- Email: [email protected]
Eco-logic: Logic-based approaches to ecological modelling
TREE vol. 7, no. 4, April
1992
difficult; newly initiated colonies rarely succeed. Even if individuals do not gain much inclusive fitness as workers, they nonetheless may gain more than if they were trying to nest alone. The difficulties that single wasps face in defending a nest against conspecifics, parasites and predators (particularly ants), in rebuilding a damaged nest, in provisioning the brood (particularly when adults are killed) and in constructing and maintaining the nest, make social life moresuccessful than nesting alone. Throughout this book, the 314 relatedness hypothesis is dismissed as an important factor in the evolution of sociality. Rather, the focus is on the different ecologies of wasps that shape their widely divergent patterns of social life. For those who think of a social insect colony as a finely tuned ‘supera surprise is in store. organism’, Nearly every chapter describes discord: workers bite, pull on and apparentlytest queens; queens bite and chase workers; workers maul and dominate each other; and workers force larvae to feed them a nutritious saliva, underthreatof being cannibalized. Even in highly social species with few obvious signs of competition, removal of the queen results in immediate changes: workers become quarrelsome and restless, their ovaries develop and they soon begin laying and eating each others’ eggs. Upon the queen’s return, the workers may no longer recognize her and she may be attacked. The colony is under
-
constant tension and harassment from the queen, from other members of the group and from marauding conspecifics. Intra- and intergroup competition have been a major force in wasp social evolution. The diversity of wasp nesting behavior provides a wonderful opportunity for comparisons, but comparative studies of wasps have been hindered by the fact that the phylogenetic relationships between taxa have not been known. The opening chapter brings together previously published cladograms and presents a new one. Using these Carpenter evaluates Westtrees, Eberhard’s model of wasp social evolution and concludes that polygynous (multiple queen) family groups with rudimentary castes are the likely ancestral condition for all social vespids. His analysis comes to the interesting conclusion that group living and some division of labor were characterisic of the common ancestor that stenogastrines share with polistines and vespines. This means that the unusual nesting biology of the stenogastrines, with their abdominal secretions, ant guards and diverse nest architecture, evolved after they separated from their common ancestor. This presents an interesting puzzle: Why did the stenogastrines fail to evolve high levels of sociality and why did they evolve ‘reversions’ to more primitive levels of social organization, unlike the polistines and vespines? Turillazzi speculates
that the limits to social complexity among stenogastrines can be found in their rudimentary prey-masticating behavior, in their poor nest materials and in the limitations imposed by their dependence on abdominal secretions for brood rearing. These are arguments of constraint; stenogastrines simply never invented the sorts of adaptations that arose in the other lineage. One could also look for differences in ecology that favored larger colony size and greater levels of integration in polistines and vespines as compared with stenogastrines. Modern comparative studies are a powerful technique for sorting out evolutionary hypotheses but, of cause, the approach requires reliable phylogenies for each group. As West-Eberhard observes in the introduction, conceptual progress in a field depends on ‘a continuous interweaving of fact and theory’. Descriptions of biology and behavior are woven together into patterns and those patterns are given adaptive explanations. The field advances when generality is found among the explanations from diverse groups. The Social Biology of Wasps identifies a fascinating array of patterns and it proposes explanations, but it does not pull together the common threads nor discuss the generalities.
