A Theory of Habitat Selection Michael L. Rosenzweig Ecology, Volume 62, Issue 2 (Apr., 1981), 327-335. Stable URL: btp//links jstor.org/sici?sic!=0012-9658% 28 198 104% 2962%3A2%3C327%3AATOHS%3E2.0,CO%3B2-B ‘Your use of the ISTOR archive indicates your acceptance of JSTOR’s Terms and Conditions of Use, available at hhup:/www.jstor org/about/terms.html. JSTOR’s Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the sereen or printed page of such transmission. Ecology is published by The Ecological Society of America. Please contact the publisher for further permissions regarding the use of this work, Publisher contact information may be obtained at hitp:/wwwjstor.org/journalsiesa.htnl, Ecology ©1981 The Ecological Society of America ISTOR and the ISTOR logo are trademarks of ISTOR, and are Registered in the U.S. Patent and Trademark Office For more information on ISTOR contact jstor-info@umich.edv, ©2002 JSTOR bupslwwww jstor.org/ ‘Sun Oct 27 07:08:28 2002‘Sri ey ne Bnagear Seay of Ane A THEORY OF HABITAT SELECTION? MICHAEL L. RoseNzweIe Department of Ecology and Evolutionary Biology, University of Arizona, Tueson, Arizona 85721 USA Abstract. A graphical theory of habitat selection is built in steps. The theory treats two species, in an environment with two usable patch types in matrix of unusable space. The fst step assumes habitat selection is density independent and free of search costs. The second assumes Jensty nde pendence, and the third assumes nether. The fist two steps produce resus already known from farier theories. The third, however, requires a new analytical device, the Isoleg, which Ts line in 1 (wo-dimensionalstae space of the two species’ densities. An isoleg is a set of points in such & Sensity space, such that On one side ofthe set, individuals of species optimize their foraging by being siriet habitat selectors, whereas on the other sie, they do so by using atleast a it of poorer patch. The population dynamics of the competitors s analyzed using ther isolegs. The isolgs allow Us to deduce that the zero isoctines of the species are warped into nodlineat forms capable of pro ‘ducing competitive coexistence. Its shown that at the equlbrium point of this coexistence, there may be no overt competition remaining. The difficulties this presents to the eld investigator ate mentioned, and 1 modified definition of interspecific competition is suggested Key words fof another. Most have considered only the optimal choice which should be made by an individual (e.¢., MacArthur and Pianka 1966, Rosenzweig 1974, Char: ‘nov 1976), or allowed that choice tobe influenced only by intraspecific competition (Fretwell and Lucas 1970). Ina major advance, Lawlor and Maynard Smith (1976) consider optimal habitat selection as a function of population densities. Their model, however, as- sumes that habitat selection is free of costs (i.e., the ‘ime and energy required to search for preferred hab- itats), an appropriate assumption in only a restricted sel of cases, Also, their model affords us a view of their system only near equilibrium, which can make it difficult to predict the effects of perturbations (nat- ural oF experimental) ‘This paper reviews some previous contributions (Fretwell and Lucas 1970, Rosenzweig 1974, 1979, Charnoy 1976) and sets about to detail the derivation ‘of a model for a simple system in which densities, behaviors and costs of habitat selection are all vari ables. Many of its conclusions are unsurprising, but some challenge widely held beliefs about mature sys- tems, As required, the model gives the same results 8 those of previously published models (Rosenzweig. 1974, Lawlor and Maynard Smith 1976, Charnov 1976) When its assumptions are collapsed to match previous Let us begin by considering density-independent, cost-free habitat selection. Then I shall add search costs to the model and use it to discuss the cireum- ' Manuscript received 6 July 1978: revised 1 April 1980: accepted 4 June 1980, competition; foraging theory; habitat selection. stances under which the habitat selector is favored by natural selection. Finally, I shall introduce intra~ and interspecific competition and note how habitat selec- tion reduces competitive alphas, sometimes to zero, tnd warps competitive isoclines So that the probability ‘of coexistence is increased. CONVENTIONS AND DEFINITIONS Let us assume the existence of an environment with two types of useful habitat patches in a matrix of non- utilizable space. The matrix may be an empty set. Let us further assume that any change in phenotype which improves an individual's fitness in one of the patch Iypes simultaneously reduces its fitness in the other, In