- Email: [email protected]
How monkeys see the world: Inside the mind of another species
TREE vol. 5, no. 12, December
chapter. This separate and rather fragmentary treatment of modular clones muddles synthesis of the ecological significance of clonality, and is the only major disappointment in the volume. The review of obligate cloning is the highlight of the book. It is the most lucid and economical treatment anywhere of the characteristics and taxonomic distribution of obligately cloning animals. Virtually all obligate cloners are parthenogenetic, having arisen by mutation in a bisexual lineage or by hybridization. Hybridization greatly increases heterozygosity, probably well beyond the limits of heterosis. Most obligately cloning parthenogens are polyclonal or polyphyletic, either condition originating by mutation or hybridization. Astoundingly rapid shifts in genomic frequency are possible via interclonal selection, particularly in polyphyletic species, which may equal or exceed evolutionary responsiveness of bisexual populations. Obligately cloning species clearly do not lack genotypic variation or responsiveness to selection, but progress is by ‘freezing’ pleiotropic gene com-
plexes that are mere snapshots of recombinant genotypes pending new inputs of variation by further hybridization. Hughes defines two basic groups of obligately cloning species, distinguished by mode of origin and ecological characteristics. Monophyand polyphyletic forms letic clones, with elty,
infrequent live in
input
of genetic
nov-
extreme habitats and display wide ecological and physiological tolerances. Such ‘generalpurpose genotypes’ result from predominance of physical over biological selection, and may persist indefinitely. These are the majority of obligately cloning species in all taxa. In contrast, polyphyletic clones with frequent recombinant input by hybridization are ecological specialists. These develop as arrays of narrowly adapted clones with distinct ecological niches that appear to result from intense competition in physically unstable environments (e.g. the guppy Poecillopsis spp.) or by coevolution between herbivores and their hosts pometaris). (e.g. the moth Alsophila Lurking in these patterns are the (apomictic?) seeds of individual-
1990
selectionist explanations awaiting more formal treatment. Hughes’ important contribution is to have so successfully clarified the patterns themselves. Jeremy B.C. Jackson Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Republic of Panama
References 1 Ghiselin, M.T. (1974) The Economy of Nature and the Evolution of Sex, University of California Press 2 Williams, G.C. (1975) Sexand Evolution, Princeton University Press 3 Smith, J.M. (1981) The Evolution of University Press Sex, Cambridge 4 Bell, G. (1982) The Masterpiece of Nature: The Evolution and Genetics of Sexuality, University of California Press 5 Stearns, S.C., ed. (1987) The Evolution Birkhatiser of Sex and Its Consequences, Verlag 6 Grosberg, R.K. and Patterson, M.R. (1989) Paleobiology 15,67-73 7 Jackson, J.B.C. (1985) in Population Biology and Evolution of Clonal Organisms (Jackson, J.B.C., Buss, L.W. and Cook, R.E., eds), pp. 297-356, Yale University Press
CognitiveEcology How Monkeys See the World: Inside the Mind of Another Species edited by D.L. Cheney and R.M. University of Chicago Seyfarth, Press, 1990. $24.95 in USA ($28.75 elsewhere) hbk (x + 359 pages) ISBN 0226102459 As parliamentarians say, I must at the outset declare ‘an interest’. I lived and worked in the field with the subjects of this book, and its authors, for many months. Despite all this time spent observing vervet monkeys (Cercopithecus aethiops), it wasn’t until I read this book that I felt I could retrospectively have a greater insight into what the behaviour I watched might mean to the animals themselves and there were some surprises. Cheney and Seyfarth have posed the question asked by most humans watching animal behaviour: what do the animals themselves ‘think’ or ‘know‘ about what they are doing? They also ask whether it is appropriate to pose such questions. The focus on primates in this book is a result of a specifically evolutionary question: is the way we humans see the world and interpret, question and respond to others’ behaviour similar to, or part of a continuum with, that of non-humans? 426
