TREE vol. 6, no.

7, July

1991

Effectsin Community Ecology: Their Definition,Study and Importance

Sharon Y. Strauss
The diversity of indirect interactions tlzaf cdfl occur within communities is large. Recent research on indirect interactions is scattered in tCle literature under numerous labels. Tke definition of indirect effects is an important aspect of their study, and clarifies some of tCre subtle differences among indirect effects found in natural communities. Choosing which species to study, ctow to manipulate species and for wkat duration, wkic/i attributes to Measure and, finally, wfiick analytical tec/iniques to use are all proMems facing the cofflmunitg ecologist. Ukimatelg, we tire striving for the Gest meuns of determining tCze relative importance of direct and indirect effects in structuring comnzunities. species could mistakenly be construed as competitors when, in fact, the mechanism behind their reciprocal negative interactions was a shared predator. Holt termed this phenomenon apparent competition because in the absence of the predator, the species did not compete; this work came to exemplify the quintessential indirect effect. How interactions among species at the same trophic level can be mediated by other species at an adjacent trophic level reflects a slightly different emphasis from studies such as Vandermeer s that consider interactions within trophic levels, or those that consider cascading indirect effects across multiple trophic levels5,6. Definition of indirect effects Indirect effects are generally defined as how one species alters the effect that another species has on a thirdI or, similarly, how and to what degree pairwise species interactions are influenced by the presence and density of other species in the community 7. For both of these definitions, three is the minimum number of species required for indirect effects. Such effects are usually achieved through trophic and/or competitive interactions between two species that effect changes in population densities or per capita growth rates of a third species@. A multitude of phenomena, bearing diverse labels, can fall under the heading of indirect effects: exploitative competition, apparent competition, facilitation, some mutualisms, cascading effects, tri-trophic-level interactions, higher-order interactions and non-additive effects, to name the most common. Perhaps one of the more controversial interactions included under the heading of indirect effects is that of exploitative competition. An informal survey of colleagues revealed that most of them considered interspecific competition to be a direct interaction. But the majority of studies on interspecific competition indicate that resource exploitation, rather than

All organisms exist as part of a community, and yet relatively few experiments investigate interspecies within the actions among coexisting context of multiple species. The importance of the community context becomes apparent with the observation that the strength and often the direction of the interaction between two species may change in the presence of others - these are indirect effects. Whether the effects of species in combination can be predicted from knowledge of their pairwise interactions was first addressed experimentally by Vandermeer . His experiments with communities of up to four competing protozoans were designed to determine whether the effects of species together were additive, or whether higher-order interactions occurred when more than two species were present; such interactions would indicate that indirect effects occurred within this trophic level. In fact, Vandermeer found no higher-order interactions in these laboratory communities, but a good short review of studies on non-additive effects is presented by Wilbur and Fauth2. Holt3, following work by Levine4. that two theoretically argued
Sharon Strauss is at the Illinois Natural History Survey, 607 E Peabody Drive, Champaign, IL 61820, USA.

direct interference, is the mechanism behind competition9. Since exploitation of one species (the prey) by another can have a negative influence on a third, such competition better fits the definition of an indirect effect than a direct effect. Species interactions can be mediated by a nonliving resource. just as two herbivores can interact indirectly by modifying the quality, as opposed to quantity, of their shared host plant via induced chemical or physical plant responses 2, 1, other organisms can interact analogously through changes in a physical resource. Alteration of water quality in temporary ponds effected by the first species to colonize may affect the subsequent success of other species13. Such interactions are usually termed priority effects or historical effects , and do not necessarily fit the definitions of indirect effects given above. One could argue that to distinguish interspecific interactions that are mediated by qualitative changes in living resources such as plants, as opposed to nonliving resources such as water or soil, is to split hairs. The interactions among organisms associated with carrion fit into neither category comfortably, although the species composition of carrion-feeding communities is also vulnerable to priority effectsI 7 Wilsoni gives a comprehensive list of nontrophic interactions, including interactions in which one species modifies a nonliving resource that subsequently affects the success of a second species. In general, because a species can potentially exert a selective force on another through nontrophic interactions, such effects should probably be included under the broad heading of indirect effects. They are especially appropriately called indirect effects since often the species effecting resource modification are temporally separated from the object(s) of such modification. Ultimately, the designation ot an interaction as an indirect effect becomes almost trivial, since all interactions-aside from predation (and subcategories parasitism and herbivorylh), some mutualisms and interference competition - can be construed as indirect. To organize this broad field, Miller and Kerfoot7 propose categories to distinguish