other words, we assume that itis possible to define the coordinate axes of some fitness set (Levins 1962) such that the fitness set's border has negative slope along that part of it which connects the phenotype ‘with highest fitness on the ordinate to the phenotype with highest fitness on the abscissa, Fig. 1. Negative slope simply means tradeoff Discussion about specialization in ecology has been hampered by a certain degree of imprecision regarding some important terms: generalist, specialist, oppor- nist, coarse-grained, etc. Two types of distinctions, here in such terms. First, the terms refer to the abi ities of phenotypes. As such, they reflect fitness. Sec- ‘ond, the terms refer to the behavior of phenotypes, ‘These distinctions have been allowed to overlap in ecology. The resulting ambiguity may be profitably climinated by defining, first, a set of terms restricted, to ability, and, second, a set restricted to behavior fence, every generalist lies on the line ‘of slope one wi sth fitness set Conversely,ws MICHAEI Tine of slope one are generalists. Those along the double line fate the phenotypes from which natural selection must select ‘optima they fall slong the edge of nepative slope which con necis the phenotype with highest In We to the one with high- estin Wy and the fitness set. These two definitions serve to pro- vide a gradient of specialization, and even though some of the degrees of specialization may be absent from a given population (e.g. the generalist itsel, the gradient will be a useful concept in developing the theory The b follows. the organis fe in proportions different from those it encounters in the environment. It can be very difficult to discriminate pickiness from opportun- ism in the field, Cost-FREE DENSITY-INDEPENDENT Haprrat SELECTION This topic is treated in Rosenzweig (1974), but the reader may find the following review of some conve- ‘A collection of more than one fine-grained pheno: type in a two-patch environment is not stable. Natural selection will eventually result in only one phenotype remaining (Levins 1962). The successful phenotype ‘depends on the proportion, Po = AcllAg + Aus where Ag is the amount of one patch and Ay the ‘amount ofthe other. As P,, declines from one to zero, the best phenotype changes (gradually or in discrete jumps) from the one which has the highest fitness in, patch G to the one which has the highest in patch B. Sometimes an intermediate value of P,, will favor the best of the generalist phenotypes. In order to ex. LL. ROSENZWEL Ecology, Vol 62, No.2 Phenotype whose fic news fa const st Phenotypes with positive slopes are patch G specaliss. Those with negative sh fre patch # specialists, As Pp grows Irom 220 Phenotype changes from the extreme B specialist co 2 fxtreme one, tothe generalist, 10a G speci hi {o the most extreme G specialist. Simultaneously, the actu finess rst declines to the level of the generalist, then rises tolthat ofthe extreme G specialist, amine habitat selection, one needs to know how the ‘mean absolute fitness of such an optimum compares With the mean absolute fitnesses of other optima achieved at other values of P. The answer is quickly discovered. ‘The generalist’s fitness does not vary with Po. Hence, any Pz that causes the generalist to be re- placed by any specialist must improve average fitness in the population. Similarly, replacement of a less ex- treme patch i specialist by a more extreme one also increases fitness. The latter follows from Levins’ (1962) demonstration that any such replacement could ‘be brought about only by increasing the proportion of the patch type i in which the two specialists excel ‘Thus the less extreme specialist's fitness must in- crease, yet its replaced by a more extreme specialist (whose fitness must therefore increase even more). These considerations allow us to construct Fig. 2 which shows that equilibrial fitness in a population of fine-grained alternatives is a U-shaped function of with a minimum at the value(s) of P, which favors the ‘generalist. (Note that this is not necessarily Py, = 0.5 nor is the U-shaped function necessarily smooth or symmetrical.) ‘The U-shape immediately resolves the question of habitat selection if population density is not consid- cred as a variable, and if organisms, by selecting hab itat, can adjust the value of Pa to its extremes. All phenotypes specialized for habitat G have optimal ft 1.00, All those specialized for habitat B wal fitness if Py is 0.00April 1981 Gf | Fro, 3 Bian a te dion he ED ‘an invigual timed from the poin when encoun patch B, through an encounter with patch G, tthe instant it encounters another patch B. Note that it may encounter ‘ny number of B's before the G, and any: number of G's before the next 8. While in patch type B, the time it spends Foraging is timed and tis f,. While in patch type Othe time itspends foraging fs timed and is The remainder fst HaBirat SELECTION WiTHt 4 SEARCHING Cost ‘This topic has been treated independently by myselt (Rosenzweig 1974) and Charnov (1976), Considering the infinite array of possible models, itis significant that our models are identical and reach identical con- clusions. ‘The functions of this section are to review and update the model and to show that the two treat= ments of it, when reduced to common symbols, are the same. ‘Charnov’s treatment, however, is preferable, be- ccause he deduces that an average fitness does in Fact exist, while I had only assumed so, He also specifies the amount of time an organism should stay ina patch Although we know that the ultimate common cur rency of biological transactions is fitness, Charnov {and others) have employed energy as currency. This may be close to the ideal in many cases, and it cer= tainly encourages field testing. In fact, one may trans- late what follows into energy currency with the un- {derstanding that it may not always be appropriate; the ‘models may be valid in units of fitness currency even though they are not always valid when formulated in ‘energy currency. Those contemplating field tests of the theory in energy currency should take care to en- sure that their subjects are in fact subject to energetic constraints during foraging activities. Assume that a forager exhibits three activities: searching, actually foraging (after resources are sensed) in patch type B, and actually foraging in patch type G. Let ty be the average time it spends foraging ima single set of B patches (ie., one or more B patches encountered without interruption by an encounter with a G patch). Let r, be the average time it spends foraging in a single set of G patches. (The subscript A THEORY OF HABITAT SELECTION a9 .« is used to denote the variable 1, because this is the variable with respect to which ull differentiation will bbe conducted, and this is the variable which we as- sume to be alterable by changes in foraging behavior.) ‘The time spent searching, t4, is the time the forager spends in going from the initial border of the first patch B of a set t0 the initial border of the first patch B in a second set (having encountered at least one patch, G in the meantime) minus [ty + t] (Fig. 3). Let In Z be the forager’s average (negative) fitness ue to metabolism, predation, etc.) while searching; In W, will be its average net fitness while foraging in 2B; and In W, will be its average net fitness while for- aging in G. Overall average fitness is therefore + toln Wy + ie + te (ay The animal also incurs costs while foraging. Let us assume those costs are incurred at the same rate as juring searching, InZ. Then the net profit rates, In W, and In W,, may be expressed as sums In W, = In Wy + nz In W, = In We + InZ and Eq. 1a becomes nw (ab) Ifthe costs are really the same while searching and foraging, In W, and In W are rates of gross profits But costs need not be the Same for the model (Eq. 1b) to hold. If they are not, then In Wy and In We are {gross profit rates adjusted for the difference in costs between foraging in a patch and searching. For ex- ample, if costs while foraging in G exceed InZ by a rate In D, then In Wis the rate of gross profit minus In. ‘The definition of the parameters in Eq. 1 utilizes the holistic or sufficient variables approach (Levins 1966) Each variable subsumes a considerable amount of in- formation about the environment and the organisms which are exploiting it. For example, the 1 values will be alfected by the average size and shape of the patches as well as their abundance of resources. Rath- er than attempt to define and then to measure all the details which go into these variables, and moreover to hhope that enough will also be known about the inter- actions of the details to construct an equation for the variable, the sufficient variables approach simply uti- lizes whatever variables are necessary to model the actual question at hand. However, the holistic method ‘does not result in the omission of the components of each sufficient variable. The value of ty still incotpo- rales the speed of the animal, the difficulty of the ter rain, the renewal rates of the resources, ele. That, in fact, is the whole point; the sufficient variable already includes all its components and does so automatically30 in precisely the right manner although no one yet knows what that manner may be. Now, suppose a phenotype is a patch B specialist ‘Then it should decrease its use of patch G if and only it \ ainw Be In WoL ° Since 1 is absent from the criterion, use of the poorer habitat should either be totaly fine grained or else nonexistent. It depends on the degree of specializa- tion (ln W,/ln W,), the search time (14), and the re- ward for finding the better patch (Le., fq, the time available to exploit it. This result is identical with Charnoy’s (1976), who, showed that a patch should be abandoned once its rate of profit return had fallen below the average for the whole set of patches. (Charnov, following the terminology of economics, terms this the Marginal Value Theorem.) In the symbols of this paper, the average is Sain, nw i +k Hence, for two patches, the rate of return in the poorer is less than the average if and only if o where j represents the better patch. Eq. 5 is, of ‘course, the same result as Eq. 3 In-a given environment, not all phenotypes need to specialize. The jack-of-all-trades has only costs to pay ‘and nothing to gain, The most extreme specialist has most to gain. Which phenotype(s) will succeed? Surprisingly, it takes only a slim edge for the ex- treme specialist to be successful. Rosenzweig (1974) showed that a specialist whose fitness in its special patch is twice that of a generalis’s should replace the latter with virtual certainty regardless of pateh abun- dance and distribution. Even advantages of less than a factor of two will usually be sufficient ‘The criterion for the specialist's success is approx imately In Wy a TW? 1+ leafy + 6.01 o where In Wy isthe generalists fitness. Note that since the variable fraction on the right side grows with fy and declines with t,, this criterion becomes easier t0 Satisfy given the same environmental conditions that make the habitat selection the preferred behavior for the specialist in the first place. However, the criteria (Eqs. 3 and 6) are not identical and so itis possible to MICHAEL L, ROSENZWEIG Ecology, Vol. 62, No.2 imagine @ generalist outcompeting a coarse-grained specialist. In fact, it is easy to show that for this 10 occur, itis necessary and sufficient for dM +2 In Wy ~ Tn Wyte ne i) IN Wey +h) o DENSITY-DEPENDENT HABITAT SELECTION Rosenzweig (1979) has considered in general terms the consequences of habitat selection in a two-species ‘competitive system. In the remainder of the paper, I investigate and develop the arguments in support of ‘one of the eases discussed in that paper. It is the case whose basic assumption is the same as that of this paper: more effective utilization of either habitat re- duires reduction of effectiveness at using the other. ‘Assume that resources per capita decline as popu lation density increases. If habitat selection has a cost, the need to be coarse grained clearly depends on those sources because In W, depends on the resource density of patch i Noting such relationships, Fretwell and Lucas (1970) predicted that habitat selection must be aban- doned after population density grows beyond some critical point. They determined this point to be the density such that the specialist's fitness was the same in both patch types. Fretwell and Lucas ought to be correct for the cost- free environment implicit in their theory. And they are ‘qualitatively correct even for other environments. But their quantitative criterion is too conservative if hab- itat selection has a cost (L.e., fy > 0). With ty > 0, habitat selection should be abandoned even before density is high enough to equalize fitnesses in the two patches. It should be abandoned as soon as fitness, In We, has fallen far enough so that In Wy M, for all 7 > j, where {and j are values of NN, and My is the isoleg density of M at N= k. This ives us the segment of the M isoleg labelled | in Fig 4, We must now turn to the NV isoleg because at N= No, we cannot yet specify whether N is coarse grained in'the vicinity of the M isoles. Now we imagine an infinitesimal M = ¢, At (e, 0) -M makes a habitat choice. As before, raising N only increases this tendency since N < N,, Hence, in the vicinity of (, Ng), Eq, 9 does not hold for species N. N must be raised to achieve the isoleg point (e, N,). We proceed as with the M isoleg with the same result which appears as line segment 2 on Fig, 4). Now we can return to the M isoleg and use the same arguments to obtain line segment 3. We next return to, 'N and get segment 4. This process can continue ini- nitely and so both isolegs have been shown to have positive slope Of course Fig. 4 should not be taken to indicate the second derivatives of the isolegs; about these, I have ‘no information yet. I also do not know whether the isolegs converge, diverge or remain