Cheney and Seyfarth have been observing behaviour and doing field experiments with their subjects over the past 15 years. They were drawn to vervets by an intriguing system of anti-predator vocalizations, which appeared to be ‘naming’the predator and thus providing specific information in a social context. Their earliest work demonstrated that this naming had referential characteristics - it didn’t simply provoke a specific response, but provided information which could then be acted on depending on the context, degree of threat, possible escape routes and even characteristics of the individual who was vocalizing. They then went further, by demonstrating that the calls had first-order intentionality, in that the caller was vocalizing to influence the behaviour of others, not simply responding to a high-level stimulus provoking a vocal response. Their subsequent extensive work with social vocalitations has strengthened this conclusion. Sceptics will need to read the detailed procedures for field experimentation and see for themselves the careful analysis and interpretation of the results presented in this book. The authors have also provided a compendium of social knowledge among vervets and other non-human
primates. The social skills of monkeys and apes continue to astonish humans with their complexity and apparent sophistication. In the light of this book, we should now be able to take for granted the capacity of primates to solve social problems. But the question of what the monkeys intend to achieve and whether they have a mental representation of these intentions is not easily answered. Cheney and Seyfarth conclude that abstract concepts of relationships are not part of vervet monkeys’ mental capacities - they reserve these for humans (with our use of language) and the apes who have been trained either with language or computers. While higher primates recognize the relationships existing between others - dominance and subordinance, mothers and offspring, malefemale friendships, etc. - as well as their own relationships with others, these relationships do not exist as abstract concepts. They are context dependent and individually specific. In delving into the mind of a monkey, Cheney and Seyfarth have broadened the basis for approaches to studies of animal cognition. Firstly, and I think most importantly, they have investigated the meaning of behaviour and vocalizations in a natural context. Rather than asking the ver-
TfiFE vol. 5, no. 12, December
1990
vets to solve problems that humans have designed, or cope with human language, they have focused on problems that vervets themselves have to solve in their social and physical environment. In order to understand hc,w monkeys see the world, it is impc’rtant that the world is the typical acaptive context for that species. Hence, the language studies of apes at-d the problem-solving tests of olier species ask what the animals know or think in a context far rem Dved from any that may have had a selective role during the species’ evolutionary history. Results from naturalistic experimentation such as that of Cheney and Seyfarth can then bti extended to generate new and possibly more contextually appropriate experimental questions.
-
Secondly, they have illuminated the important and controversial issues of where the non-human primates lie on the continuum of cognitive capacities. Human linguistic and social capacities did not arise de novo. This book shows that they are more firmly rooted in the primate past than is thought by some. Representation, intentionality, deception and manipulation of others’ behaviour are all present in the communication and behaviour patterns of non-human primates. Equally important is what is lacking: a knowledge of others’ minds and motives abstractions of representations. Such concepts may have only arisen with language, and language is thought to be a recent evolutionary phenomenon, possibly arising as late as
the origin of anatomically modern humans some 200000 years ago. If this is correct then an understanding of the mind of monkeys and apes is fundamental to understanding the mind of humans during much of their five-million-year evolutionary history. It also provides us with the starting point for establishing the kinds of selective traits that would lead to the capacity for abstraction and the development of beliefs, and the origins of language.
