- Email: [email protected]
Evolutionary paleobiology of behavior and coevolution
TREE vol. 6, no. 7, January
7997
For instance, the role of disturbance in modifying the outcome of competition is not given adequate attention, the competitive dynamics of species utilizing patchy and ephemeral resources are ignored, and I looked in vain for anything on indirect competition. I expected references to papers such as those by Huston2, Atkinson and Shorrocks3, HoIt and Jeffries and Lawton5, but was disappointed. The little that is said about population cycles in small mammals could have been written in 1980. In view of the eloquent case made in the introduction for enlightened stewardship of the natural world, Ricklefs’ book is strangely lacking in applied aspects of ecology. There is very little on the management or harvesting of natural populations; the only reference I could find to acid rain was a mention of its ability to erode marble statuary, and the greenhouse effect features only in the introduction. References to biological control seem to be restricted to the OpuntiaCactoblastis story in the chapter on predation, and the replacement series of Aphytis species under the heading of competition. Otherwise, the role of biological control in pest management is largely overlooked and the modern concept of integrated pest management ignored. There is also an unevenness in the depth of treatment of topics. BHT appeared first in 1986, so the publication of a second edition in 1990 may well have set a speed record in the ecological literature. It maintains a narrower view of the scope of ecology than does Ricklefs, adopting the Krebs6 definition. The result is a book that makes only brief forays into evolutionary and behavioural territory; coevolution, for instance, merits a full chapter in Ricklefs, but here only a few rather non-committal paragraphs. The new edition contains two additional chapters. One deals at length with human management of populations, and the other with the flux of matter through ecosystems. Other changes include improvements to diagrams and, curiously, a complete change of subject matter in the four pages of colour plates. Previously, these featured an eclectic set of images from rosette treelets to aposematic butterflies. Now we are offered snapshots of a selection of ecosystems. I doubt whether the didactic value of any of these warrants the cost of their inclusion. The new chapters are comprehensive, well written and welcome. Like the rest of the book, they are thoroughly up-to-date. The risk is that the authors may be obliged to con-
tinue revising frequently, as the fortunes of new ideas wax or wane. I was a little surprised to find that supplyside influences on community composition7 have not been included, but otherwise the volume contains all that it should. The features of the previous edition have been retained, most notably the important distinction between the population dynamics of unitary and modular organisms, but also the take-home messages in the margins of the text. Two other minor, but very useful, attributes are the provision of separate indexes for organisms and subjects and the inclusion of section numbers after each bibliographic reference. I wish thatthese were standard in ecology texts. So, two books that appeared at first sight to be similar turn out to be very different. No doubt Ricklefs will continue to appeal to those who require coverage of aspects of behavioural ecology and evolution, though it faces formidable competition from specialist texts in these areas. I shall continue to recommend BHT to ad-
vanced undergraduates and postgraduates in ecology. Unlike nouvelle cuisine, it has proved to be more than a short-lived novelty; it is also a great deal more sustaining intellectually than the former is nutritionally. Small quibbles apart, it is a row de force that has yet to be surpassed.
John H.R. Gee Deptof BiologicalSciences, The University College of Wales, Aberystwyth, Dyfed SY23 3DA, UK
References 1 Smith, R.H. (1986)
Nature 319,809 2 Huston. M. (1979) Am. Nat. 113. 81-101 3 Atkinson, W.D. and Shorrocks, 6. Ecol. 50,461-471 (1981) J. Aim. 4 Holt, R.D. (1977) Theor. Pop/. Biol. 12, 197-229 5 Jeffries, M.J. and Lawton, J.H. (1984) Biol. J. Linn. Sot. 23, 269-286 6 Krebs, C.J. (1972) Ecology, Harper & Row 7 Roughgarden, J., Gaines, SD. and Pacala. S.W. 11987) in Oraanization of Comnktitiek Pas; and present (Gee,
J.H.R. and Giller, P.S., eds), pp. 491-518, Blackwell Scientific Publications
Paleo biology: Beyond Morphology Evolutionary Paleobiology of Behavior and Coevolution by A.J. Boucot, Elsevier, $748.75/Dfl 290.00 hbk (xxiii pages) ISBN 0 444 88034 8
7990. + 724