parallel. Should they converge and actually meet, they may begin to slope negatively. However, the presence or absence ‘of such convergence, or even of regions of negative isoleg slope, are beyond the scope of this paper be- cause they do not influence its qualitative conclusions. Consequently, hereinafter the isolegs are drawn as straight parallel lines, Isoclines of fine-grained competitors In order to study the influence of habitat selection "upon two-species competitive systems, we now turn ‘our attention toa brief review of competitive isoclines. Afterwards, we shall overlay the isoclines upon isolegs and determine how the latter warp the former. Competition is often studied by means of Lotka- Volterra equations, linear differential equations whose solutions ae still unknown. For didactic purposes, the zero isoclines of these equations are often plotted in the state space of two species, That is, in a two-co- ‘ordinate system whose axes are the species densities, (M, N), one plots the set of points (M,, N)) such that Midt or dNidt is zero. These are termed the M and 1N iisoclines respectively. Slobodkin (1961) plotted these linear isoctines which he then used for illustrat- ing the meaning of things like the competition coeff- cient (a) \ MacArthur (1958) implies that this procedure can be profitably reversed. He plots the isoclines first and ‘makes deductions from them. This allows one to han: dle differential equation systems which not only re-am 5, The four sypes of competitive isocne systems, In each, the N isocline emanates from Ky, the M isocline ‘emanates from Ky. Stippled areas delimit population com- positions at which dNidi > 0; in hatched areas, dMidt > 0. Vectors indicate the direction in which population compos tions will move. In part a, NV always excludes M. In part b. 1M always excludes. in parte, there is a saddle point caused by some allelopathogen oF usurpation, and that Species wins which is Ast common. Only in pat do the main unsolved, but also remain unwritten. From qual- itative properties of the system, one goes directly to the qualitative properties of its solution (Rosenzweig 1973), ‘There are four basic competition graphs (Fig. 5). ‘Three of them are possible with fine-grained compet- itors; either of the two competitors can outcompete the other (Fig. Sa or b) deterministically. Or, each can produce an allelopathogen (chemical or behavioral), So that the one which first becomes abundant is the ‘winner. The fourth graph, coexistence (Fig. Sd), is not possible without some form of habitat or resource al- Jocation in space or time. Isoclines of habitat selectors ‘The slopes of the isoclines are determined by the intensity ofthe competitive interaction; the more com- petition, the more the isoclines approach the slope of the axis of their own species. The intensity of com- petition is measured by coefficients, a, the interactive effect of species j on species 7 where uy = (In WianM(@ In WAN). LN; is the density of / and In Wis its per capita rate of increase 1) MICHAEL L, ROSENZWEIG cology, Vol. 62, No.2 % Fic. 6. Warping isclines across an isoleg field: see text, for derivation. Suipling and shading a in Fig. Sd, but omit- {ed for clarty. The vectors show thatthe system converges fon the coexistence equilibrium shere (in this case). ay = 0 Often, ecologists try to estimate alphas by means of resource utilization overlaps (e.g., MacArthur and Levins 1967, May and MacArthur 1972, Roughgarden 1074), The fact that these are only estimates of Eq. 10 is emphasized by their sometime failure under field conditions (Dayton 1973, Schroder and Rosenzweig. 1975). Nevertheless, it should tend to be true that two species with similar ecological requirements which forage in precisely the same habitats have higher al- phas than an otherwise similar pair which selects somewhat different habitats. We shall assume this to be so, and use it to determine the effect of habitat selection on isoctine slope. ‘We can now track an isocline across an exemplary pair of isolegs (Fig. 6). We begin at a point arbitrarily lose £0 (Ky, 0). Let this point lie to the right of My 0 that a reasonably complex and interesting example may be derived and used for illustrative purposes. Near (Ky, 0}, M is not habitat selecting, but N is. ‘Thus there is some overlap, hence competition. The isocline slopes negatively. ‘As the M isoleg is approached, M's behavior is, probably optimized at smaller and smaller values of ‘ithe time it spends in N's special habitat declines. ‘Thus the M isocline would be concave upwards. When it reaches the isoleg, both species are habitat selecting, so the isocline becomes vertical Because there