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effect: how relationship the between species A and species B is affected focal Table I. Typesof indirect by a third species, C among the manifold mechanisms behind indirect effects (Table 1). Studies of indirect effects have to date hinged primarily on changes in abundance of the target species. However, indirect effects may result in biologically significant changes in a species that are not reflected in its abundance. Vannis found that direct effects of fish predation changed the population structure of cladoceran zooplankton populations. Larger individuals were eaten by fish, thus favoring individuals that metamorphosed at a smaller size. Because smaller individuals had more resources available to them as a result of decreased intraspecific competition, their reproductive rates increased. The overall effect was that cladoceran populations did not change in density, but did change in mean body size and population structure. In a separate similar changes in both study, cladoceran size and density were a result of indirect effects when piscivores were introduced to this community. How might demographic changes in one species affect community structure? Since smaller individuals may eat different prey species than larger individuals, they may alter the relative abundance of species in prey communities. Another potential indirect effect that has received little explicit attention is the alteration of a species genotypic composition as opposed to its abundance. For example, in the Bahamas, buttonwood trees (Conocarpus erectus) with silver (pubescent) leaves receive less leaf damage from herbivorous insects than do those with smooth leavesIs. On islands where lizards occur, the proportion of buttonwood trees that are silver is low; on islands where lizards are absent-and herbivorous arthropods abundant - more trees are silver. Schoener suggests that * the mechanism responsible for the positive correlation between the proportion of buttonwood trees with silver leaves and lizard absence is the indirect effect of the lizards reduction of herbivore populations. Since leaf attributes of buttonwood are probably heritable (Semple as cited in Ref. 191, an indirect effect of lizard introduction may be to change the genetic composition, as opposed to the density, of buttonwood populations.
Type of indirect 1. Abundancea effect Effect of sp. C Change in abundance of sp. 6 through predation, competition or facilitationb Response of focal sp. A Change in abundance, per capita growth rateb, demographyb or genotypeb via change in sp. 6 Change in abundance, per capita growth rate, demography or genotype via change in sp. 6

2. BehavioraP. morphological, or physiological

chemical

Change in behavior, morphology, chemistry physiology, but not necessarily abundance, sp. B Change in quality resource

or of

3. EnvironmentaP

of abiotic

Change in abundance, per capita growth rate, demography or genotype via change in shared resource Change in abundance, per capita growth rate, demography or genotype as a result of altered response of sp. A to sp. B caused by the presence of sp. C

4. Response

None

a Terms altered from original b Added by author.

discussion

in Ref. 7 (T. Miller,

pers. commun.).

How genotypic changes within one species can affect the presence or behavior of others within the community is less obvious. Changes in genotypic composition may alter a species susceptibility to disease or other environmental factors, thereby affecting its (and potentially other species ) density. In addition, recent experimental evidence suggests that genotypic changes in one species may be matched by changes in others20, a subject that will be discussed in more detail below. In all, the ways in which species interact indirectly are diverse and numerous, and require careful observation and experimental manipulation for full enumeration. How important are indirect effects? The relative importance of direct versus indirect effects in structuring communities is a key question and an area in which results are generally lacking. Painezl discusses how the existence of modules - groups of species that interact with each other, but that affect few other species outside the module - can depend on the indirect effects of strongly interacting co-occurring, species (e.g. those suppressing competitively dominant species). WiIson15J2 and others2 ,23.24 have suggested that indirect effects may be very important, and some say the most important, processes affecting the success of a species. Species

may be adapted not only to feed efficiently and to protect themselves from predators, but also to modify their community in ways that loop back beneficially to themselves 22. Indirect effects are thus postulated to be the unit interaction of community evolution. Over evolutionary time, the effect of the community on the members comes to reflect the effect of that member on the community, and only those that positively contribute to community welfare persist.5 Recent work by GoodnightZo lends credence to this view. Using laboratory populations of two species of Ebolium, competing he found that by selecting for attributes in one species (emigration rate, population size) he obtained correlated concurrent genetic changes in the other species. These results suggest that species may be able to adapt to indirect effects and that communities, cemented together by such interactions, may themselves succeed and fail based on the dynamics of all species taken in toto. Studying indirect effects: selecting species In studying indirect effects, investigators tend to manipulate species they suspect will have strong effects on the community. An often implicit assumption is that if a species has strong direct effects, it will be more likely to exhibit strong indirect 207

