- Email: [email protected]
Comparative Primate Socioecology
BOOK REVIEWS
specifically the manufacture of handaxes, shows that imitation does not always lead to cultural evolution. The volume succeeds in the editors’ stated goal of extending the comparative database on mammalian social learning. In more than one instance a chapter appears to represent the first time that social learning has even been addressed in a species or group (e.g. Higginbottom & Croft’s chapter on marsupials). The editors believe that field studies are necessary in showing the likely functions or survival value of social learning, whereas controlled experiments are better suited to teasing out specific mechanisms. King argues in her chapter that identifying mechanisms of social learning is a nearimpossible task for most field researchers, and that a functional perspective based on the acquisition and donation of information is preferable. It follows that the many field studies summarized here stand as a useful testing ground for theoretical work that looks at the conditions under which social learning should be selected for (e.g. Laland et al. 1996). On the negative side, the book’s origin as a collection of conference papers is sometimes a little too obvious. The editors have prepared introductory comments for each section that go some way towards linking the different chapters. However, it would seem that many of the authors did not have a chance to read each other’s contributions: there are few cross-references between chapters. Furthermore, the inclusion of 21 chapters in a book of about 400 pages means that the average chapter is quite short. Perhaps as an inevitable result of this brevity, those authors who focused on their own work (as opposed to presenting a review) have tended to summarize previously published material rather than offering a novel or extended treatment. Phrases such as ‘much more research is needed in this area’ occur often in this collection. This is not a criticism per se; indeed, such phrases occur throughout the scientific literature. However, in an area like social learning, where we know so little about so many taxa, some strategic thinking about realistic future research is probably called for, and this is largely absent from the book. For example, given a lot of time, an unlimited budget and various technological advances in focal animal tracking and video recording, it is easy to conceive of experiments and fieldwork that would allow us to explore social learning in a difficult species such as the lion. But this is unlikely to happen. If our goals are to find out more about the functions and mechanisms of social learning in general, perhaps the most prudent course would be to focus on species where both field studies and naturalistic laboratory work are practicable: the chapters by Hudson et al., Laland and Faulkes on rabbits, rats and naked molerats, respectively, are inspiring in this regard. The book also exhibits differing perspectives on how much cognitive complexity we should ascribe to an animal that is shown to be capable of social learning. For example, Broom claims that the social learning seen in domestic animals can only be explained in terms of conscious awareness and cannot be reduced to ‘automatic responses’. In contrast, Hudson et al. (page 152) emphasize that isolating the mechanisms involved in social learning ‘can help counterbalance more mentalistic accounts’. Ultimately this is an empirical question, but it seems likely that in many cases investigators have overestimated the cognitive machinery required for one animal to learn from another (Noble & Todd, in press).
In summary, this book would be useful to anyone who needs to be reminded of just how little we know about social learning in most mammals. Readers seeking a more integrated treatment of theoretical and experimental work in the area of social learning might do well to start with Heyes & Galef (1996) before reading this volume. JASON NOBLE Center for Adaptive Behavior and Cognition, Max Planck Institute for Human Development, Lentzeallee 94, 14195 Berlin, Germany References Heyes, C. M. & Galef, B. G., Jr. (Eds) 1996. Social Learning in Animals: The Roots of Culture. San Diego: Academic Press. Laland, K. N., Richerson, P. J. & Boyd, R. 1996. Developing a theory of animal social learning. In: Social Learning in Animals: the Roots of Culture (Ed by C. M. Heyes & B. G. Galef, Jr), pp. 129–154. San Diego: Academic Press. Noble, J. & Todd, P. M. In press. Imitation or something simpler? Modelling simple mechanisms for social information processing. In: Imitation in Animals and Artifacts (Ed. by K. Dautenhahn & C. Nehaniv). Cambridge, Massachusetts: MIT Press.