The fossil record yields its secrets grudgingly. It tells us most readily about hard tissue morphology, but information on behavior, ecological interactions and other biological properties is given more reluctantly and with disturbing vagueness. Then why devote an entire book to the paleobiology of behavior and ecological interactions? Boucot’s answer, initially, is that he wants to show that behavior and coevolutionary interactions between species evolve rapidly, but once established are stable for geologically long intervals. In the concluding chapter, he reveals his more ambitious objective of synthesizing his views on community evolution, coevolution, and behavioral and morphological evolution into a unified model for the tempo of evolutionary change. The middle 90% of the book is a fascinating review of evidence on behavior and ecological interactions from the fossil record. Boucot argues that ecological communities episodically undergo rad-
ical reorganizations within geological instants and then experience prolonged intervals of stability. In his view, community reorganization disrupts coevolutionary relationships and stimulates quantum evolution (i.e. evolution of genus- and familylevel characters) of morphological and behavioral innovations. These innovations rapidly adapt organisms to the new community. As long as community structure is stable, however, only trivial (i.e. species-level) phenotypic evolution occurs. This is an intriguing view of macroevolution and its relation to ecological change. Unfortunately, Boucot fails to substantiate his claim that behavioral evolution and coevolution are strongly episodic. Although numerous behavioral and ecological phenomena from the fossil record are presented, they are rarely accompanied by convincing information on their evolutionary tempo. Boucot fails to show that stratigraphic sequences of fossil taxa with a behavioral or ecological characteristic represent a single lineage rather than a set of phenotypically similar lineages. Also, he rarely tracks the appearance of new behaviors and ecological interactions through geological time, nor does he even show from their stratigraphic distribution 35
TREE Vol. 6,
that such characters must have evolved within a short interval. Finally, claims for stability (stasis) of behaviors and ecological interactions are based on observation at too few (usually two) points in geological time. These methodological deficiencies are glaring because the problems of recognizing lineages1-3 and of characterizing rates4 and patterns5J of morphological evolution in fossils have been debated extensively. Such questions are serious and as pertinent to behavioral and ecological phenotypes as they are to morphology. Because of these deficiencies, Boucot cannot claim to have demonstrated that behavioral evolution and coevolution are episodic, and thus their tempos cannot be used as support for his model of evolutionary tempo. Equally disappointing is Boucot’s failure to make adequate use of the literature. For example, omission of Vermeij’s’ recent book, which also argues for control of phenotypic innovation by community structure, is unfortunate. Ehrlich and Raven’s8 argument that phenetic cohesion of species is imposed by community structure rather than by gene flow could have been used by Boucot to support his thesis. Van Valen’sg Red Queen hypothesis holds that change in the biotic environment exerts con-
tinuous pressure for evolution rather than causing the episodic crises envisioned by Boucot. Conflicts and congruencies between the author’s views and those presented by others are inadequately explored. Similarly, Boucot sometimes overlooks recent contributions, citing, for example Romer’s’O classic but aging Vertebrate Paleontology instead of Carroll’s” recent comprehensive review of vertebrate paleontology. On the positive side, the volume is packed with interesting and wellorganized examples of behavioral and ecological properties of fossil organisms. Collection of this information in one place is the primary value of the book. Echoing recent suggestions by others’*, Boucot criticizes studies that estimate numbers of taxa and their rates of extinction and origin through geological time without taking into account membership of these taxa in different ecological communities. Aggregation of such data among communities would obscure the episodic nature of evolution and extinction if the episodes are, as Boucot claims, asynchronous among communities. Although Boucot has discussed some interesting issues and gathered a wealth of information on behavior and ecology of fossil organisms, he has not built the empirical edifice he had hoped would support his argu-
~0,
ment for the control tempo by community
7,
J'anuary : ,“!~ ?