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effects. Such an assumption is probably correct in most systems. Abundant species are often taken to be strongly interacting species; however, less-common species often exert considerable influence on community structure and can be easily overlooked25. A classic example of an influential but only moderately abundant species is the mussel Mytilus californianus, which can eliminate many sessile species at multiple trophic levels once released from Pisaster starfish predation26. Several authors point out that identifying strong interactors in the field is not a trivial matter and cannot be based solely on species abundance25-27. Some investigators do larger community manipulations by removing trophic species - guilds of species that feed, and are fed upon, in a similar manner 8,28,29-and in the process may get around difficulties of selecting individual species. Removal of trophic species is likely to result in indirect effects that extend beyond a single trophic level, since, by definition, no other species perform the same trophic role in the community29. Guilds have occasionally been defined by other parameters. Menge et a1.30based their guilds on a combination of trophic habit and species size and vagility; consequently, they found some trophic overlap between guilds, a fact that they felt may weaken the direct and indirect effects of any one guild. Fowler27 compared community responses to deletion of guilds as well as species. She found that classification of plants into guilds based on taxonomic and morphological traits fdicots versus grasses) had little meaning with respect to community in structure; contrast, grasses grouped according to growing season (C, versus C, species) did appear to form competing guilds. Guilds may offer a useful way of including many species in a study, but because of the hazards of placing species into guilds25, and because the characters used to define guilds are not always meaningful with respect to community structure, they may not be a substitute for manipulations of individual species. This is in spite of the fact that, in many cases, one species will dominate the behavior of a guild through its sheer abundance 8,30.

Selecting species and guilds for the study of direct and indirect effects leads inexorably to the study of food webs. A large body of theory has grown around the structure and stability of food webs3r-33. The presence or absence of indirect effects has been postulated to be linked to food-web attributes such as connectance, strengths of interaction among species, numbers of omnivorous species and the competitive stature of preferred prey species2 . For example, several authors have suggested that in complex webs with omnivorous species, more species and greater numbers of interactions among species, indirect effects may become diluted due to noise in transmission34-36. In general, however, the food-web literature is filled with controversy, and space does not allow presentation of the diverse viewpoints here (see Refs 37 and 38 for recent TREE reviews and comments). Studying indirect effects: experimental design The vast majority of experiments use species removals to investigate community structure. Removals allow one to determine the effect of a particular species at that density on the rest of the community. Because natural populations undergo fluctuations in density, impact on the community of the same species removal may vary between seasons or through time. Few investigators use a continuum from zero to high densities to determine the role of a species in a community (but see Refs I3,39,40). Threlkeld40 found, in artificial communities, that although low densities of fish altered zoaplankton species composition and density, it took high densities of fish to alter the phytoplankton community at the next trophic level. Threlkeld goes further to suggest that use of density continua may point out nonlinearities in foodweb responses, and may reconcile studies whose results appear contradictory (see also Ref. 2). Since natural populations often fluctuate widely in their densities, the same arguments may apply to the effects of species removals over different years or seasons. Bender et a/.4 propose that the manner in which species are removed from communities may aid in separating direct from indirect

effects. They suggest that in pulse experiments, where a species is removed completely only once, the change in per capita growth rate of remaining species reflects the direct effects of the removed species. In contrast, press experiments maintain removals over several generations of the target species; longer-term responses of the community to such removals, measured through changes in species abundance, are proposed to be a combination of both direct and indirect effects. This model makes the biological assumption that indirect effects require more time than direct effects. Whether such an assumption is justified needs further investigation; at least one studyJ2 finds that indirect effects occur very rapidly. An early approach to examining community properties was to build communities of increasing complexity Wilbur and Fauth2 have used . this building-block approach to examine the relationships between two anuran prey species and two predaceous species, using a factorial design. Their purpose was to determine the additivity of interactions among four species at two trophic levels. They estimated the strength of intraspecific competition from single-species treatments at several densities for both prey species. The experimental design used all combinations of the four species. With no predators, the two prey species competed; in treatments with one or two predator species, indirect effects resulted in the two prey species acting mutualistically by spreading the risk of predation. In contrast to the prey species, the effects of the two predators together on a single prey species were additive. The lack of fit of predictions based on pairwise interactions between species - i.e. linear, additive effects - reflects the general importance of indirect effects in this community. Detecting indirect effects When a community is perturbed and no indirect effects of that removal or addition are found, what conclusions are to be drawn? it is likely that any species removal or addition will have indirect effects, given the complex nature of communities. However, if these effects are swamped by the natural varia-