is complete habitat separation be- tween the isolegs, the M isocline remains vertical until it crosses the N isoleg. At this point, N begins 0 invade M's special habitat, and competition reasserts itself. As the isocline departs more and more from the 1V isoleg, its slope probably increases, soit should be ‘convex upward, Beginning near the point (O, Ky), we may now trace the N isocline in the same manner. The conclusions are the same except that as competition declines, the‘April 1981 ky. N ° Ke M Fio. 7. Habitat selection promotes the possibilty of co- existence, The straight soclines are for fine-grained system, whereas habitat selection produces the curved isoclines. becomes extinct in the fine-grained alternative, but coexists with M in the coarse-grained one 1 iisocline’s slope increases. Thus it is horizontal where M's is vertical, concave where M's is convex, and convex where M's is concave. Discussion In one respect, the model produces a conventional conclusion. Habitat selection is indeed a force for achieving competitive coexistence. If one begins with ‘fine-grained system in which M outcompetes Nand then allows for habitat selection, one will quickly see this. The values Ky and Ky are fixed, with or without the habitat selection, because at saturation densities, each species is an opportunist in the absence of the other. However, the habitat selection, as we have seen, steepens the M isocline, and reduces the abso- lute value of the slope of the NV isoctine. This cannot but increase the probability that the two will intersect and form a stable node (Fig. 7). ‘There are numerous ways to combine isoclines and isolegs. It is not my intention for this paper to become ‘catalog of them. The important features of Fig, 6 and the reason it was chosen as an example is that in it, the isoclines cross at a place where both species are extreme habitat selectors and a, = 0. Standard dy- ‘namic analysis of the equilibrium so formed reveals a stable node (see the vectors drawn in Fig. 6). Hence the net effect of natural selection in this case is to produce an a of zero despite the facts that, (1) competition is one of the principal formative forces in the system; (2) because of the competition, each species’ steady- state population is well below its carrying capacity (K). ‘This obviously creates some serious problems for the student of population interactions. In the first A THEORY OF HABITAT SELECTION a3 place, there is the minor semantic problem of how to define competition. Previously, one might have said there was competition between two species if and only it a In WylaN <0; a In Wola ay where both are not permitted to be zero (Rosenzweig 1970). Now it appears we must adopt the following definition: Two species compete if and only ifthe sum of their equilibrium densities is less than the sum of their carrying capacities, at least partially because ‘each depresses the other's net per capita reproductive rate, at least at some density combinations, Repro- ductive rate depression is merely a translation of In- equality 11 above. So the novelty in this definition is the recognition that the relationship which brings about the interaction does not have to prevail at every possible density pair, but does have to produce an overall depression of densities. ‘Such a definition of competition, however correct, is little consolation to the field ecologist. Shoud he try to measure “niche overlap” in the vicinity of the sta- ble node in Fig. 6, it will be zero, Should he try a perturbation experiment, he will again come up with ay = 0. That, in fact, is probably just what happened to Schroder and Rosenzweig (1975) ‘The theoretician, too, should be troubled by a stable alpha of zero between two strong competitors. MacArthur and Levins (1967) coined the concept of a “limiting similarity” between two competitors, the idea being that two species may have too high an alpha to coexist. They encouraged us to believe, and many ‘others have agreed, that communities ought to be packed with competitors until close competitors reach limiting similarity. The mechanism by which this is to be achieved is random generation of new species cou- pled with extinction of those too close together, May ‘and MacArthur (1972) estimated the value of limiting similarity t0 be 0.56, However, as MacArthur and Levins (1967) also pointed out, niche positions are subject to natural se- lection. In fact, what has been done in this paper is to ‘examine the consequence of natural selection’s action fon a habitat selection system. The result was shown to include the possibility of a, = 0, Since natural se- lection will separate any randomly generated new