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Cultural Transmission and Evolution: A Quantitative Approach, L.L. Cavalli-

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956 ANIMAL BEHAVIOUR,
The six chapters in Volume 13 cover a wide range of
topics, including hibernation, activity patterns, reproduc-
tion, life histories and growth. Bennett's chapter on
energetic constraints on activity covers an area that
should be considered much more deeply than it is by those
who are concerned with functional explanations of be-
haviour. Functional arguments often ignore the fact that
animals are limited in what they can do by both environ-
mental and physiological constraints. This is particularly
apparent in reptiles. Like Huey, Bennett adopts a cost-
benefit approach, in this case to aerobic and anaerobic
respiration. Whereas aerobic respiration can support low
levels of activity over long periods, activities that are, of
necessity, very vigorous, such as escape from a predator
or fighting, involve a switch to anaerobic respiration
which, because of its physiological costs, can only be
sustained for short periods.
Duvall, Guillette & Jones review work on the environ-
mental control of reptilian reproductive cycles, which has
gone a long way towards understanding the interactions
that occur between proximal environmental factors artd
various endocrine and neuroendocrine events. There are
interesting comparisons to be made here with work on
birds and mammals, though these are not considered in
this chapter. Congdon, D u n h a m & Tinkle discuss energy
budgets and life history strategies and explore the ener-
getic 'choices' made by individuals, according to their sex
and age, in relation to their environment and life history.
They conclude that, though developments in this area are
promising, future work requires study of the energy
budgets of individual animals, which, they suggest, is not
yet technologically feasible. Again, those who conduct
studies of this kind on birds and mammals will find much
of interest in this chapter.
All the chapters in these two books are reviews and
generally provide very thorough and up to date coverage
of their various topics. The reference sections are impres-
sively extensive. The emphasis throughout both books on
the ecological aspects of reptile physiology leads naturally
to frequent considerations of behaviour. Many etho-
logists, sadly, have only a rudimentary understanding of
physiology. If there is to be a major synthesis of physio-
logy and ethology, then I suggest that it is very likely that
it will have its origins in studies of reptiles. For this
reason, I recommend that ethologists take the time to
browse in volumes which they might otherwise pass by.
TIM ~IALLIDAY
Animal Behaviour Research Group,
Department of Biology,
The Open University,
Milton Keynes MK7 6AA, U.K.
Animal Behaviour: Ecology and Evolution. By C. J.
BARNARD. London: Croom Helm (1983). Pp. 339.
Price s hardback, s paperback.
This book has many desirable features. As a review of the
current state of the art of ethology it is good and compre-
hensive. I know now where to turn in order to find a
reference to almost anything. The style is, in general,
friendly and enthusiastic. Reading this book gives a vivid
impression of the many revolutionary theoretical insights
and tectmiques that have recently entered ethology (e.g.
ESS, optimal foraging, arms race, Beau Geste, systems
models, extended phenotype). Starting with neuro-
physiology, presented (as with most authors) in a rather
dull, unimaginative and 'fact-dense' manner, the book
steadily improves, through learning theory and motiva-
tion models (quite well described) to mating, cormaauni-
31, 3
cation and social behaviour, where Barnard is, in my view,
at his best. As an account of evolutionary processes in
animal behaviour, written with an ecological ftavour, I can
certainly recommend it.
However, I seriously wonder whether Barnard has
taken on too much. The spectrum of subjects covered is
vast in the extreme, ranging from the sodium pump to
cognitive learning theory and from human language to
dung fly mating. This means that, so often, only a super-
ficial passing reference is possible. For instance, I don't
believe that one can usefully teach either the action
potential or cognitive learning theory in a page or so. Yet,
herein lies the author's dilemma. Not to include cognitive
learning theory would convey the impression that P a r i t y
had the last word on the subject of conditioning.
I have a few specific criticisms. Many of the diagrams
are inadequately explained or the text fails to integrate
with the diagram. Miller's model of the parsimony of a
single drive is introduced, without it being said that
Miller's evidence was against such a model (page 66).
A distinction is drawn between hormonal effects on the
nervous system and their effects on sensory perception.
However, some of the effects described under 'sensory
perception' work via the nervous system (pp. 36-37).
In short, many of my criticisms result from packing too
much into too little space. For better or worse, ethology
has simply grown too fast, and yet, paradoxically, a strong
emergent property of this book is that it conveys very
well indeed the richness of ideas and observations in
contemporary ethology. It was a pleasure to read it.
FREDERICK TOATES
Biology Department,
The Open University,
Walton Hall,
2VIilton Keynes MK7 6AA, U.K.