are the time and intellectual energy needed to take in all the heavy stuff, and the pain one suffers until models actually work on the screen of the computer and can be compared satisfactorily with observations. The books attack these two problems, respectively. Renshaw has done an excellent job, sailing between the Scylla of lacking necessary mathematical rigor and the Charybdis of being biologically vacuous. To be frank, those who cannot stand basic algebra and elementary calculus should not open the book, but those who are willing to struggle through such formalism in the hope of understanding deterministic and stochastic, homogeneous and spatially structured models will gain a lot. Derivations that are easy to follow are presented explicitly, whereas painful parts are only referred to. Eminent biological case studies, often beauti-
ful and classic ones, are amply cited and explained. Yet this is not all. The reader is aided by the presentation of brief example programs, written in FORTRAN and using the NAG routines. The PC package MINITAB can also be used to elaborate on the examples or one’s own problems. A future edition of this book might usefully include examples generated by the packages SCOP and MATHEMATICA. Renshaw’s book offers far more than immediate practical use; it is also thoughtfully written. Having read it, one is likely to have a more balanced view of the (complementary) roles of deterministic and stochastic models. His comments on deterministic chaos versus stochastic fluctuations are highly interesting: he argues, for example, that different patterns in deterministic dynamics are relevant only if they are not
H. Jane Brockmann Deptof Zoology,Universityof Florida,Gainesville, FL32611,USA
Theory and Programming Modelling Biological Populations in Space and Time
by Eric Renshaw, Cambridge University Press, 1991. f60.00/$110.00 hbk (xvii + 403 pages) ISBN 0 521 30388 5 Eco-Logic: Logic-BasedApproaches to Ecological Modelling by David Robertson, Alan Bundy, Robert Muetzelfeldt, Mandy Haggith and Michael Uschold, The MIT Press, 1991. $35.OO/f31.50 hbk (243 pages) ISBN 0 262 18143 6 Mathematical modelling in ecology has a long tradition. Verhulst published the logistic growth curve in 1838, and Lotka and Volterra were very active in the 1920s. Since then, a formidable arsenal of mathematical tools has been developed. Two of the many difficulties arising from this
141
TREE vol. 7, no. 4, April
destroyed by the effect of genuine stochasticity. (Here, I regret that Renshaw did not consider the links between chaos and epidemiology, although he deals with them separately; see Ref. 1 for a recent overview.) In the second book, the Edinburgh Artificial Intelligence group argue that ecological modelling could be given a novel basis by logic programming, using various versions of the language PROLOG. In their view the snag with traditional programs, using an imperative language, is that the structure of the model itself and the commands for the execution of the particular program (which is not a unique implementation of the model) are hopelessly tangled, making life difficuIt,especiaIlyforecologistswith less formal or algorithmic training. The major problem is that the book will convince very few people that
they should choose Eco-Logic instead. One reason for this is implicit in the group’s concluding remarks: ‘We believe that further progress in improving model comprehension will be best achieved by promoting collaboration between researchers in simulation, software engineering, log programming and artificial intelligence, and that this research must be motivated by the demands of a non-trivial domain of application’ (p. 215). One has the (maybe unjustified) impression that in fact during the project of writing the book, the authors did not collaborate enough with ecologists, and they did not consider nontrivial cases in ecology. People having ‘formulae phobia’ and a dislike of programming will not find this book easy. Despite these criticisms, I would be interested to see how this approach develops in the future. The authors
Review copies of the following books have been received. The appearance of a book in the list does not preclude the possibility of it being reviewed in TREE in the future.