doi:10.1006/anbe.1999.1324, available online at http://www.idealibrary.com on
Comparative Primate Socioecology. Edited by P. C. LEE. Cambridge: Cambridge University Press (1999). Pp. xii+412. Price $74.95. ‘What can be more curious,’ wrote Darwin, ‘than that the hand of man, formed for grasping, that of a mole for digging, the leg of a horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include similar bones in the same relative position?’ (1859a, page 334). Comparative morphology was a powerful pillar for formulating the concept of natural selection. Comparative socioecology adopts an evolutionary approach in seeking to identify how ecological variables shape social behaviour. Phyllis Lee has succeeded in producing a valuable book that demonstrates how difficult it is to answer this question. The diversity of primate social relationships, mating systems, and life history profiles cannot be pigeonholed into standard ecological models of social systems. The 15 chapters are organized into three sections (Comparative Methods, Comparative Life Histories, Comparative Socioecology) that probe six themes. MacLarnon opens the volume by outlining the rules, and problems, of comparative analyses. Her caveats provide an antidote to the assumption that adopting sophisticated quantitative comparative analyses in behaviour and ecology necessarily produce the most reasonable answer to a hypothesis. Contradictory conclusions can emerge when different variables are incorporated into testing a hypothesis, when different methods are used for comparative analysis, or when different statistical packages are used (e.g. Williamson & Dunbar’s chapter). Robson-Brown follows with a methodological chapter emphasizing the role of cladistics in comparative analysis, and concludes that sleep patterns in primates depend more upon phylogeny than ecology. Purvis & Webster complete the trio of methodological chapters with a concise description of the Comparative Analysis by Independent Contrasts
2000 The Association for the Study of Animal Behaviour
659
660
ANIMAL BEHAVIOUR, 59, 3
(http://evolve.zps.ox.ac.uk/CAIC/CAIC.html; Appendix 3.1). Criticisms of their approach are discussed at length, but ((their (contribution) ((suffers in the) (end when ((they (provide a)) (phylogeny of) (((extant Primates) that is)) virtually) (incomprehensible)) (due to (layers of (parentheses embedded))) in (parentheses within) (parentheses)))). All three methodology chapters reiterate that the fundamental problem with comparative analyses is that related species share traits derived from a common ancestor, so species within a genus are not ‘independent’ units for analysis. Application of life history models to primates is the basis of the second theme. Ross & Jones concur with many studies that show that, for their body size, primates mature slowly and reproduce at late ages. The authors suggest that female primates adopt a variety of behavioural strategies, possibly mediated by brain power, to circumvent physiological constraints on reproductive rate. Lee subsequently reasons that female alternative reproductive strategies provide mechanisms to maximize infant growth and survival while minimizing energetic and reproductive costs. She links her ‘weaning weight hypothesis’ and female strategies to maternal metabolic efficiency and risk level in the environment. Barton integrates brain evolution with primate socioecology by proposing that neocortical expansion of the parvocellular region was instrumental in primate evolution because it enhanced colour vision (which fostered incorporation of frugivory into the diet) and visual acuity (which fostered facial recognition and processing of social information). Two chapters explore sexual selection. Van Schaik et al. conclude that infanticide risk is the predominant force molding female sexuality, a conclusion opposite that of Dixson (1998). They reason that females seek a balance between paternal certainty, to elicit infant protection, and paternal uncertainty, to prevent infanticide, with enlarged sex skin swellings evolving because females ‘needed’ [sic] exaggerated signals to achieve this balance in some multimale mating systems. Their chapter is marred by a strident biological determinism. For example, they write (page 209) that ‘it [is] impossible for natural selection to design a male decision rule’ that enables males to detect ovulation. It is impossible for evolution to design an elephant that soars like an eagle, but no biological law precludes evolution of an ovulation detector. Natural selection has produced birds that swim and fish that ‘fly’. van Schaik et al.’s premise that ‘. . . the risk of male infanticide should have selected [my emphasis] for female counterstrategies’ (page 206) conflicts with Darwin’s inference (1859a, page 180) that ‘The belief that any given structure [e.g. female counterstrategies] which we think . . . would have been beneficial . . . would have been gained under all circumstances through natural selection, is opposed to what we can understand of its mechanism of action.’ If, in fact, natural selection ‘should’ produce female counterstrategies, then why is it ‘impossible’ for males to evolve countercounterstrategies? Plavcan’s chapter reveals that both canine and body size dimorphism are associated with levels of male–male competition and the operational sex ratio, but that the two measures of sexual dimorphism are not identical within a species. He suggests that multiple mechanisms contribute to the evolution of sexual dimorphism and points out that measuring size dimorphism is straightforward, but measuring sexual selection is not. Challenging current ecological models of primate social organization forms the next theme. Kappeler establishes a