Cultural Transmission and Evohttion: A Quantitative
Approach. By L. L. CAVALLI-SFoRZA ~6 M. FELDMAN.
Princeton, New Jersey: Princeton University Press
(1981). Pp. 368. Price $25.00 hardback, $10.50 paper-
back.
The study of human social behaviour has been, for the
most part, the province of anthropologists and socio-
logists. With the rapid rise of sociobiology, however,
many biologists became interested in analysing h u m a n
cultural phenomena with techniques and theories which
had been successfully applied to other species (such as kin
selection in social insects). The reaction to these attempts
in traditional circles was luke-warm at best, and occasion-
ally openly hostile. Fortunately, with the publication of
Cultural Transmission and Evolution, the process of apply-
ing biological models to anthropology has entered a new
and potentially more productive phase. The excesses and
occasional naivet6 of the more radical sociobiologists
have now given way to a much more cautious and con-
servative approach to h u m a n behaviour. In fact, it is not
clear whether these authors wish to be identified with
sociobiology at all. For the most part their work, much
of which has been published before, has followed its own
course, remaining untouched by the recent development
of sociobiology. Indeed there is practically no mention
made of genetics in this book, and the authors clearly
wish to avoid the issue of genetic determinants of be-
haviour, which so clearly pervades other recent work on
human social behaviour. Since the book contains only
the briefest discussions on the universal characteristics of
human behaviour and culture and very little on within-
species cultural variation, it is in no way an essay on
 BOOK REVIEWS
H u m a n Nature. Instead, it is a mathematically-oriented
treatise on the population dynamics of traits which are
not transmitted via traditional genetic mechanisms. As
defined rather broadly by Cavalli-Sforza & Feldman,
cultural transmission refers to any of these non-genetic
mechanisms of transmission.
Rather than delving into the details of the cultural
transmission process, the authors concentrate on broader
issues. Recognizing how little is known about the mech-
anisms of cultural transmission, Cavalli-Sforza &
Feldman postulate a set of abstract transmission co-
efficients which express the probability that an individual
will adopt a trait as a function of its contacts with other
members of the social group. This kind of approach can
be criticized as mere phenomenolngy, but the insights
gained later on in the work justify (at least up to a point)
the level of abstraction. In addition, as the authors point
out, Darwin developed his ideas on biological evolution in
the absence of any correct knowledge of the mechanisms
of genetic transmission, and modern cultural theorists
find themselves in a somewhat similar situation.
The book deals with three broad categories of cultural
evolution: vertical (from parent to offspring), horizontal
(between members of the same age group or generation),
and oblique (t~om non-parental members of the parental
generation to offspring). By invoking certain essential
simplifying assumptions about the nature of the trans-
mission coefficients (e.g. that they remain constant over
long periods of time), the authors are able to construct
increasingly complex models of cultural evolution. Even
the simplest of these, which involves no natural selection,
random drift, 'mutation' (innovation), migration, or
assortative mating, generates more complicated dy-
namics than would a corresponding genetic model,
largely because of the very general nature of the trans-
mission coefficients (in fact, Mendelian inheritance can be
shown to be a special case of vertical transmission). But,
in due course, all of these 'forces' of cultural evolution are
introduced, and before long general conclusions become
extremely difficult to extract from the equations. In such
cases, the authors focus on special cases which admit
analytic solutions. In the last chapters the transmission
process is extended to include multiple state and con-
tinuous traits. As in the earlier chapters, the authors
apply their models to interesting phenomena such as
stratification, hierarchies, parent versus teacher learning,
and other factors which may have played a significant role
in human cultural evolution. Emphasis on more involved
models does make the results more difficult to absorb,
but this is preferable to severe oversimplification.
The large number of cultural transmission processes
stands in contrast to the relatively few modes of genetic
inheritance. Although the details of genetic transmission
do constrain the evolutionary process (as is becoming
increasingly evident), the lack of any single set of rules
governing cultural transmission indicates that the trans-
mission process itself will play a more central role in
cultural evolution than it appears to have played in bio-
logical evolution. The book does an excellent job of re-
vealing the vast repertoire of dynamics which can result
from cultural transmission. For example, peculiarities of
the transmission process can produce evolution which
runs counter to natural selection, and this book therefore
provides some theoretical justification for questioning the
role of adaptation in cultural evolution. Anthropologists
who pursue a functionalist (ecological) approach to
cultural change are aware of the limitations of the influ-
ence of adaptation, but they may be surprised to hear
these admonitions from biologists.