of evolutionary structure. Michael A. Bell
Deptof Ecologyand Evolution,StateUnivewry ai New York, Stony Brook,NY 11794.5245,USA
References 1 Schaeffer, B., Hecht, M.K. and Eldredge, N. (1972) Evol. Biol. 6, 31-46 2 Gingerich, P.D. (1979) in Phylogeneric Analysis and Paleontology (Cracraft, J. and Eldredge, N., eds), pp. 41-79, Columbia University Press 3 Bell, M.A. (1988) Trends Ecol. Evol. 3, 320-325 4 Gingerich, P.D. (1983) Science 222, 159-161 5 Bookstein, F.L., Gingerich, P.D. and Kluge, A.G. (1978) Paleobiology4, 120-l 34 6 Bell, M.A. and Legendre, P. (1987) Syst. Zool. 36. 52-61 7 Vermeij, G. (1987) Evolution and Escalation, Princeton University Press 8 Ehrlich, P.R. and Raven, P.H. (1969) Science 165,1228-l 232 9 Van Valen, L. (1973) Evol. Theory 1, l-18 10 Romer, A.S. (1966) Vertebrate Paleontology, University of Chicago Press 11 Carroll, R.L. (1987) Vertebrate Paleontology and Evolution, W.H. Freeman 12 Behrensmeyer, A.K. (1988) Paleogeogr. Paleoclimatol. faleoecol. 63, 183-199
Financed by research ecologists and non-profit-making, the in2ernationel Ecology kWitute (ECI) has a staff of 41 ecologists - marine, terrestrial and limnetic - oil of high pro&s&& rsput&on. N&her the direetw nor any member of the scientific staff receives remuneration. Every year a jury campowd of ECt members s&eta - in a rot&#@ wrn - prize winners among marine, terrestrial or iimnetic ecologists. In 1991, prize winner will be selected in the fictldof limn&k esalogy. The winner of ths Ecology Institute Prize is requested to author a 2011to 300 print&pqe book, to be published by ECI in the series ‘Excellence in Ecology’(EE) and to be made available worldwideat cost price. Ei?s cct~nw@ t8 different from that oftextbooks. In addition to reviewing a cert&~ f&id rif knowledge, itgives the authors a chme to Msatheir pe~~rtelviews on importirnte&ogical issues, to interp~etrrent ~ti~~~owttsdgeon the basis oftheir own mqerience and in&#, an&o tell us what, in their opinion, should b&done in the future. The Ecology Institute Prize is endow&d with a stipend of US$&XlO. Asecond prize may*ba awarded honoring a young ecologist who has conduct@ and pubKsh& res#s of uniqurprtyind research efforts representing an important scientific brMtt&rqjh: the IlppEPR!ZEftritematio ix) is endowed with a stipend of USQSO. Nomktaticms we m alt resslerreh ecol@ts. iersonof tb&!I Jury (see beiow) before July 31, 1991. E#ig&Ieare ait gtaggylin scientific rese I’rtd~~~ranc)t9lt,~ry’schairpereon: Jury mernbersnominated SK&M& The Jury w&El 1x~‘\MFnner&sN t&~nominatibns received, as well as their o&n will be replaced by knowladgrr of top pwb~fnefs, and their own professton& @@m%nt. C&to WWWJ (Director EC.0 Ecology Enstitute,Nordbilnte 23, D-2124 Oldendorf/Luhe, FRG. Tel: 04132i727. Fax: Q413MB83 ECI Limnetic Jury 1991: i, S&oPaula, BF&I; 0. Uhimann, Dresden,
7997
For instance, the role of disturbance in modifying the outcome of competition is not given adequate attention, the competitive dynamics of species utilizing patchy and ephemeral resources are ignored, and I looked in vain for anything on indirect competition. I expected references to papers such as those by Huston2, Atkinson and Shorrocks3, HoIt and Jeffries and Lawton5, but was disappointed. The little that is said about population cycles in small mammals could have been written in 1980. In view of the eloquent case made in the introduction for enlightened stewardship of the natural world, Ricklefs’ book is strangely lacking in applied aspects of ecology. There is very little on the management or harvesting of natural populations; the only reference I could find to acid rain was a mention of its ability to erode marble statuary, and the greenhouse effect features only in the introduction. References to biological control seem to be restricted to the OpuntiaCactoblastis story in the chapter on predation, and the replacement series of Aphytis species under the heading of competition. Otherwise, the role of biological control in pest management is largely overlooked and the modern concept of integrated pest management ignored. There is also an unevenness in the depth of treatment of topics. BHT appeared first in 1986, so the publication of a second edition in 1990 may well have set a speed record in the ecological literature. It maintains a narrower view of the scope of ecology than does Ricklefs, adopting the Krebs6 definition. The result is a book that makes only brief forays into evolutionary and behavioural territory; coevolution, for instance, merits a full chapter in Ricklefs, but here only a few rather non-committal paragraphs. The new edition contains two additional chapters. One deals at length with human management of populations, and the other with the flux of matter through ecosystems. Other changes include improvements to diagrams and, curiously, a complete change of subject matter in the four pages of colour plates. Previously, these featured an eclectic set of images from rosette treelets to aposematic butterflies. Now we are offered snapshots of a selection of ecosystems. I doubt whether the didactic value of any of these warrants the cost of their inclusion. The new chapters are comprehensive, well written and welcome. Like the rest of the book, they are thoroughly up-to-date. The risk is that the authors may be obliged to con-
tinue revising frequently, as the fortunes of new ideas wax or wane. I was a little surprised to find that supplyside influences on community composition7 have not been included, but otherwise the volume contains all that it should. The features of the previous edition have been retained, most notably the important distinction between the population dynamics of unitary and modular organisms, but also the take-home messages in the margins of the text. Two other minor, but very useful, attributes are the provision of separate indexes for organisms and subjects and the inclusion of section numbers after each bibliographic reference. I wish thatthese were standard in ecology texts. So, two books that appeared at first sight to be similar turn out to be very different. No doubt Ricklefs will continue to appeal to those who require coverage of aspects of behavioural ecology and evolution, though it faces formidable competition from specialist texts in these areas. I shall continue to recommend BHT to ad-
vanced undergraduates and postgraduates in ecology. Unlike nouvelle cuisine, it has proved to be more than a short-lived novelty; it is also a great deal more sustaining intellectually than the former is nutritionally. Small quibbles apart, it is a row de force that has yet to be surpassed.