number of differences between lemuroid and anthropoid societies; the most prominent is that multimale– multifemale lemur societies tend to have a uniform adult sex ratio with females frequently dominant to males. He concludes that lemur societies are either in a state of ‘evolutionary transition’ (page 291), or that adapting to the ecology of Madagascar might have involved incorporation of some unusual metabolic strategies. Strier assaults current socioecological models by focusing on the lack of cohesive bonds among female New World monkeys. She stresses that phylogeny and demography, as well as ecology, exert an impact on the structure of social relationships. Both authors emphasize that ecological models of ‘primate’ society are inapplicable to all primates. Foraging energetics forms the foundation for the fifth theme. Williams & Dunbar present a model accounting for the upper and lower limits of group size based upon a handful of ecological variables influencing time budgets. Their methods have important implications for conservation biology by establishing potential species distributions and ranges in group size based upon dietary and habitat requirements. Bean describes how food competition and body size dimorphism are related to daily energetic requirements and form a bidirectional pathway contributing to four different social systems found among the great apes. The final theme is human socioecology. Blurton-Jones et al. champion the ‘grandmother hypothesis’, which attributes the lengthy postreproductive life span of women to the value of females provisioning weaned, but semidependent, grandchildren. Foley speculates about the evolution of human behaviour based upon the fossil record, archaeological evidence, and environmental surroundings. He concludes that male kin bonding was of crucial importance in hominid evolution as a mechanism promoting survival in predator-rich environments or in intergroup antagonistic confrontations, a viewpoint closely echoing that of Alexander’s (1974) suggestion that hostile intergroup interactions formed the basis of human social evolution. Finally, Mace & Holden analyse about 80 sub-Saharan human cultures tracking the evolution of matrilineal inheritance patterns, but I do not understand why their chapter is included in this volume. Most of the contributions are intellectually stimulating, presenting new analyses as well as summarizing earlier research. But the editorial scrutiny is substandard. Words are inexplicably hyphenated (e.g. ‘difficult-ies’, ‘gran-dmothers’, ‘re-productive’), and, often, spelt incorrectly (e.g. ‘retention of give digits’, ‘agnostic competition’). The text contains a number of tables and appendices, but use of these is compromised by two oversights. First, in some cases, primates are listed in alphabetical order by genus and species, which hinders analysis by subfamily. Second, the taxonomy is inconsistent (e.g. Presbytis entellus (Table 4.2)=Semnopithecus entellus (Appendix 5.1); Lemur fulvus (Appendix 8.1)=Eulemur fulvus (Table 10.1)=Petterus fulvus (Appendix 3.1)). Primate societies did not evolve independently of ecological constraints, but it is absurd to think that the enormous flexibility and diversity of social systems can be mapped on to simple ecological factors. Furthermore, many solutions exist to solving the same problem: 100 pennies will buy the same item as 10 dimes or four quarters or one dollar bill. Wright (1969, 1977) was probably right: multiple adaptive peaks characterize the evolutionary landscape. Comparative Primate Socioecology will disappoint you if you want to know exactly how
BOOK REVIEWS
environmental variables have molded primate social behaviour and life history profiles, but it will excite you if you are challenged to learn about variation in social behaviour in nature, and if you are captivated and puzzled by Darwin’s (1859b) question to T. H. Huxley: ‘what the devil determines each particular variation?’ FRED B. BERCOVITCH Caribbean Primate Research Center, P.O. Box 1053, Sabana Seca, PR 00952, U.S.A. References Alexander, R. D. 1974. The evolution of social behavior. Annual Review of Ecology and Systematics, 5, 324–383. . Darwin, C. R. 1859a. On the Origin of Species. London: John Murray. Darwin, C. R. 1859b. Letter to T. H. Huxley [25 Nov 1859]. In: The Autobiography of Charles Darwin (Ed. by F. Darwin), pp. 226–227. New York: Dover. Dixson, A. F. 1998. Primate Sexuality. Oxford: Oxford University Press. Wright, S. 1969. Evolution and the Genetics of Populations. Vol. 2. The Theory of Gene Frequencies. Chicago: University of Chicago Press. Wright, S. 1977. Evolution and the Genetics of Populations. Vol. 3. Experimental Results and Evolutionary Deductions. Chicago: University of Chicago Press.