Charles E. Dinsmore ted.) A History ation Research: Milestones in the a Science Cambridge University f30.00C654.95 hbk (x + 228 pages) 39271 3
S. Blackmore and R.B. Knox teds) Microspores: Evolution and Ontogeny Academic Press, 1990. f49.50C699.00 hbk (x + 347 pages) ISBN 0 12 103458 5 Robert W. Blake ted.) Efficiency and Economy in Animal Physiology Cambridge University Press, 1991. f30.00/$59.95 hbk (xii + 187 pages) ISBN 0 521 40066 X British Museum (Natural History) Man’s Place in Evolution (2nd edn) Natural History Museum Publications and Cambridge University Press, 1991. f7.95/$12.95 pbk (103 pages) ISBN 0 521 40864 4
of RegenerEvolution of Press, 1991. ISBN 0 521
William H. Durham Coevolution: Genes, Cuhure, and Human Diversify Stanford University Press, 1991. $65.00 hbk (xxii + 629 pages) ISBN 0 8047 1537 8 Frank Ellis Agricultural Policies in Developing Countries Cambridge University Press, 1992. f40.00/$69.95 hbk, fl4.95/$29.95 pbk (xv + 357 pages) ISBN 0 521 39584 4 Ft. W. Elwood and S.J. Neil Assessments and Decisions: A Study of information Gathering by Hermit Crabs Chapman & Hall, 1992. f29.95 hbk (ix + 192 pages) ISBN 0 412 39110 4
....I.... ‘“‘,““,.~,,~““,~ >,....“:_~...“..‘.“.“..** .~” .......“.Y,..“.y ,...L,.... P..S ,..... y:* ........x.*.,,q*~
7992
conjecture: ‘By analogy with Knuth’s concept of metafonts, from which any particular font can be implemented by a process of specialisation, we could think of the superparadigm as being a meta-model, from which (in a similar way) any particular model can be built’ (p. 16). I suspect that either the domain of such a meta-model will be restricted or, by the application of GGdel’s theorem, there will be particular models that cannot be built from the meta-model by specialization.
EiirsSzathmziry Laboratoryof Mathematical Biology, MRCNational Institute for Medical Research, The Ridgeway, Mill Hill, London NW7 lAA, UK
Reference 1 Olsen, L.F. and Schaffer, W.M. (1990) Science 249,499-504
Verne Grant The Evolutionary Process: A Critical Study of Evolutionary Theory (2nd edn) Columbiauniversity Press, 1991.$60.00 hbk(xviii +487 pages) ISBN 0 231 07324 0 Martin Lockley Tracking Dinosaurs: A New Look atanAncienr WorldCambridge University Press, 1991. f27.50/$39.50 hbk, f9.95/$14.95 pbk (xii + 238 pages) ISBN 0 521 42598 0 Frank K. McKinney and Jeremy B.C. Jackson Bryozoan Evolution (pbk edn) The University of Chicago Press, 1991. $15.95/fl2.75 pbklxii + 238 pages) ISBN 0 226 56047 3 H. Frederik Nijhout The Development and Evolution of Butterfly Wing Patterns Smithsonian Institution Press, 1991.$53.95/f34.95 hbk, $23.95/f15.50 pbk (xvi + 297 pages) ISBN 0 87474 921 2
*.... ‘...‘.y.*,.“ *... .*..U**.*r .. ......‘..,.... i.r......*.s.r .......*... ‘..*..* l”.......... ....*..,“*L*I” ................... ? ..,’
.........n,~.~-..r+C..ll.*.,~~u...~ .....“*“‘* .....*..,,a..... *** ......“.*~\r.~.i-.‘“y,‘~,*.*.-‘ ....*.*+.. *...r*.*-u.~r.*.u* .....1........I..I,........*..... ......................*,....I..4..I......I.I......I..........I Addres.s ....”.......*I,“‘*,**.“,.u .I_ . . . . . . . . . . *,**~..I...,“ .I,.. .* *.*.*4...* ..,,.a.,....‘...........I....% ...**“,.*l.*.,.l.l .,..*.,,.,....,, ****.** ,..,....,... ,.../.... . ..*.~~.z.,....i..~.*X .a.* u.nn;.i..* . . . . . “ .,...‘ “ “ “ .... ,....,... _ ....,,.#..,.,,....,. %...^,, ....*..a. .......,,,“I..I.n‘ .’ 1 .~““”.......V.....‘~*..~L.*“‘ )..“ ,,.,_ ..‘.....I.....I....... >.*.,* II.....a.1 .......*.,:*,................ *.,.~..l*.‘.+,....... *.** .........*..*..... ,“.*1...n.r.,.**~.c.~..,~~~*~..~..~.~.~.~~~.~” ...o.... .“....““,II__, ....~~...‘ .~,. ..~.“‘ Name of ~~~~~~ .’ >
142
1992