957
Readers familiar with the theory of population genetics
will also be familiar with most of the models presented in
this book. For instance, diffusion equations are em-
ployed to study the evolution of culturally-transmitted
traits in small populations in much the same fashion as
they are employed in population genetics. One weakness
in patterning the models after models from traditional
population genetics is evident in the sections in which
natural selection is applied to cultural evolution. Recent
work in theoretical sociobiology has demonstrated that
most social interactions result in frequency-dependent
selection. In their discussion of selection, however, the
authors always assume constant selection coefficients.
They do note that some forms of transmission are ex-
pressly frequency-dependent, but they ignore the very
relevant fact that many cultural traits which affect the
individual's social behaviour will also affect fitness in a
frequency-dependent manner, thus complicating matters
further (oddly enough, both authors have developed
the theory of kin selection by applying models of
frequency-dependent selection to the evolution of social
behaviour).
The utility of Cavalli-Sforza & Feldman's theory is
limited by our ignorance concerning the transmission
process itself. Because we as yet do not know the con-
straints on the transmission coefficients, we cannot make
many conclusive statements about which of the interesting
dynamic processes discussed in the book actually occur.
The authors use the term 'cultural selection' to refer to
the decision-making process by which behaviour is
learned, but aside from brief discussions in the introduc-
tion and epilogue, there is no attempt to model any of the
fine structure of this learning process. Admittedly some
cultural traits are transmitted under conditions which
involve little or no selection on the part of the recipient
(neonatal circumcision, for example), and these can
appropriately be analysed with simple transmission co-
efficients. However, for behaviour which involves signifi-
cant learning it is probable that the transmission process
can be decomposed further than has been done here. A
substantial body of literature exists in sociology and
psychology which deals with learning theory, and in the
past few years there has also been some significant work
in evolutionary biology on the importance of learning and
learning rules. Such work should certainly be incor-
porated into the structure of any theory of cultural evolu-
tion (the authors have promised a second volume which
may deal with this problem).
A refreshing aspect of this book lies in its application
of theory to data. Although empirical examples are
spread rather thin, they are each discussed carefully and
in some detail. The authors successfully convey a sense of
excitement about these early attempts at applying a
mathematical theory to cultural phenomena. Interestingly,
the most convincing examples axe ones in which the de-
tails of the transmission process are fairly well understood.
These include an analysis of the evolution of surnames in
Italy's Parma valley, the history of linguistic change, and
certain aspects of epidemiolngy. It may appear odd at
first to think of a communicable disease as a cultural
entity, but the authors make consistent reference to what
might be called the 'epidemiology of culture', which is an
interesting metaphor. As the diffusion of a n innovation is
analogous to the spread of an infectious disease, so may
the maintenance of customs and traditions be analogous
to endemic diseases. Some other interesting examples of
the theory are in the area of linguistics, where 'traits' are
at least semi-quantifiable. There is a detailed discussion
of the gradual evolution of languages over long periods of
 958 ANIMAL BEHAVIOUR,
time, and several interesting analogies are found to
principles in evolutionary biology. For instance, the
divergence of two languages from an ancestral source re-
sembles in many ways the classical model of allopatric
speciation, in which the divergence is mediated by cul-
tural isolation. Another similarity is seen between local
geographic variation in dialects and models of genetic
distance brought on by geographic isolation.
It is a rather curious fact that in none of the recent
works on cultural evolution written by biologists has any
mention been made of cultural transmission in non-human
species. Since all these authors have defined cultural
transmission broadly (as non-genetic transmission), they
have in no way excluded species other than man in which
many types of behaviour are transmitted socially. If
cultural transmission turns out to be an important factor
in the behavioural development of other species, socio-
biologists may actually find it useful to study the dynamics
of human cultural transmission as a key to other
organisms, instead of attempting the reverse. On the other
hand, field studies of cultural transmission in other species
(the celebrated Japanese monkeys of Koshima Islet come
to mind) might also serve as paradigms for the general
study of cultural transmission in humans. Fortunately
Cultural Transmission and Evolution provides an elegant
theoretical framework for either approach.
The authors are aware of both the theoretical and
experimental limitations of their theory, but have chosen
a somewhat simplified mathematical model because, as
they state in the preface, a mathematical approach is
always more precise than an expository one, if only
because each element of the theory must be rigorously
defined and related to the whole. Naturally, any attempt
to model a phenomenon with the inherent complexity of
h u m a n culture is invariably bound to be either an approxi-
mation or an over-simplification. But whereas matbe-
matical approximations can be steadily improved, verbal
arguments can often only be debated. It is in this spirit
that the book should be appreciated.