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bility of populations, we must assume that they are of minor importance to community structure. There are, however, a number of reasons why one might fail to detect important indirect effects that are occurring. First, as mentioned earlier, the magnitude of the perturbation may govern the nature of the response. Second, direct and indirect effects may be of opposite sign and similar magnitude, thus obscuring both effects. Third, as discussed in the definitions section above, indirect important effects may not always be exhibited as changes in species abundance. There are situations in which the magnitude of indirect effects is easy to measure. For example, suppose that species C eats only species B, which in turn eats species A; then if removal of C affects the abundance of A, that effect must be solely indirect. If direct and indirect effects affect the same species, but are opposite in direction, detecting either or both effects is not straightforward. For example, the grazing activity of Chironomus larvae on lake bottoms decreased bacterial densities (direct effect)43. At higher densities of larvae, however, decreases in bacteria were not proportional to Chironomus densities. Bioturbation of the sediments by Chironomus stimulated bacterial growth (indirect effect) and thus compensated for some of the decrease in bacterial densities due to the direct effects of grazing. Had both effects been equal in magnitude, one might have concluded that there was no effect of grazing on bacterial density. Similarly, a predator may have a direct negative effect on a prey species by consuming it, along with an indirect positive effect by consuming a competing prey species. The future There is still a long way to go in the study of indirect effects. Experimental protocol remains a crucial area of research. How to measure and quantify indirect effects, and how to distinguish them from direct effects, is a subject hotly debated (for the most recent contributions in a series see Refs 44,451. Each experimental method to date yields some information, but none is fully capable of discerning the relative contributions of direct and indirect effects free of artifact. Factorial

designs analysed with ANOVA can often detect the presence of higherorder interactions2; however, as mentioned earlier, lack of detection of higher-order interactions does not necessarily indicate that they are absent (if, for example, direct and indirect effects are equal and opposite in magnitude). As advocated by some, path analysis may go further as a tool for determining the relative strengths of indirect versus direct effects2*23.In such analysis, it is assumed that all relationships among species are known beforehand, and they are specified in a food web (path) that includes cross-links within trophic levels to accommodate competitive and facilitative interactions. Direct impacts of species upon one another are estimated using standardized partial regression coefficients (usually based on species abundances) in pair-wise comparisons; indirect effects are predicted through addition of pairwise effects along a specified path. In theory, this technique should allow better separation of the magnitudes of direct and indirect effects. However, the accuracy of these estimates is predicated on complete prior knowledge of all interactions within the food web (not a trivial matter), and on how paths are counted; it is also vulnerable to assumptions about both the directional and the mechanistic properties of interspecific relationships (see Refs 45,46 for more comprehensive lists). In wrestling with the problem of experimental protocol, Loeh1e43 suggests giving up on any attempt to separate direct from indirect effects. This is perhaps the result of the painful recognition that, to date, knowledge of exactly how important indirect effects are has only been gained from intimate knowledge and detailed experimentation within individual systems. Studying indirect effects will remain difficult at best. Experiments examining the structure of multispecies communities are necessarily elaborate, and their statistical analysis can be equally so. Nonetheless, without such painstaking studies we cannot decide whether, or when, indirect effects are important forces in determining biological diversity. We also must try to determine how indirect an effect can be and still be

influential. In a time when perturbation of natural communities is common, it behooves us to persevere and to learn as much as possible about the forces that affect species persistence.
Acknowledgements I thank T. Schoener and T. Miller for extensive discussions. Helpful comments on various versions of the manuscript were gratefully received from T. Miller, T. Schoener, M. Schwartz, D. Simberloff, P. Stiling, C. Toft and anonymous reviewers.