doi:10.1006/anbe.1999.1333, available online at http://www.idealibrary.com on
Ecobeaker 2.0 Laboratory Guide and Manual. By E. MEIR. Ithaca, New York: BeakerWare (1999). Pp. i+372. Price $85.00 single disk, $2000 site licence (25 copies). If the students at your institution are like those at mine, one of the most challenging aspects of your job is getting them to think critically and empirically. Ecobeaker is an excellent set of interactive computer simulation laboratories that can help you to achieve this goal. Ecobeaker is divided into two sections. The first section contains laboratory exercises, and the second contains a tutorial and manual to customize the program. Because I teach at an undergraduate institution, I was most interested in reviewing the prepared laboratories, although I also spent some time manipulating parameters such as graphs and sampling methods. I will first comment on the labora tories, then briefly discuss the tutorial and manual. Ecobeaker contains 21 laboratories. They are divided into what most would consider the fields of population ecology (oil spills and logistic bacteria; competitive exclusion principle; R* competition, succession and death; varieties of competition and biological pest control), community ecology (sick fish, intermediate disturbance hypothesis, keystone predation, barnacles and tides, aquatic trophic cascades, sewage, predator avoidance in aquatic systems, island biogeography, corridors, stepping stones and butterflies), evolution (islands and natural selection, the heterozygous advantage) and sampling (quadrat sampling, sampling under pressure, mark and recapture, breeding bird survey sampling design and varieties of spatial models). Each laboratory contains a background, an outline of the laboratory, the laboratory itself, notes and comments, and references. The background section contains a brief introduction to the ‘big picture’ of the exercise. I found these sections easy and fun to read,
and I imagine that students will enjoy them too since they are written at a general level and describe real-life applications to ecological theory. The outline sets the stage for the computer simulation and, again, Meir does an excellent job of excluding jargon and explaining things at a general level. The laboratory section contains the instructions for executing the actual laboratory. The directions are clear and straightforward, and a student can have any of the laboratories up and running in minutes. The final section (notes and comments) contains an example of how the laboratory relates to a real-life situation. Some laboratories also have a section called ‘More Things to Try’, where students are encouraged to change the simulation by adding more or different guilds of species or by changing the rate of change or distribution of individuals. The reference section usually contains one or two sources for further information. The thing that I like most about Ecobeaker is the congruity of organization, not only in the arrangement of the text but also on the computer screen. The computer screen setup is consistent across all of the laboratories: multiple windows show a grid in which students can see the simulation run; a graph that displays the numeric results; a control panel and a window that permits changing parameters. Students can run the program one simulation at a time, or have the steps occur quickly in order to observe trends over time. Computer simulations can be powerful teaching tools, and I have used a variety of them to help students visualize what happens when variables in mathematical models are changed. This is the first software package I have come across that can model many species at a time while also allowing students to modify the behaviour of individual species. Although I am overwhelmingly positive about these computer simulations, I warn faculty that they will probably have to spend some time developing supplemental materials for a laboratory. For example, the Corridors, Stepping Stones and Butterfly laboratory addresses a problem faced by conservation biologists: is it better to add habitat to make a bigger ‘patch’ or to create a corridor between small patches? The goal of the laboratory is to outline a restoration plan that maximizes butterfly numbers. Meir estimates that this laboratory will take 4–6 h, but as written, I cannot imagine students taking the time to change systematically the location of the area to figure out which configuration is best, instead of running one or two simulations and calling it quits. To use this laboratory effectively I would develop a handout to guide students along; for example, what happens if 50% of the new area is added to patch 1? What happens if 100% of the area is stretched between patch 1 and patch 2? What happens to bufferfly numbers when both patch 1 and patch 2 are increased in size? As in the laboratory section, the tutorial and manual section is very well written. Ecobeaker is currently available only in Macintosh format (the author expects the Windows version to be available in February 2000). Although I am not an accomplished Mac user, Meir’s directions are so complete that I easily saved runs, changed graphs, varied my sampling method, marked individuals to follow over time, changed habitat types in the butterfly laboratory and had fun manipulating the grid on which the simulation ran. Ecobaker’s online site (http://www.ecobeaker.com) contains a few more laboratories, as well as information on ordering and creating a customized laboratory guide. If the Windows version is as good as that of the Mac, you