Vernon A. Harris Sessile Animals of the Sea Shore Chapman & Hall, 1990. f37.50 hbk (x + 379 pages) ISBN 0 412 33760 6
J.A. McNeely, K.R. Miller, W.V. Reid, R.A. Mittermeier and T.B. Werner Conserving the World’s Biological Diversity IUCN, WRI, Cl, WWF-US and the World Bank, 1990. (193 pages) ISBN 0 915825 42 2
P.C. Lee University of Deptof BiologicalAnthropology, Cambridge,DowningSt, CambridgeCB23DZ, UK
Books Received
Review copies of the following books have been raceived. The appearance of a book in the list daes not preclude the possibility of it being reviewed in TREE in the future. John Alcock Sonoran Desert Summer The University of Arizona Press, 1990. $19.95 hbk (x + 187 pages) ISBN 0 8165 1150 0 Tim M. Berra Evolution and atronism: A Basic Guide to Erolution Debate Stanford 1990. $29.50 hbk, $7.95 pbk IS’3N 0 807 1548 3
rhe Myth of Crerhe Facts in the University Press, (xx + 198 pages)
D;i niel B. Botkin Discordant Harmonies: A New Ecologyforthe Twenty-firstCenturyOxford University Press, 1990. f17.50/$19.95 hbk (xii + 241 pages) ISBN 0 19 505491 1 A Z. Brown and A. McLachlan Ecology ofSandy Shores Elsevier, 1990. $97.50/Dfl.190.00 hbk (xii + 328 pages) ISBN 0 444 88661 3 Lynton Keith Caldwell Between Two Worlds: Science, the Environmental Movement and P~licv Choice Cambridae Universitv Press (C:smbridge Studies in EGvironmental’ Policy), 1990. f32.50/$44.50 hbk (xv + 224 pages) ISBN 0 ‘j21 33152 8 Neil A. Camobell Bioloov (2nd edn) Beniamin/ Cl.immings, i990. f19.9ghbk (xxx + 1165 pages) ISBN 0 8053 1800 3 S 4. Drury A Guide to Remote Sensing: Interpreting Images of the Earth Oxford University Press, 1990. f40.00 hbk. f20.00 pbk (viii + 199 pcges) ISBN 0 19 854495 2 B J. Ens, T. Piersma, W.J. Wolff and L. Zwarts (a Js) Homeward Bound: Problems Waders face w?en Migrating from the Bane D’Argun Mauritania. to their Northern Breedino Grounds in Spring international Wader and -Waterfowl Research and the Research Institute for Nature Management, 1990. (xii + 364 pages) ISBN 90 71”202 131 C. Harcourt and J. Thornback Lemurs of Madagascar and the Comoros: The /UCN Red D&a Book IUCN - The World Conservation U lion, 1990.
W. Hawthorne Field Guide to the Forest Trees of Ghana National Resources Institute, 1990. (vi + 275 pages) ISBN 0 902500 34 1 R. Hengeveld Dynamic Biogeography Cambridge University Press (Cambridge Studies in Ecology), 1990. f30.00/$54.50 hbk (xiv + 249 pages) ISBN 0 521 38058 8
G. Tyler Miller, Jr Living in the Environment (6th edn) Wadsworth Publishing Company, 1990. f16.95 (xviii + 620 pages) ISBN 0 534 12222 1 Dennis M. Power ted.1 Current Ornithology (Vol. 7) Plenum Press, 1990. $75.00 hbk (xiv + 388 pages) ISBN 0 306 43307 9
Roger N. Hughes ted.) Behavioural Mechanisms of Food Selection Springer-Verlag, 1990. DM 388.00 hbk (xii + 886 pages) ISBN 3 540 51762 6 D.H. Hutson and T.R. Roberts teds) Environmental fate of Pesticides (Progress in Pesticide Biochemistryand Toxicology, Vol. 7) John Wiley & Sons, 1990. f65.00 hbk Ix + 286 pages) ISBN 0471917117 Robert Edward Lee Phycology (2nd edn) Cambridge University Press, 1990. f17.50/ hbk (xv + 645 pages) $29.95 pbk, f37.50/$59.50 ISBN 0 521 36744 1
R.J. Lincoln and G.A. Boxshall The Cambridge lllusrrated Dictionary of Natural History (pbk edn) Cambridge University Press, 1990. f9.95/$14.95 pbk (iv + 413 pages) ISBN 0 521 39941 6 Francesca Lvman with lrvina Mintzer. Kathleen Courrier andJames MacKeniie The Greenhouse Trap: What We’re Doing to the Atmosphere and How We Can Slow Global Warming Beacon Press, 1990. (xvii + 190 pages) ISBN 0 8070 8503 0 J.M. Lynch led.) The Rhizosphere John Wiley & Sons (Wiley Series in Ecological and Applied Microbiology), 1990. f60.00/$110.40 hbk (xiv + 458 pages) ISBN 0 471 92548 9 Jennifer A. Marshall Graves, Rory M. Hope and Desmond W. Cooper teds) Mammals from Pouches and Eggs: Genetics, Breeding and Evolution of Marsupials and Monotremes CSIRO, 1990. A$50.00 in Australia (US$50.00 elsewhere) (xi + 337 pages) ISBN 0 643 05020 5
427