John H.R. Gee Deptof BiologicalSciences, The University College of Wales, Aberystwyth, Dyfed SY23 3DA, UK
References 1 Smith, R.H. (1986)
Nature 319,809 2 Huston. M. (1979) Am. Nat. 113. 81-101 3 Atkinson, W.D. and Shorrocks, 6. Ecol. 50,461-471 (1981) J. Aim. 4 Holt, R.D. (1977) Theor. Pop/. Biol. 12, 197-229 5 Jeffries, M.J. and Lawton, J.H. (1984) Biol. J. Linn. Sot. 23, 269-286 6 Krebs, C.J. (1972) Ecology, Harper & Row 7 Roughgarden, J., Gaines, SD. and Pacala. S.W. 11987) in Oraanization of Comnktitiek Pas; and present (Gee,
J.H.R. and Giller, P.S., eds), pp. 491-518, Blackwell Scientific Publications
Paleo biology: Beyond Morphology Evolutionary Paleobiology of Behavior and Coevolution by A.J. Boucot, Elsevier, $748.75/Dfl 290.00 hbk (xxiii pages) ISBN 0 444 88034 8
7990. + 724
The fossil record yields its secrets grudgingly. It tells us most readily about hard tissue morphology, but information on behavior, ecological interactions and other biological properties is given more reluctantly and with disturbing vagueness. Then why devote an entire book to the paleobiology of behavior and ecological interactions? Boucot’s answer, initially, is that he wants to show that behavior and coevolutionary interactions between species evolve rapidly, but once established are stable for geologically long intervals. In the concluding chapter, he reveals his more ambitious objective of synthesizing his views on community evolution, coevolution, and behavioral and morphological evolution into a unified model for the tempo of evolutionary change. The middle 90% of the book is a fascinating review of evidence on behavior and ecological interactions from the fossil record. Boucot argues that ecological communities episodically undergo rad-
ical reorganizations within geological instants and then experience prolonged intervals of stability. In his view, community reorganization disrupts coevolutionary relationships and stimulates quantum evolution (i.e. evolution of genus- and familylevel characters) of morphological and behavioral innovations. These innovations rapidly adapt organisms to the new community. As long as community structure is stable, however, only trivial (i.e. species-level) phenotypic evolution occurs. This is an intriguing view of macroevolution and its relation to ecological change. Unfortunately, Boucot fails to substantiate his claim that behavioral evolution and coevolution are strongly episodic. Although numerous behavioral and ecological phenomena from the fossil record are presented, they are rarely accompanied by convincing information on their evolutionary tempo. Boucot fails to show that stratigraphic sequences of fossil taxa with a behavioral or ecological characteristic represent a single lineage rather than a set of phenotypically similar lineages. Also, he rarely tracks the appearance of new behaviors and ecological interactions through geological time, nor does he even show from their stratigraphic distribution 35
TREE Vol. 6,
that such characters must have evolved within a short interval. Finally, claims for stability (stasis) of behaviors and ecological interactions are based on observation at too few (usually two) points in geological time. These methodological deficiencies are glaring because the problems of recognizing lineages1-3 and of characterizing rates4 and patterns5J of morphological evolution in fossils have been debated extensively. Such questions are serious and as pertinent to behavioral and ecological phenotypes as they are to morphology. Because of these deficiencies, Boucot cannot claim to have demonstrated that behavioral evolution and coevolution are episodic, and thus their tempos cannot be used as support for his model of evolutionary tempo. Equally disappointing is Boucot’s failure to make adequate use of the literature. For example, omission of Vermeij’s’ recent book, which also argues for control of phenotypic innovation by community structure, is unfortunate. Ehrlich and Raven’s8 argument that phenetic cohesion of species is imposed by community structure rather than by gene flow could have been used by Boucot to support his thesis. Van Valen’sg Red Queen hypothesis holds that change in the biotic environment exerts con-
tinuous pressure for evolution rather than causing the episodic crises envisioned by Boucot. Conflicts and congruencies between the author’s views and those presented by others are inadequately explored. Similarly, Boucot sometimes overlooks recent contributions, citing, for example Romer’s’O classic but aging Vertebrate Paleontology instead of Carroll’s” recent comprehensive review of vertebrate paleontology. On the positive side, the volume is packed with interesting and wellorganized examples of behavioral and ecological properties of fossil organisms. Collection of this information in one place is the primary value of the book. Echoing recent suggestions by others’*, Boucot criticizes studies that estimate numbers of taxa and their rates of extinction and origin through geological time without taking into account membership of these taxa in different ecological communities. Aggregation of such data among communities would obscure the episodic nature of evolution and extinction if the episodes are, as Boucot claims, asynchronous among communities. Although Boucot has discussed some interesting issues and gathered a wealth of information on behavior and ecology of fossil organisms, he has not built the empirical edifice he had hoped would support his argu-
~0,
ment for the control tempo by community
7,
J'anuary : ,“!~ ?