difficult; newly initiated colonies rarely succeed. Even if individuals do not gain much inclusive fitness as workers, they nonetheless may gain more than if they were trying to nest alone. The difficulties that single wasps face in defending a nest against conspecifics, parasites and predators (particularly ants), in rebuilding a damaged nest, in provisioning the brood (particularly when adults are killed) and in constructing and maintaining the nest, make social life moresuccessful than nesting alone. Throughout this book, the 314 relatedness hypothesis is dismissed as an important factor in the evolution of sociality. Rather, the focus is on the different ecologies of wasps that shape their widely divergent patterns of social life. For those who think of a social insect colony as a finely tuned ‘supera surprise is in store. organism’, Nearly every chapter describes discord: workers bite, pull on and apparentlytest queens; queens bite and chase workers; workers maul and dominate each other; and workers force larvae to feed them a nutritious saliva, underthreatof being cannibalized. Even in highly social species with few obvious signs of competition, removal of the queen results in immediate changes: workers become quarrelsome and restless, their ovaries develop and they soon begin laying and eating each others’ eggs. Upon the queen’s return, the workers may no longer recognize her and she may be attacked. The colony is under
-
constant tension and harassment from the queen, from other members of the group and from marauding conspecifics. Intra- and intergroup competition have been a major force in wasp social evolution. The diversity of wasp nesting behavior provides a wonderful opportunity for comparisons, but comparative studies of wasps have been hindered by the fact that the phylogenetic relationships between taxa have not been known. The opening chapter brings together previously published cladograms and presents a new one. Using these Carpenter evaluates Westtrees, Eberhard’s model of wasp social evolution and concludes that polygynous (multiple queen) family groups with rudimentary castes are the likely ancestral condition for all social vespids. His analysis comes to the interesting conclusion that group living and some division of labor were characterisic of the common ancestor that stenogastrines share with polistines and vespines. This means that the unusual nesting biology of the stenogastrines, with their abdominal secretions, ant guards and diverse nest architecture, evolved after they separated from their common ancestor. This presents an interesting puzzle: Why did the stenogastrines fail to evolve high levels of sociality and why did they evolve ‘reversions’ to more primitive levels of social organization, unlike the polistines and vespines? Turillazzi speculates
that the limits to social complexity among stenogastrines can be found in their rudimentary prey-masticating behavior, in their poor nest materials and in the limitations imposed by their dependence on abdominal secretions for brood rearing. These are arguments of constraint; stenogastrines simply never invented the sorts of adaptations that arose in the other lineage. One could also look for differences in ecology that favored larger colony size and greater levels of integration in polistines and vespines as compared with stenogastrines. Modern comparative studies are a powerful technique for sorting out evolutionary hypotheses but, of cause, the approach requires reliable phylogenies for each group. As West-Eberhard observes in the introduction, conceptual progress in a field depends on ‘a continuous interweaving of fact and theory’. Descriptions of biology and behavior are woven together into patterns and those patterns are given adaptive explanations. The field advances when generality is found among the explanations from diverse groups. The Social Biology of Wasps identifies a fascinating array of patterns and it proposes explanations, but it does not pull together the common threads nor discuss the generalities.
