animated
sausage, the serpentine weasel can penetrate into the inner sanctum of a vole’s
home-adding insult to injury by eating the occupant, snug in the comfort of the
deceased’s own nest. Nowhere is safe for a vole when weasels are around: anywhere a
weasel’s head can go, its body can follow. However, as Carolyn King puts it, in nature
death is part of life. The need to squeeze through crannies keeps weasels small, and the
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95
need to minimize food requirements during the breeding season reinforces the
advantages of miniaturization to females. Males, on the other hand, who do nothing
whatsoever for their young, aside from supplying their genes, operate in a fierce arena of
sexual rivalry. In the battle between males for females, muscle power is an asset, but at
odds with the demands of burrow hunting. The solution is a compromise, with males
being almost twice the weight of females. Although the book includes many excursions
up interesting byways, the route charted by the author steadily clocks up the mileposts in
her argument: almost everything about weasels can be explained in terms of their
specialization in a high-risk, high-reward, diminutive rodent prey.
So the demands of food and sex together provide coherent explanation of the
differences in size between the sexes, but what about the differences in size between the
species? How can big species, like the stoat, co-exist with small ones like the common
weasel? Carolyn King’s suggestion is that they can only co-exist in an unstable world:
when rodents become scarce the common weasel, with its smaller demands for food, can
hang on for longer in the face of shortage, and is quicker off the reproductive chocks
when conditions improve. In contrast, under conditions of plenty, the larger stoat’s brute
force can easily bully its diminutive cousin off the scene. So, the argument runs, their
coexistence depends on a salient feature of life as a mouse hunter: numbers of prey
fluctuate widely.
Both weasels and stoats have found ways of coping with this boom and bust economy,
but their solutions are very different. Weasels can have two litters a year, so in a good
year, females born early in the spring can be producing grandchildren for their mother
by the time she is producing their younger sisters 5 months later. Stoats do things
differently, probably because they are bound to a system called delayed implantation:
that is, eggs fertilized in one year go through a period of suspended animation for 10
months before being born the next. They, therefore, cannot produce two litters a year,
but they can get a head start in a most remarkable way. Not only do female stoats come
into oestrus within 3 weeks of giving birth, but their nestling daughters do so too! A male
who forces his way into the family nest not only mates with the mother, but cradle
snatches her daughters too.
Dr King’s synthesis is convincing. It is also clearly and charmingly written and the
author openly applauds her subjects. The book is a personal narrative of her own voyage
of discovery, from her days as a graduate student a t Oxford. The opening pages include
a tribute to her then supervisor, H. N. (Mick) Southern, and those of us who knew him
realize how justified that tribute is. The book is illustrated with line drawings by Larry
Douglas. It is not seriously marred by the fairly numerous typographical errors that
smell of editorial carelessness in contrast to the meticulously worded text-two maps
only make sense when you realize that their legends are the wrong way around, and part
of the only formula in the book is missing. O n the assumption that Dr King knows which
way up North America goes, I hazard the guess that she was not too pleased with
whoever printed it upside down in her book.
My final thought about this very interesting book concerns the anonymity of the main
characters. The story of weasel society is clearly told, but always in terms of
generalizations. Students of weasel behaviour d o not seem to have explored their subjects
as individuals. There are none of the case histories of machiavellian interpersonal
intrigue that has emerged from studies of alliance and enmity in the social cliques of, say,
lions and wolves. This may be because the solitary sexist world of weasels is simply less
socially idiosyncratic than that of clannish Carnivora, but I doubt this is the explanation.
Rather, I guess that small size and elusive habits have protected the weasel’s anonymity
and thwarted the fieldworker’s voyeuristic enthusiasms. It is not to detract from this
excellent book one jot by saying that I hope in some future edition to discover more
about weasels as individuals.
DAVIDMACDONALD
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Bird Flight Performance: A Practical Calculation Manual, by Colin J. Pennycuick. Oxford:
Oxford Science Publications, 153 pp. Hardback: E25.00. ISBN 0-19-857721-4.
Over the past 10 years there has been great interest among ecologists in estimating
daily energy budgets in birds and also in estimating the costs of different activities.
Everybody knows that flight is a common habit in birds during foraging as well as for
transportation, but to estimate the cost of flight usually gives the investigator a headache.
Colin Pennycuick, who is an experienced pilot on motorcraft as well as gliders, has been
working on bird flight for more than 20 years. He was one of the first who used
aerodynamic principles to estimate the power consumption of animal flight, and he has
made flight mechanics, morphology, and behaviour understandable for biologists and
others in admirably innovative and clear scientific papers.
Bird Flight Performance includes Colin Pennycuick‘s aerodynamic models for estimating
the power requirements for flapping, gliding and soaring flight. His various aerodynamic
models for calculating flight performance have previously been compiled in a chapter of
Avian Biology, Vol. V (D. S. Farner and J. R. King, Eds, Academic Press, 1975),
‘Mechanics of Flight’, and this has been the basis for the computer programs of the disc
included in this book. In the first chapters the author gives an instructive and lucid
introduction to flight mechanics and describes the fundamental variables important for
flight, as well as the work and powers required and available for flight. The total power
is the sum of several components and the theoretical graph of the sum power curve as a
function of flight speed is U-shaped. From this curve the minimum power and maximum
range speeds can be defined in still air as well as in a head-wind or in a tail-wind. Using
only a few variables, such as body mass, wing span, flight speed, air density and profile
drag coefficient (estimated or measured), Colin Pennycuick has developed simple
expressions for the various power components and optimal flight speeds, using
momentum jet and steady-state aerodynamic theories.
After the three first chapters the reader should be ready to use the enclosed BASIC
programs for flight calculations. The fourth chapter tells in detail how to run the
different programs. Program 1 is designed for those who wish to calcualate the energy
requirements of a flying bird, carrying food back to its nest, and to estimate the flight
range of a long-distant migrant, given the amount of body fat available as fuel. Program
1A is an alternative that includes mechanical power only, for which muscle performance
is the point of interest. Program 2 is for gliding performance calculations. Chapter 6
outlines a basis for this program and is recommended to be read before making use of it.
Appendix 1 shows the listings of all these programs. Careful instructions on how to use
the programs are given both in the book text and on the disc. The user can change
variables in a pre-recorded program disc, such as air density and the added mass of a
radio transmitter.
In chapter 5 the physical processes underlying the aerodynamic model used are
explained in more detail, so that the reader can “appreciate the nature of the
simplifications that have been made, and see in qualitative terms where errors are likely
to arise”. This includes Reynolds number, inertial and biscous forces, drag forces and
drag coefficients, airflow around the wings, and vortex wakes. Chapter 7 deals with the
basic physical principles, of force, work, and power generation by muscles, and the
scaling of the power required and available. The last chapter (8) gives example
calculations.
Bird Flight Performance is lucid and stimulating to read, and I highly recommend this
book to any biologist who wants to get insight into bird flight or to calculate flight
performance. The book also contains several splendid black-and-white photographs of
flying birds, taken by the author, showing various interesting details of the wings in
various modes of flight.
ULLAM. NORBERG