chapter. This separate and rather fragmentary treatment of modular clones muddles synthesis of the ecological significance of clonality, and is the only major disappointment in the volume. The review of obligate cloning is the highlight of the book. It is the most lucid and economical treatment anywhere of the characteristics and taxonomic distribution of obligately cloning animals. Virtually all obligate cloners are parthenogenetic, having arisen by mutation in a bisexual lineage or by hybridization. Hybridization greatly increases heterozygosity, probably well beyond the limits of heterosis. Most obligately cloning parthenogens are polyclonal or polyphyletic, either condition originating by mutation or hybridization. Astoundingly rapid shifts in genomic frequency are possible via interclonal selection, particularly in polyphyletic species, which may equal or exceed evolutionary responsiveness of bisexual populations. Obligately cloning species clearly do not lack genotypic variation or responsiveness to selection, but progress is by ‘freezing’ pleiotropic gene com-
plexes that are mere snapshots of recombinant genotypes pending new inputs of variation by further hybridization. Hughes defines two basic groups of obligately cloning species, distinguished by mode of origin and ecological characteristics. Monophyand polyphyletic forms letic clones, with elty,
infrequent live in
input
of genetic
nov-
extreme habitats and display wide ecological and physiological tolerances. Such ‘generalpurpose genotypes’ result from predominance of physical over biological selection, and may persist indefinitely. These are the majority of obligately cloning species in all taxa. In contrast, polyphyletic clones with frequent recombinant input by hybridization are ecological specialists. These develop as arrays of narrowly adapted clones with distinct ecological niches that appear to result from intense competition in physically unstable environments (e.g. the guppy Poecillopsis spp.) or by coevolution between herbivores and their hosts pometaris). (e.g. the moth Alsophila Lurking in these patterns are the (apomictic?) seeds of individual-
1990
selectionist explanations awaiting more formal treatment. Hughes’ important contribution is to have so successfully clarified the patterns themselves. Jeremy B.C. Jackson Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Republic of Panama
References 1 Ghiselin, M.T. (1974) The Economy of Nature and the Evolution of Sex, University of California Press 2 Williams, G.C. (1975) Sexand Evolution, Princeton University Press 3 Smith, J.M. (1981) The Evolution of University Press Sex, Cambridge 4 Bell, G. (1982) The Masterpiece of Nature: The Evolution and Genetics of Sexuality, University of California Press 5 Stearns, S.C., ed. (1987) The Evolution Birkhatiser of Sex and Its Consequences, Verlag 6 Grosberg, R.K. and Patterson, M.R. (1989) Paleobiology 15,67-73 7 Jackson, J.B.C. (1985) in Population Biology and Evolution of Clonal Organisms (Jackson, J.B.C., Buss, L.W. and Cook, R.E., eds), pp. 297-356, Yale University Press
CognitiveEcology How Monkeys See the World: Inside the Mind of Another Species edited by D.L. Cheney and R.M. University of Chicago Seyfarth, Press, 1990. $24.95 in USA ($28.75 elsewhere) hbk (x + 359 pages) ISBN 0226102459 As parliamentarians say, I must at the outset declare ‘an interest’. I lived and worked in the field with the subjects of this book, and its authors, for many months. Despite all this time spent observing vervet monkeys (Cercopithecus aethiops), it wasn’t until I read this book that I felt I could retrospectively have a greater insight into what the behaviour I watched might mean to the animals themselves and there were some surprises. Cheney and Seyfarth have posed the question asked by most humans watching animal behaviour: what do the animals themselves ‘think’ or ‘know‘ about what they are doing? They also ask whether it is appropriate to pose such questions. The focus on primates in this book is a result of a specifically evolutionary question: is the way we humans see the world and interpret, question and respond to others’ behaviour similar to, or part of a continuum with, that of non-humans? 426