MICHAEL J. SANDERSON
RICHARD E. MIC~OD
Department of Ecology and Evolutionary Biology,
University of Arizona,
Tucson, AZ 85720,
U.S.A.
The Behavior and Natural History of the Caribbean Reef
Squld (Sepioteuthis sepioidea). By MARTIN MOYNIHAN &
ARCADIO F. RODANICHE. Berlin & Hamburg: Paul
Parey (1982). Pp. 152. Price $21.60 (paperback).
This is a remarkable book about an extraordinary animal,
for even to those of us who have studied them for a lifetime,
cephalopod molluscs are out of the ordinary and this
account of the natural history and social life of one of
them is arguably 'more nearly complete' than for any other
species. It is based on observations of the Caribbean reef
s q u i d - the name has been coined by the authors, I
g a t h e r - - w h i l e snorkeling in and around the San Bias
Islands on the Atlantic coast of Panama. What better way
for an ethologist to spend his days away from the office
and laboratory over a number of years, mostly in the
early 1970s. Moynihan is known for his contribution to
the literature on bird and primate displays and the verte-
brate ethologist has been able to bring to the description
of the behaviour observed a depth of meaning that others
would have missed and some, frankly, will find startling,
even unbelievable.
View publication stats
31, 3
Amongst cephalopods, whose organization is so remi-
niscent of vertebrates, squids generally show less elaborate
behaviour than do octopuses and cuttlefish. Sepioteuthis
sepioidea--whieh could equally well be called the
'cuttlefish s q u i d ' - - i s an exception and an example of
convergence within the cephalopods. As the many expert
photographs and drawings illustrate, this squid
( T e u t h i d a e ) - which has the wide body and circum-
ferential fins of the cuttlefish (Sepiidae) - - is capable of a
range of shapes, movements, colour patterns and ritua-
tized behaviour (categorized by the authors as social
signals, displays and antidisplays) as wide as that of any
cuttlefish. Each is described in detail and the description
lengthened many times by examination of the contexts
in which each occurs, including the context of other
animals. Groups of the Caribbean reef squid are often
found associated with goatfish and the groups may be
joined by other squid species e.g. Loligo.
The authors then proceed to an elaborate breakdown
of the meaning of the signals and displays, ritualized and
unritualized, based on the frequency and relative position
of each observed element, employing computer programs
to construct dendrograms.
It is of course no accident that these squids, which are
social and live in the richly visual environment of a
sheltered tropical reef, have rich repertoires of visual
behaviour. The authors have gone to the logical extreme
of assuming that every sequence and combination of
patterns is communicative and 'pursuing the grammatical
a n a l o g y . . , can be assigned to three different classes
which we will call signifiers, modifiers and positionals'.
Readers will want to judge for themselves whether this
bold attempt at semantic a n a l y s i s - trying to make the
signals i n t e l l i g i b l e - is justified at this stage of our
ignorance, but it is a refreshing change from the excessive
caution of so many accounts. Given the preponderance of
camouflage patterns in the repertoires of other cephalo-
pods (octopuses and cuttlefish) I am puzzled by the
almost complete absence of any assignment of crypsis to
the patterns of Sepioteuthis sepioidea. Perhaps the seman-
tic interpretation should include the possibility that the
squids are escaping detection as prey by sending un-
intelligible signals. That is what crypsis is about.
The illustrations are the best single feature of this
monograph. The photographs provide a more complete
documentation of the behaviour of a cephalopod in its
natural environment than we have for any of its better-
known relatives and Moynihan's graceful and meticulous
drawings carry the conviction due to an artist.
ANDREW PACKARD
Department of Physiology,
University Medical School,
Teviot Place,
Edinburgh EH8 9AG, U.K.
Avian Incubation: Egg Temperature, Nest Humidity, and
Behavioral Thermoregulation in a Hot Environment
(Ornithological Monographs, No. 30). By GILBERT S.
GRANT. Washington, D.C. : American Ornithologists'
Union (1982). Pp. ix+75. Price $9.00.
This monograph overviews thermoregulatory problems
faced by birds incubating their eggs in hot, arid environ-
ments. The study focuses on the behavioural and physio-
logical adaptations used during incubation by seven
Charadriifornaes (black-necked stilt, Himantopus mexi-
canus: American avocet, Recurvirostra americana; snowy
plover, Charadrius alexandrinus; killdeer, C. vociferus;
gull-billed tern, Gelochelidon nilotica; Forster's tern,