species, and since all the available resources of the ‘two patches will be fully utilized when it finishes, and since it appears that quite a plausible end result is ay = 0, itis hard to accept the significance of limiting simitarity as the end product of evolutionary matura- tion, ‘A necessary property of a scientific theory is test- ability, Testability means falsifiabiity (Medawar 1967). The present theory makes predictions which are of the sort I believe t0 be most readily testable in ecology, but it also points out a caveat for those anx-m ious to reject competition as an influential force in the ‘evolution of community structure. Let us consider the latter first ‘Suppose, by virtue of the kinds of considerations in this theory, two species find themselves coexisting with ain «sof zero. One can use the definition of com petition herein proposed to discover if their equilibria are below their carrying capacities. This probably re- quires grand perturbations of density I their equilibria are less, there is no problem, The species fit the definition of competition. But suppose they do not? What is one to conclude? The conclusion must be that there is ar present n0, ‘competition. Moreover, if one equilibrium is higher ‘and the other lower, then we can agree to have ob- served an exploitation. And if both are higher than their K°s, itis @ mutualism. But what if there seems tobe no interaction at all? What if both equilibria equal their K's? ‘We cannot change the conclusion that there is no competition, But we may not want to reject the pos- sibility that competition was instrumental in producing the species relationships which we observe. For ex- ‘ample, suppose a system is so stable that natural per~ turbations virtually never cause densities to cross iso- legs. Is it not possible under such stable conditions that natural selection will have eliminated the vestigial ‘behavioral flexibility which allows utilization of the alternative habitats? If so, even the most dramatic per turbation will not sueceed in demonstrating that, were it not for the second species, the first would use a larger range of habitats and have a higher density. To succeed in such a case, the perturbation would not only have to be grand, but it would also have to be sustained long enough for the behavioral flexibility 10 re-evolve. That seems so impractical that it seems to me best to conclude that the field biologist has litle hope of conducting experiments on species whose c's are zero that falsify the hypothesis that competition caused them to be that way. At least I cannot now envision such experiments based upon the present the- ory. (On the more optimistic side of the ledger, one may see bright possibilities for field tests. Ecological the- ories which predict precise values for coefficients and parameters are not as practical as physico-chemical theories which do so. There is so much noise in eco- systems that it is always possible to wonder if the measurement failed to fit the theory because of the noise and not because of the inadequacy of the theory. (On the other hand, theories which make qualitative predictions are often too easily fit. Most qualitative predictions simply state the existence and direction of fa pattern (e.g., clutch size should rise with latitude) All too often, even these are discovered before the theory is advanced, and no further predictions from the theory are made to allow the pattern to be tested, MICHAEL L. ROSENZWEIG Ecology, Vol. 62, No.2 ‘The current theory’s predictions are neither as pre cise as quantitative predictions, nor as simple and eas- ily fitas most qualitative ones. The theory predicts the shapes and relationships of seven different regions of fa two-species state space. It predicts the existence of three behaviorally distinguishable regions separated by borders of positive slope. And it predicts three seg- ‘ments each of two different zero isoclines. The shape of cuch of these isoclines is nonlinear and radically dissimilar from any other Ihave ever seen. Moreover, the isoclines are supposed to articulate with the isolews where the former change from infinite or zero slope to some negative slope. 'AIL of these features add up to vulnerability for the theory. Should any fail to prove true (given that the ‘assumptions of the theory are met), the theory would at least need modification and perhaps would need to be rejected AcKNOWLEDOMENTS Drs. 7. Caraco, S. Fretwell, H.R. Pulliam, W. M. Sch: fer, and C. C. Smith made stimolating and valuable com- ‘ments. The continued support ofthe National Seience Foun ation made the research possible. 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