References

I Vandermeer, I.H. II9691 Ecology 50, 362-371 2 Wilbur, H.M. and Fauth, I.E. (19901 Am. Nat. 135, 176-204 3 Holt, R.D. 119771 Theor. Popul. Biol. 12. 197-227 4 Levine, S.H. (19761 Am. Nat. I IO. 903-910 5 Kerfoot, WC. and Sih. A. f 198 Predation 71 Direct and indirect Impacts on Aquatic Communities, University Press of New England 6 Abrams, P. ( 19871 in Predation: Direct and Indirect Impacts on Aquatic Communities IKerfoot, W.C. and Sih, A., edsl, pp. 38-54, University Press of New England 7 Miller, T.E. and Kerfoot, W.C. f 1987) in Predation: Direct and Indirect impacts on IKerfoot, WC. and Aquatic Communities Sib, A., edsl, pp. 33-37, University Press of New England 8 Vanni, M.f. ( 1986) Ecology 67, 337-354 9 Schoener, T.W. ( 19831 Am. Nat. 122, 240-285 IO Kerfoot, W.C. and Sih, A. ( 19871 in Predation: Direct and indirect Impacts on (Kerfoot, W.C. and Aquatic Communities Sih, A., edsl, pp. vii-viii, University Press of New England I I Karban, R. and Myers, J.H t 1989) Annu. Rev. Ecol. Syst. 20. 33 l-348 I2 Strauss, S.Y. (I991 I Ecology 72. 543-558 I3 Alford, R.A. and Wilbur, H.M. #I 1985) Ecology 66, 1097-I 105 I4 Kneidel, K.A. (19831 Ecol. Entomol. 8, 163-169 15 Wilson, D.S. II9801 The Natural Selection of Populations and Communities Benjamin/ Cummings I6 Toft, C.A. ( 19861 in Communit,y Ecology (Diamond, 1. and Case, T.I.. edsl, pp. 445-464, Harper & Row I7 Turner, A.M. and Mittlebach. G.E ( 19901 Ecology71, 2241-2254 II? Schoener, T.W. (I9881 Proc. Nat/ Acad. Sci. USA 84, 7992-7995 I9 Schoener, T.W. II9881 Oikos 53, 253-266 44, 20 Goodnight, C.J. II9901 Evolution 1614-1624 21 Paine, R.T. (1980) /. Anim. Eco,l. 49, 667-685 22 Wilson, D.S. (19861 in Community Ecology (Diamond, j. and Case, T.]., eds), pp. 437-445, Harper & Row 23 Higashi, M. and Patten, B.C. (19891 Am. Nat. 133, 288-302 24 Vandermeer, I. t 1980) Am rVat. I 16. 44 l-448 25 Paine, R.T. (1988) Ecology69, 1648-1654 26 Paine, R.T ( 1966) Am. Nat 100. 65-75 209

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27 Fowler, N. 11981 I/. Ecol.69, 843-854 28 Duggins, D.O. 11981) Oecologia 48, 157-163 29 Brown, I.H. and Heske, E.I. i 19901 Science 250, 1705-l 707 30 Menge, B.A., Lubchenco, I. and Ashkenas, L.R. ( 1986) /. Exp. Mar. Bio/. Eco/. 100, 225-269 31 May, R.M. ( 1973) Stability and Complexiry in Model Ecosystems, Princeton University Press 32 Pimm, S.L. (19821 Food Webs, Chapman & Hall 33 Briand, F. and Cohen, I.E. ( 19871 Science 238,956-960

34 Kerfoot, W.C. in Predation: Direct and Indirect Impacts on Aquatic Communities (Kerfoot, W.C. and Sih, A., edsl, pp. 57-70, University Press of New England 35 Schoener, T.W. (1989) Ecology 70, 1559-1589 36 Mills, E.L., Forney, I.L. and Wagner, K.I. ( 1987) in Predation: Direct and /direct impacts on Aquatic Communities (Kerfoot, WC. and Sih, A., eds), pp. Il8-131, University Press of New England 37 Lawton, I.H. and Warren, P.H. 11988) Trends &co/. ,901. 3, 242-245 38 Kerfoot, W.C. and De Angelis, D.L. i 19891 Trends Ecol. Evol. 4, 167-l 7 I