Cheney and Seyfarth have been observing behaviour and doing field experiments with their subjects over the past 15 years. They were drawn to vervets by an intriguing system of anti-predator vocalizations, which appeared to be ‘naming’the predator and thus providing specific information in a social context. Their earliest work demonstrated that this naming had referential characteristics - it didn’t simply provoke a specific response, but provided information which could then be acted on depending on the context, degree of threat, possible escape routes and even characteristics of the individual who was vocalizing. They then went further, by demonstrating that the calls had first-order intentionality, in that the caller was vocalizing to influence the behaviour of others, not simply responding to a high-level stimulus provoking a vocal response. Their subsequent extensive work with social vocalitations has strengthened this conclusion. Sceptics will need to read the detailed procedures for field experimentation and see for themselves the careful analysis and interpretation of the results presented in this book. The authors have also provided a compendium of social knowledge among vervets and other non-human
primates. The social skills of monkeys and apes continue to astonish humans with their complexity and apparent sophistication. In the light of this book, we should now be able to take for granted the capacity of primates to solve social problems. But the question of what the monkeys intend to achieve and whether they have a mental representation of these intentions is not easily answered. Cheney and Seyfarth conclude that abstract concepts of relationships are not part of vervet monkeys’ mental capacities - they reserve these for humans (with our use of language) and the apes who have been trained either with language or computers. While higher primates recognize the relationships existing between others - dominance and subordinance, mothers and offspring, malefemale friendships, etc. - as well as their own relationships with others, these relationships do not exist as abstract concepts. They are context dependent and individually specific. In delving into the mind of a monkey, Cheney and Seyfarth have broadened the basis for approaches to studies of animal cognition. Firstly, and I think most importantly, they have investigated the meaning of behaviour and vocalizations in a natural context. Rather than asking the ver-
TfiFE vol. 5, no. 12, December
1990
vets to solve problems that humans have designed, or cope with human language, they have focused on problems that vervets themselves have to solve in their social and physical environment. In order to understand hc,w monkeys see the world, it is impc’rtant that the world is the typical acaptive context for that species. Hence, the language studies of apes at-d the problem-solving tests of olier species ask what the animals know or think in a context far rem Dved from any that may have had a selective role during the species’ evolutionary history. Results from naturalistic experimentation such as that of Cheney and Seyfarth can then bti extended to generate new and possibly more contextually appropriate experimental questions.
-
Secondly, they have illuminated the important and controversial issues of where the non-human primates lie on the continuum of cognitive capacities. Human linguistic and social capacities did not arise de novo. This book shows that they are more firmly rooted in the primate past than is thought by some. Representation, intentionality, deception and manipulation of others’ behaviour are all present in the communication and behaviour patterns of non-human primates. Equally important is what is lacking: a knowledge of others’ minds and motives abstractions of representations. Such concepts may have only arisen with language, and language is thought to be a recent evolutionary phenomenon, possibly arising as late as
the origin of anatomically modern humans some 200000 years ago. If this is correct then an understanding of the mind of monkeys and apes is fundamental to understanding the mind of humans during much of their five-million-year evolutionary history. It also provides us with the starting point for establishing the kinds of selective traits that would lead to the capacity for abstraction and the development of beliefs, and the origins of language.