are the time and intellectual energy needed to take in all the heavy stuff, and the pain one suffers until models actually work on the screen of the computer and can be compared satisfactorily with observations. The books attack these two problems, respectively. Renshaw has done an excellent job, sailing between the Scylla of lacking necessary mathematical rigor and the Charybdis of being biologically vacuous. To be frank, those who cannot stand basic algebra and elementary calculus should not open the book, but those who are willing to struggle through such formalism in the hope of understanding deterministic and stochastic, homogeneous and spatially structured models will gain a lot. Derivations that are easy to follow are presented explicitly, whereas painful parts are only referred to. Eminent biological case studies, often beauti-
ful and classic ones, are amply cited and explained. Yet this is not all. The reader is aided by the presentation of brief example programs, written in FORTRAN and using the NAG routines. The PC package MINITAB can also be used to elaborate on the examples or one’s own problems. A future edition of this book might usefully include examples generated by the packages SCOP and MATHEMATICA. Renshaw’s book offers far more than immediate practical use; it is also thoughtfully written. Having read it, one is likely to have a more balanced view of the (complementary) roles of deterministic and stochastic models. His comments on deterministic chaos versus stochastic fluctuations are highly interesting: he argues, for example, that different patterns in deterministic dynamics are relevant only if they are not
H. Jane Brockmann Deptof Zoology,Universityof Florida,Gainesville, FL32611,USA
Theory and Programming Modelling Biological Populations in Space and Time
by Eric Renshaw, Cambridge University Press, 1991. f60.00/$110.00 hbk (xvii + 403 pages) ISBN 0 521 30388 5 Eco-Logic: Logic-BasedApproaches to Ecological Modelling by David Robertson, Alan Bundy, Robert Muetzelfeldt, Mandy Haggith and Michael Uschold, The MIT Press, 1991. $35.OO/f31.50 hbk (243 pages) ISBN 0 262 18143 6 Mathematical modelling in ecology has a long tradition. Verhulst published the logistic growth curve in 1838, and Lotka and Volterra were very active in the 1920s. Since then, a formidable arsenal of mathematical tools has been developed. Two of the many difficulties arising from this
141
TREE vol. 7, no. 4, April
destroyed by the effect of genuine stochasticity. (Here, I regret that Renshaw did not consider the links between chaos and epidemiology, although he deals with them separately; see Ref. 1 for a recent overview.) In the second book, the Edinburgh Artificial Intelligence group argue that ecological modelling could be given a novel basis by logic programming, using various versions of the language PROLOG. In their view the snag with traditional programs, using an imperative language, is that the structure of the model itself and the commands for the execution of the particular program (which is not a unique implementation of the model) are hopelessly tangled, making life difficuIt,especiaIlyforecologistswith less formal or algorithmic training. The major problem is that the book will convince very few people that
they should choose Eco-Logic instead. One reason for this is implicit in the group’s concluding remarks: ‘We believe that further progress in improving model comprehension will be best achieved by promoting collaboration between researchers in simulation, software engineering, log programming and artificial intelligence, and that this research must be motivated by the demands of a non-trivial domain of application’ (p. 215). One has the (maybe unjustified) impression that in fact during the project of writing the book, the authors did not collaborate enough with ecologists, and they did not consider nontrivial cases in ecology. People having ‘formulae phobia’ and a dislike of programming will not find this book easy. Despite these criticisms, I would be interested to see how this approach develops in the future. The authors
Review copies of the following books have been received. The appearance of a book in the list does not preclude the possibility of it being reviewed in TREE in the future.