39 Wilbur, H.M. (I9721 Ecology 53, 3-21 Direct 40 Threlkeld, S.T. (1987) in Predation: and Indirect Impacts on Aquatic Communities (Kerfoot, W.C. and Sih, A., edsl, pp. 161-173, University Press of New England 41 Bender, E.A., Case, T.I. and Gilpin. M.E. (1984) Ecology65, l-13 42 Dungan, M.L. ( 19871 Am. Nat. 127, 292-316 43 johnson, R.K., Bostrom, B. and Van de Bund, W. II9891 Limnol. Oceanogr. 34, 992-1003 44 Loehle, C. ( 19901 Ecology, 71, 2382-2386 45 Patten, B.C. (19901 Ecology, 71, 23862393 46 Pilette, R. ( 19891 Am. Nat. 133, 303-307

BirdFlightand OptimalMigration
Thomas Alerstam
storage capacity, speed and range, can be derived. Research into vertebrate flight is developing rapidly, as reviewed extensively in two important recent books, by Pennycuick and by Norberg2. Theoretical predictions about bird flight are tested at many levels: investigating the aerodynamic properties of birds bodies and wings, analysing the vortex wakes of flying birds, measuring flight metabolism or registering speed, climbing capacity, glide ratio and range, etc. These efforts provide fascinating insights into the principles and conditions of bird flight, with important bearing on the ecology and evolution of bird migration. Theory of bird flight A comprehensive theory of bird flight was first developed 20 years ago by Pennycuick3 on the basis of classical aerodynamic theory. Tucker4 put forward an expanded version of the theory, and a closely related interpretation was presented by Creenewalt5. Pennycuick has developed further the basic theoretical frameworkl,6, taking care to shape the theory so as to provide a great amount of predictive power about flight behaviour, which can be used by field ecologists7. Of key importance in this theory is the power curve for flapping flight (the power required to fly as a function of speed). Pennycuick s approach permits the power curve to be estimated for a given bird on the basis of its body mass and wing span only. Enclosed with Pennycuick recent book is a pres

TCle developnzent tCle mechanical of and aerodynamical theory of bird flight has greatly stimulated research at widely different levels in tCrefield of bird nzovernent. Recentwork has shown that the drag of bird bodies less tkan was previouslyassunzed. is Furthermore, the structure and circulation of wingtip vortices in the wake offlying birds have been revealed, with implications for estimating flight performance on the basis of vortex theory. Predictions about optimal speed and flight behaviour have been successfully tested 6y field studies using optical and radar registration. Flight theory also allows predictions about optimal fuel deposition rules for migrating Girds. Research about bird flight, with the dynamic interplay between theoretical development and empirical work in biophysics, physiology and ecology, representsa fine example of a highly successful use of the optimality approach in biology.

Flapping flight is by far the most expensive method of locomotion, considering energy costs per unit of time. In the light of this, the longdistance migrations of multitudes of birds may seem paradoxical. But flight brings a distinct advantage in comparison with movement on the ground or in the sea: speed. The normal speed by flight is often more than ten times higher than that by sustained running or swimming, for similar-sized animals. As a conseThomas Aierstam is at the Dept of Ecology, Animal Ecology, University of Lund, Ecology Building, S-223 62 Lund, Sweden.

quence, the cost of transport, i.e. the energy cost per unit of distance travelled, is reasonably low for flight, clearly lower than for running and only a bit higher than for swimming. The combination of speed and reasonable transport costs enables birds to make the most astounding seasonal journeys. Our knowledge of bird migration is steadily growing, thanks to the accumulation of ringing results and to field, radar and radiotelemetry studies (which nowadays also involve satellite tracking). But although a lot is known about birds feats of travelling across the globe, our understanding of the evolutionary possibilities and limitations in bird migration, and of the adaptive significance of migratory habits, is comparatively meagre. Which is of highest priority to migrants in different situations: to minimize the energy costs of transport, or to travel as fast as possible? For migrating birds, hampered in their flight performance because of heavy fuel burdens, predation risk may be highly significant. It may be that, in many cases, birds are primarily adapted to minimize the risk of mortality from predation during migration. The aerodynamical theory of bird flight must provide the foundation for analysing these and similar open questions about optimization criteria in bird migration. On the basis of this theory, crucial predictions about the performance of travelling birds, their energy costs, fuel

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