2000 The Association for the Study of Animal Behaviour
661
specifically the manufacture of handaxes, shows that imitation does not always lead to cultural evolution. The volume succeeds in the editors’ stated goal of extending the comparative database on mammalian social learning. In more than one instance a chapter appears to represent the first time that social learning has even been addressed in a species or group (e.g. Higginbottom & Croft’s chapter on marsupials). The editors believe that field studies are necessary in showing the likely functions or survival value of social learning, whereas controlled experiments are better suited to teasing out specific mechanisms. King argues in her chapter that identifying mechanisms of social learning is a nearimpossible task for most field researchers, and that a functional perspective based on the acquisition and donation of information is preferable. It follows that the many field studies summarized here stand as a useful testing ground for theoretical work that looks at the conditions under which social learning should be selected for (e.g. Laland et al. 1996). On the negative side, the book’s origin as a collection of conference papers is sometimes a little too obvious. The editors have prepared introductory comments for each section that go some way towards linking the different chapters. However, it would seem that many of the authors did not have a chance to read each other’s contributions: there are few cross-references between chapters. Furthermore, the inclusion of 21 chapters in a book of about 400 pages means that the average chapter is quite short. Perhaps as an inevitable result of this brevity, those authors who focused on their own work (as opposed to presenting a review) have tended to summarize previously published material rather than offering a novel or extended treatment. Phrases such as ‘much more research is needed in this area’ occur often in this collection. This is not a criticism per se; indeed, such phrases occur throughout the scientific literature. However, in an area like social learning, where we know so little about so many taxa, some strategic thinking about realistic future research is probably called for, and this is largely absent from the book. For example, given a lot of time, an unlimited budget and various technological advances in focal animal tracking and video recording, it is easy to conceive of experiments and fieldwork that would allow us to explore social learning in a difficult species such as the lion. But this is unlikely to happen. If our goals are to find out more about the functions and mechanisms of social learning in general, perhaps the most prudent course would be to focus on species where both field studies and naturalistic laboratory work are practicable: the chapters by Hudson et al., Laland and Faulkes on rabbits, rats and naked molerats, respectively, are inspiring in this regard. The book also exhibits differing perspectives on how much cognitive complexity we should ascribe to an animal that is shown to be capable of social learning. For example, Broom claims that the social learning seen in domestic animals can only be explained in terms of conscious awareness and cannot be reduced to ‘automatic responses’. In contrast, Hudson et al. (page 152) emphasize that isolating the mechanisms involved in social learning ‘can help counterbalance more mentalistic accounts’. Ultimately this is an empirical question, but it seems likely that in many cases investigators have overestimated the cognitive machinery required for one animal to learn from another (Noble & Todd, in press).
In summary, this book would be useful to anyone who needs to be reminded of just how little we know about social learning in most mammals. Readers seeking a more integrated treatment of theoretical and experimental work in the area of social learning might do well to start with Heyes & Galef (1996) before reading this volume. JASON NOBLE Center for Adaptive Behavior and Cognition, Max Planck Institute for Human Development, Lentzeallee 94, 14195 Berlin, Germany References Heyes, C. M. & Galef, B. G., Jr. (Eds) 1996. Social Learning in Animals: The Roots of Culture. San Diego: Academic Press. Laland, K. N., Richerson, P. J. & Boyd, R. 1996. Developing a theory of animal social learning. In: Social Learning in Animals: the Roots of Culture (Ed by C. M. Heyes & B. G. Galef, Jr), pp. 129–154. San Diego: Academic Press. Noble, J. & Todd, P. M. In press. Imitation or something simpler? Modelling simple mechanisms for social information processing. In: Imitation in Animals and Artifacts (Ed. by K. Dautenhahn & C. Nehaniv). Cambridge, Massachusetts: MIT Press.