Charles E. Dinsmore ted.) A History ation Research: Milestones in the a Science Cambridge University f30.00C654.95 hbk (x + 228 pages) 39271 3
S. Blackmore and R.B. Knox teds) Microspores: Evolution and Ontogeny Academic Press, 1990. f49.50C699.00 hbk (x + 347 pages) ISBN 0 12 103458 5 Robert W. Blake ted.) Efficiency and Economy in Animal Physiology Cambridge University Press, 1991. f30.00/$59.95 hbk (xii + 187 pages) ISBN 0 521 40066 X British Museum (Natural History) Man’s Place in Evolution (2nd edn) Natural History Museum Publications and Cambridge University Press, 1991. f7.95/$12.95 pbk (103 pages) ISBN 0 521 40864 4
of RegenerEvolution of Press, 1991. ISBN 0 521
William H. Durham Coevolution: Genes, Cuhure, and Human Diversify Stanford University Press, 1991. $65.00 hbk (xxii + 629 pages) ISBN 0 8047 1537 8 Frank Ellis Agricultural Policies in Developing Countries Cambridge University Press, 1992. f40.00/$69.95 hbk, fl4.95/$29.95 pbk (xv + 357 pages) ISBN 0 521 39584 4 Ft. W. Elwood and S.J. Neil Assessments and Decisions: A Study of information Gathering by Hermit Crabs Chapman & Hall, 1992. f29.95 hbk (ix + 192 pages) ISBN 0 412 39110 4
....I.... ‘“‘,““,.~,,~““,~ >,....“:_~...“..‘.“.“..** .~” .......“.Y,..“.y ,...L,.... P..S ,..... y:* ........x.*.,,q*~
7992
conjecture: ‘By analogy with Knuth’s concept of metafonts, from which any particular font can be implemented by a process of specialisation, we could think of the superparadigm as being a meta-model, from which (in a similar way) any particular model can be built’ (p. 16). I suspect that either the domain of such a meta-model will be restricted or, by the application of GGdel’s theorem, there will be particular models that cannot be built from the meta-model by specialization.
EiirsSzathmziry Laboratoryof Mathematical Biology, MRCNational Institute for Medical Research, The Ridgeway, Mill Hill, London NW7 lAA, UK
Reference 1 Olsen, L.F. and Schaffer, W.M. (1990) Science 249,499-504
Verne Grant The Evolutionary Process: A Critical Study of Evolutionary Theory (2nd edn) Columbiauniversity Press, 1991.$60.00 hbk(xviii +487 pages) ISBN 0 231 07324 0 Martin Lockley Tracking Dinosaurs: A New Look atanAncienr WorldCambridge University Press, 1991. f27.50/$39.50 hbk, f9.95/$14.95 pbk (xii + 238 pages) ISBN 0 521 42598 0 Frank K. McKinney and Jeremy B.C. Jackson Bryozoan Evolution (pbk edn) The University of Chicago Press, 1991. $15.95/fl2.75 pbklxii + 238 pages) ISBN 0 226 56047 3 H. Frederik Nijhout The Development and Evolution of Butterfly Wing Patterns Smithsonian Institution Press, 1991.$53.95/f34.95 hbk, $23.95/f15.50 pbk (xvi + 297 pages) ISBN 0 87474 921 2
*.... ‘...‘.y.*,.“ *... .*..U**.*r .. ......‘..,.... i.r......*.s.r .......*... ‘..*..* l”.......... ....*..,“*L*I” ................... ? ..,’
.........n,~.~-..r+C..ll.*.,~~u...~ .....“*“‘* .....*..,,a..... *** ......“.*~\r.~.i-.‘“y,‘~,*.*.-‘ ....*.*+.. *...r*.*-u.~r.*.u* .....1........I..I,........*..... ......................*,....I..4..I......I.I......I..........I Addres.s ....”.......*I,“‘*,**.“,.u .I_ . . . . . . . . . . *,**~..I...,“ .I,.. .* *.*.*4...* ..,,.a.,....‘...........I....% ...**“,.*l.*.,.l.l .,..*.,,.,....,, ****.** ,..,....,... ,.../.... . ..*.~~.z.,....i..~.*X .a.* u.nn;.i..* . . . . . “ .,...‘ “ “ “ .... ,....,... _ ....,,.#..,.,,....,. %...^,, ....*..a. .......,,,“I..I.n‘ .’ 1 .~““”.......V.....‘~*..~L.*“‘ )..“ ,,.,_ ..‘.....I.....I....... >.*.,* II.....a.1 .......*.,:*,................ *.,.~..l*.‘.+,....... *.** .........*..*..... ,“.*1...n.r.,.**~.c.~..,~~~*~..~..~.~.~.~~~.~” ...o.... .“....““,II__, ....~~...‘ .~,. ..~.“‘ Name of ~~~~~~ .’ >
142