of evolutionary structure. Michael A. Bell
Deptof Ecologyand Evolution,StateUnivewry ai New York, Stony Brook,NY 11794.5245,USA
References 1 Schaeffer, B., Hecht, M.K. and Eldredge, N. (1972) Evol. Biol. 6, 31-46 2 Gingerich, P.D. (1979) in Phylogeneric Analysis and Paleontology (Cracraft, J. and Eldredge, N., eds), pp. 41-79, Columbia University Press 3 Bell, M.A. (1988) Trends Ecol. Evol. 3, 320-325 4 Gingerich, P.D. (1983) Science 222, 159-161 5 Bookstein, F.L., Gingerich, P.D. and Kluge, A.G. (1978) Paleobiology4, 120-l 34 6 Bell, M.A. and Legendre, P. (1987) Syst. Zool. 36. 52-61 7 Vermeij, G. (1987) Evolution and Escalation, Princeton University Press 8 Ehrlich, P.R. and Raven, P.H. (1969) Science 165,1228-l 232 9 Van Valen, L. (1973) Evol. Theory 1, l-18 10 Romer, A.S. (1966) Vertebrate Paleontology, University of Chicago Press 11 Carroll, R.L. (1987) Vertebrate Paleontology and Evolution, W.H. Freeman 12 Behrensmeyer, A.K. (1988) Paleogeogr. Paleoclimatol. faleoecol. 63, 183-199
Financed by research ecologists and non-profit-making, the in2ernationel Ecology kWitute (ECI) has a staff of 41 ecologists - marine, terrestrial and limnetic - oil of high pro&s&& rsput&on. N&her the direetw nor any member of the scientific staff receives remuneration. Every year a jury campowd of ECt members s&eta - in a rot&#@ wrn - prize winners among marine, terrestrial or iimnetic ecologists. In 1991, prize winner will be selected in the fictldof limn&k esalogy. The winner of ths Ecology Institute Prize is requested to author a 2011to 300 print&pqe book, to be published by ECI in the series ‘Excellence in Ecology’(EE) and to be made available worldwideat cost price. Ei?s cct~nw@ t8 different from that oftextbooks. In addition to reviewing a cert&~ f&id rif knowledge, itgives the authors a chme to Msatheir pe~~rtelviews on importirnte&ogical issues, to interp~etrrent ~ti~~~owttsdgeon the basis oftheir own mqerience and in&#, an&o tell us what, in their opinion, should b&done in the future. The Ecology Institute Prize is endow&d with a stipend of US$&XlO. Asecond prize may*ba awarded honoring a young ecologist who has conduct@ and pubKsh& res#s of uniqurprtyind research efforts representing an important scientific brMtt&rqjh: the IlppEPR!ZEftritematio ix) is endowed with a stipend of USQSO. Nomktaticms we m alt resslerreh ecol@ts. iersonof tb&!I Jury (see beiow) before July 31, 1991. E#ig&Ieare ait gtaggylin scientific rese I’rtd~~~ranc)t9lt,~ry’schairpereon: Jury mernbersnominated SK&M& The Jury w&El 1x~‘\MFnner&sN t&~nominatibns received, as well as their o&n will be replaced by knowladgrr of top pwb~fnefs, and their own professton& @@m%nt. C&to WWWJ (Director EC.0 Ecology Enstitute,Nordbilnte 23, D-2124 Oldendorf/Luhe, FRG. Tel: 04132i727. Fax: Q413MB83 ECI Limnetic Jury 1991: i, S&oPaula, BF&I; 0. Uhimann, Dresden,