doi:10.1006/anbe.1999.1324, available online at http://www.idealibrary.com on
Comparative Primate Socioecology. Edited by P. C. LEE. Cambridge: Cambridge University Press (1999). Pp. xii+412. Price $74.95. ‘What can be more curious,’ wrote Darwin, ‘than that the hand of man, formed for grasping, that of a mole for digging, the leg of a horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include similar bones in the same relative position?’ (1859a, page 334). Comparative morphology was a powerful pillar for formulating the concept of natural selection. Comparative socioecology adopts an evolutionary approach in seeking to identify how ecological variables shape social behaviour. Phyllis Lee has succeeded in producing a valuable book that demonstrates how difficult it is to answer this question. The diversity of primate social relationships, mating systems, and life history profiles cannot be pigeonholed into standard ecological models of social systems. The 15 chapters are organized into three sections (Comparative Methods, Comparative Life Histories, Comparative Socioecology) that probe six themes. MacLarnon opens the volume by outlining the rules, and problems, of comparative analyses. Her caveats provide an antidote to the assumption that adopting sophisticated quantitative comparative analyses in behaviour and ecology necessarily produce the most reasonable answer to a hypothesis. Contradictory conclusions can emerge when different variables are incorporated into testing a hypothesis, when different methods are used for comparative analysis, or when different statistical packages are used (e.g. Williamson & Dunbar’s chapter). Robson-Brown follows with a methodological chapter emphasizing the role of cladistics in comparative analysis, and concludes that sleep patterns in primates depend more upon phylogeny than ecology. Purvis & Webster complete the trio of methodological chapters with a concise description of the Comparative Analysis by Independent Contrasts
2000 The Association for the Study of Animal Behaviour
659
660
ANIMAL BEHAVIOUR, 59, 3
(http://evolve.zps.ox.ac.uk/CAIC/CAIC.html; Appendix 3.1). Criticisms of their approach are discussed at length, but ((their (contribution) ((suffers in the) (end when ((they (provide a)) (phylogeny of) (((extant Primates) that is)) virtually) (incomprehensible)) (due to (layers of (parentheses embedded))) in (parentheses within) (parentheses)))). All three methodology chapters reiterate that the fundamental problem with comparative analyses is that related species share traits derived from a common ancestor, so species within a genus are not ‘independent’ units for analysis. Application of life history models to primates is the basis of the second theme. Ross & Jones concur with many studies that show that, for their body size, primates mature slowly and reproduce at late ages. The authors suggest that female primates adopt a variety of behavioural strategies, possibly mediated by brain power, to circumvent physiological constraints on reproductive rate. Lee subsequently reasons that female alternative reproductive strategies provide mechanisms to maximize infant growth and survival while minimizing energetic and reproductive costs. She links her ‘weaning weight hypothesis’ and female strategies to maternal metabolic efficiency and risk level in the environment. Barton integrates brain evolution with primate socioecology by proposing that neocortical expansion of the parvocellular region was instrumental in primate evolution because it enhanced colour vision (which fostered incorporation of frugivory into the diet) and visual acuity (which fostered facial recognition and processing of social information). Two chapters explore sexual selection. Van Schaik et al. conclude that infanticide risk is the predominant force molding female sexuality, a conclusion opposite that of Dixson (1998). They reason that females seek a balance between paternal certainty, to elicit infant protection, and paternal uncertainty, to prevent infanticide, with enlarged sex skin swellings evolving because females ‘needed’ [sic] exaggerated signals to achieve this balance in some multimale mating systems. Their chapter is marred by a strident biological determinism. For example, they write (page 209) that ‘it [is] impossible for natural selection to design a male decision rule’ that enables males to detect ovulation. It is impossible for evolution to design an elephant that soars like an eagle, but no biological law precludes evolution of an ovulation detector. Natural selection has produced birds that swim and fish that ‘fly’. van Schaik et al.’s premise that ‘. . . the risk of male infanticide should have selected [my emphasis] for female counterstrategies’ (page 206) conflicts with Darwin’s inference (1859a, page 180) that ‘The belief that any given structure [e.g. female counterstrategies] which we think . . . would have been beneficial . . . would have been gained under all circumstances through natural selection, is opposed to what we can understand of its mechanism of action.’ If, in fact, natural selection ‘should’ produce female counterstrategies, then why is it ‘impossible’ for males to evolve countercounterstrategies? Plavcan’s chapter reveals that both canine and body size dimorphism are associated with levels of male–male competition and the operational sex ratio, but that the two measures of sexual dimorphism are not identical within a species. He suggests that multiple mechanisms contribute to the evolution of sexual dimorphism and points out that measuring size dimorphism is straightforward, but measuring sexual selection is not. Challenging current ecological models of primate social organization forms the next theme. Kappeler establishes a