Vernon A. Harris Sessile Animals of the Sea Shore Chapman & Hall, 1990. f37.50 hbk (x + 379 pages) ISBN 0 412 33760 6
J.A. McNeely, K.R. Miller, W.V. Reid, R.A. Mittermeier and T.B. Werner Conserving the World’s Biological Diversity IUCN, WRI, Cl, WWF-US and the World Bank, 1990. (193 pages) ISBN 0 915825 42 2
P.C. Lee University of Deptof BiologicalAnthropology, Cambridge,DowningSt, CambridgeCB23DZ, UK
Books Received
Review copies of the following books have been raceived. The appearance of a book in the list daes not preclude the possibility of it being reviewed in TREE in the future. John Alcock Sonoran Desert Summer The University of Arizona Press, 1990. $19.95 hbk (x + 187 pages) ISBN 0 8165 1150 0 Tim M. Berra Evolution and atronism: A Basic Guide to Erolution Debate Stanford 1990. $29.50 hbk, $7.95 pbk IS’3N 0 807 1548 3
rhe Myth of Crerhe Facts in the University Press, (xx + 198 pages)
D;i niel B. Botkin Discordant Harmonies: A New Ecologyforthe Twenty-firstCenturyOxford University Press, 1990. f17.50/$19.95 hbk (xii + 241 pages) ISBN 0 19 505491 1 A Z. Brown and A. McLachlan Ecology ofSandy Shores Elsevier, 1990. $97.50/Dfl.190.00 hbk (xii + 328 pages) ISBN 0 444 88661 3 Lynton Keith Caldwell Between Two Worlds: Science, the Environmental Movement and P~licv Choice Cambridae Universitv Press (C:smbridge Studies in EGvironmental’ Policy), 1990. f32.50/$44.50 hbk (xv + 224 pages) ISBN 0 ‘j21 33152 8 Neil A. Camobell Bioloov (2nd edn) Beniamin/ Cl.immings, i990. f19.9ghbk (xxx + 1165 pages) ISBN 0 8053 1800 3 S 4. Drury A Guide to Remote Sensing: Interpreting Images of the Earth Oxford University Press, 1990. f40.00 hbk. f20.00 pbk (viii + 199 pcges) ISBN 0 19 854495 2 B J. Ens, T. Piersma, W.J. Wolff and L. Zwarts (a Js) Homeward Bound: Problems Waders face w?en Migrating from the Bane D’Argun Mauritania. to their Northern Breedino Grounds in Spring international Wader and -Waterfowl Research and the Research Institute for Nature Management, 1990. (xii + 364 pages) ISBN 90 71”202 131 C. Harcourt and J. Thornback Lemurs of Madagascar and the Comoros: The /UCN Red D&a Book IUCN - The World Conservation U lion, 1990.
W. Hawthorne Field Guide to the Forest Trees of Ghana National Resources Institute, 1990. (vi + 275 pages) ISBN 0 902500 34 1 R. Hengeveld Dynamic Biogeography Cambridge University Press (Cambridge Studies in Ecology), 1990. f30.00/$54.50 hbk (xiv + 249 pages) ISBN 0 521 38058 8
G. Tyler Miller, Jr Living in the Environment (6th edn) Wadsworth Publishing Company, 1990. f16.95 (xviii + 620 pages) ISBN 0 534 12222 1 Dennis M. Power ted.1 Current Ornithology (Vol. 7) Plenum Press, 1990. $75.00 hbk (xiv + 388 pages) ISBN 0 306 43307 9
Roger N. Hughes ted.) Behavioural Mechanisms of Food Selection Springer-Verlag, 1990. DM 388.00 hbk (xii + 886 pages) ISBN 3 540 51762 6 D.H. Hutson and T.R. Roberts teds) Environmental fate of Pesticides (Progress in Pesticide Biochemistryand Toxicology, Vol. 7) John Wiley & Sons, 1990. f65.00 hbk Ix + 286 pages) ISBN 0471917117 Robert Edward Lee Phycology (2nd edn) Cambridge University Press, 1990. f17.50/ hbk (xv + 645 pages) $29.95 pbk, f37.50/$59.50 ISBN 0 521 36744 1
R.J. Lincoln and G.A. Boxshall The Cambridge lllusrrated Dictionary of Natural History (pbk edn) Cambridge University Press, 1990. f9.95/$14.95 pbk (iv + 413 pages) ISBN 0 521 39941 6 Francesca Lvman with lrvina Mintzer. Kathleen Courrier andJames MacKeniie The Greenhouse Trap: What We’re Doing to the Atmosphere and How We Can Slow Global Warming Beacon Press, 1990. (xvii + 190 pages) ISBN 0 8070 8503 0 J.M. Lynch led.) The Rhizosphere John Wiley & Sons (Wiley Series in Ecological and Applied Microbiology), 1990. f60.00/$110.40 hbk (xiv + 458 pages) ISBN 0 471 92548 9 Jennifer A. Marshall Graves, Rory M. Hope and Desmond W. Cooper teds) Mammals from Pouches and Eggs: Genetics, Breeding and Evolution of Marsupials and Monotremes CSIRO, 1990. A$50.00 in Australia (US$50.00 elsewhere) (xi + 337 pages) ISBN 0 643 05020 5
427