number of differences between lemuroid and anthropoid societies; the most prominent is that multimale– multifemale lemur societies tend to have a uniform adult sex ratio with females frequently dominant to males. He concludes that lemur societies are either in a state of ‘evolutionary transition’ (page 291), or that adapting to the ecology of Madagascar might have involved incorporation of some unusual metabolic strategies. Strier assaults current socioecological models by focusing on the lack of cohesive bonds among female New World monkeys. She stresses that phylogeny and demography, as well as ecology, exert an impact on the structure of social relationships. Both authors emphasize that ecological models of ‘primate’ society are inapplicable to all primates. Foraging energetics forms the foundation for the fifth theme. Williams & Dunbar present a model accounting for the upper and lower limits of group size based upon a handful of ecological variables influencing time budgets. Their methods have important implications for conservation biology by establishing potential species distributions and ranges in group size based upon dietary and habitat requirements. Bean describes how food competition and body size dimorphism are related to daily energetic requirements and form a bidirectional pathway contributing to four different social systems found among the great apes. The final theme is human socioecology. Blurton-Jones et al. champion the ‘grandmother hypothesis’, which attributes the lengthy postreproductive life span of women to the value of females provisioning weaned, but semidependent, grandchildren. Foley speculates about the evolution of human behaviour based upon the fossil record, archaeological evidence, and environmental surroundings. He concludes that male kin bonding was of crucial importance in hominid evolution as a mechanism promoting survival in predator-rich environments or in intergroup antagonistic confrontations, a viewpoint closely echoing that of Alexander’s (1974) suggestion that hostile intergroup interactions formed the basis of human social evolution. Finally, Mace & Holden analyse about 80 sub-Saharan human cultures tracking the evolution of matrilineal inheritance patterns, but I do not understand why their chapter is included in this volume. Most of the contributions are intellectually stimulating, presenting new analyses as well as summarizing earlier research. But the editorial scrutiny is substandard. Words are inexplicably hyphenated (e.g. ‘difficult-ies’, ‘gran-dmothers’, ‘re-productive’), and, often, spelt incorrectly (e.g. ‘retention of give digits’, ‘agnostic competition’). The text contains a number of tables and appendices, but use of these is compromised by two oversights. First, in some cases, primates are listed in alphabetical order by genus and species, which hinders analysis by subfamily. Second, the taxonomy is inconsistent (e.g. Presbytis entellus (Table 4.2)=Semnopithecus entellus (Appendix 5.1); Lemur fulvus (Appendix 8.1)=Eulemur fulvus (Table 10.1)=Petterus fulvus (Appendix 3.1)). Primate societies did not evolve independently of ecological constraints, but it is absurd to think that the enormous flexibility and diversity of social systems can be mapped on to simple ecological factors. Furthermore, many solutions exist to solving the same problem: 100 pennies will buy the same item as 10 dimes or four quarters or one dollar bill. Wright (1969, 1977) was probably right: multiple adaptive peaks characterize the evolutionary landscape. Comparative Primate Socioecology will disappoint you if you want to know exactly how
BOOK REVIEWS
environmental variables have molded primate social behaviour and life history profiles, but it will excite you if you are challenged to learn about variation in social behaviour in nature, and if you are captivated and puzzled by Darwin’s (1859b) question to T. H. Huxley: ‘what the devil determines each particular variation?’ FRED B. BERCOVITCH Caribbean Primate Research Center, P.O. Box 1053, Sabana Seca, PR 00952, U.S.A. References Alexander, R. D. 1974. The evolution of social behavior. Annual Review of Ecology and Systematics, 5, 324–383. . Darwin, C. R. 1859a. On the Origin of Species. London: John Murray. Darwin, C. R. 1859b. Letter to T. H. Huxley [25 Nov 1859]. In: The Autobiography of Charles Darwin (Ed. by F. Darwin), pp. 226–227. New York: Dover. Dixson, A. F. 1998. Primate Sexuality. Oxford: Oxford University Press. Wright, S. 1969. Evolution and the Genetics of Populations. Vol. 2. The Theory of Gene Frequencies. Chicago: University of Chicago Press. Wright, S. 1977. Evolution and the Genetics of Populations. Vol. 3. Experimental Results and Evolutionary Deductions. Chicago: University of Chicago Press.
doi:10.1006/anbe.1999.1333, available online at http://www.idealibrary.com on
Ecobeaker 2.0 Laboratory Guide and Manual. By E. MEIR. Ithaca, New York: BeakerWare (1999). Pp. i+372. Price $85.00 single disk, $2000 site licence (25 copies). If the students at your institution are like those at mine, one of the most challenging aspects of your job is getting them to think critically and empirically. Ecobeaker is an excellent set of interactive computer simulation laboratories that can help you to achieve this goal. Ecobeaker is divided into two sections. The first section contains laboratory exercises, and the second contains a tutorial and manual to customize the program. Because I teach at an undergraduate institution, I was most interested in reviewing the prepared laboratories, although I also spent some time manipulating parameters such as graphs and sampling methods. I will first comment on the labora tories, then briefly discuss the tutorial and manual. Ecobeaker contains 21 laboratories. They are divided into what most would consider the fields of population ecology (oil spills and logistic bacteria; competitive exclusion principle; R* competition, succession and death; varieties of competition and biological pest control), community ecology (sick fish, intermediate disturbance hypothesis, keystone predation, barnacles and tides, aquatic trophic cascades, sewage, predator avoidance in aquatic systems, island biogeography, corridors, stepping stones and butterflies), evolution (islands and natural selection, the heterozygous advantage) and sampling (quadrat sampling, sampling under pressure, mark and recapture, breeding bird survey sampling design and varieties of spatial models). Each laboratory contains a background, an outline of the laboratory, the laboratory itself, notes and comments, and references. The background section contains a brief introduction to the ‘big picture’ of the exercise. I found these sections easy and fun to read,
and I imagine that students will enjoy them too since they are written at a general level and describe real-life applications to ecological theory. The outline sets the stage for the computer simulation and, again, Meir does an excellent job of excluding jargon and explaining things at a general level. The laboratory section contains the instructions for executing the actual laboratory. The directions are clear and straightforward, and a student can have any of the laboratories up and running in minutes. The final section (notes and comments) contains an example of how the laboratory relates to a real-life situation. Some laboratories also have a section called ‘More Things to Try’, where students are encouraged to change the simulation by adding more or different guilds of species or by changing the rate of change or distribution of individuals. The reference section usually contains one or two sources for further information. The thing that I like most about Ecobeaker is the congruity of organization, not only in the arrangement of the text but also on the computer screen. The computer screen setup is consistent across all of the laboratories: multiple windows show a grid in which students can see the simulation run; a graph that displays the numeric results; a control panel and a window that permits changing parameters. Students can run the program one simulation at a time, or have the steps occur quickly in order to observe trends over time. Computer simulations can be powerful teaching tools, and I have used a variety of them to help students visualize what happens when variables in mathematical models are changed. This is the first software package I have come across that can model many species at a time while also allowing students to modify the behaviour of individual species. Although I am overwhelmingly positive about these computer simulations, I warn faculty that they will probably have to spend some time developing supplemental materials for a laboratory. For example, the Corridors, Stepping Stones and Butterfly laboratory addresses a problem faced by conservation biologists: is it better to add habitat to make a bigger ‘patch’ or to create a corridor between small patches? The goal of the laboratory is to outline a restoration plan that maximizes butterfly numbers. Meir estimates that this laboratory will take 4–6 h, but as written, I cannot imagine students taking the time to change systematically the location of the area to figure out which configuration is best, instead of running one or two simulations and calling it quits. To use this laboratory effectively I would develop a handout to guide students along; for example, what happens if 50% of the new area is added to patch 1? What happens if 100% of the area is stretched between patch 1 and patch 2? What happens to bufferfly numbers when both patch 1 and patch 2 are increased in size? As in the laboratory section, the tutorial and manual section is very well written. Ecobeaker is currently available only in Macintosh format (the author expects the Windows version to be available in February 2000). Although I am not an accomplished Mac user, Meir’s directions are so complete that I easily saved runs, changed graphs, varied my sampling method, marked individuals to follow over time, changed habitat types in the butterfly laboratory and had fun manipulating the grid on which the simulation ran. Ecobaker’s online site (http://www.ecobeaker.com) contains a few more laboratories, as well as information on ordering and creating a customized laboratory guide. If the Windows version is as good as that of the Mac, you
2000 The Association for the Study of Animal Behaviour
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