Abstract
In this chapter, we first trace the history of the concept of ecological niche and see how its meanings varied with the search for a theory of ecology. The niche concept has its roots in the Darwinian view of ecosystems that are structured by the struggle for survival and, originally, the niche was perceived as an invariant place within the ecosystem, that would preexist the assembly of the ecosystem. The concept then slipped towards a sense in which the niche, no longer a pre-existing ecosystem structure, eventually became a variable that would in turn have to be explained by the competitive exclusion principle and the coevolution of species. This concept, while more operational from an empirical point of view than the previous one, suffered from an ill-founded definition. A recent refoundation by Chase & Leibold enabled to overcome some of the definitional difficulties.
We then present how, in contemporary ecology, the niche concept is recruited to explain biodiversity and species coexistence patterns. In parallel, neutralist models, by successfully explaining some ecological patterns without resorting to explanations in terms of niche, have questioned the explanatory virtues of the niche concept.
After this presentation, it seems that the fortunes and misfortunes of the niche concept can be seen as a reflection of the difficulties of ecology to give birth to a theory that would be both predictive and explanatory.
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Notes
- 1.
Julve (2005) provides a synthetic list of actors of seemingly ecological ideas since ancient times.
- 2.
For both authors: (1) the ecological equivalents were the rationale for the concept, as an evidence that similar niches existed in different places, (2) the niche was seen as a place that existed independently of its occupant, (3) food was a major component of the niche but the niche was not restricted to food, as it also included the micro-habitat factors and the relationship to predators. However, Elton’s definition being more vague, several species could share the same niche (Griesemer 1992: 235). In addition, Elton explicitly excluded macro-habitat factors, which was not the case for Grinnell. (See Schoener 1989: 86–87 for a detailed discussion of the relationship of these two concepts.)
Griesemer (1992: 235–236) notices that the two concepts are better distinguished with respect to the research programs in which they were inserted, rather than to differences between some of their respective definitions: Grinnell focused on the environment to explain speciation, while Elton focused on the structure of the communities.
- 3.
Schoener (1989: 85), acknowledging Gaffney (1973, here cited as 1975), notices in particular the precedence of Johnson (1910). Johnson used the word in a way similar to Grinnell’s concept: species must occupy different niches in a region, because of the importance of competition in the Darwinian theory. However, Johnson observed that the lady-beetles he studied did not seem to show a clear niche distinction – an observation, Schoener remarks, that was to be repeated many times on arthropods in later studies. Hutchinson (1978: 156), who studied the books available to Grinnell from 1910 to 1914, did not find Johnson’s work in them (Schoener 1989: 85).
Schoener also reports the work of another contemporary, Taylor (1916), who worked with Grinnell, and who also focused on ecological equivalents (Schoener 1989: 84). Taylor however, Schoener notices, rather than imagining that the repetition of local adaptive radiations to similar niches between different locations would lead to convergences, suggested that the same group of organisms would fill the same niche in different geographical areas. Barriers to dispersal could thus prevent some niches to be filled.
- 4.
In their historical introduction, Chase and Leibold (2003: 7–8) give a quick and edifying portrait of such studies in vegetal ecology: “For example, Tansley (1917) performed experiments that showed how plant species competed and coexisted, in a sense vying for shared niche space. Tansley also explicitly contrasted the conditions in which a species could theoretically exist with the actual conditions in which it did exist: ideas generally attributed to Hutchinson (1957) in his discussion of “fundamental” and “realized” niches (…). Salisbury (1929) furthered this distinction and suggested that the similarity in species requirements was strongly related to the intensity of their competition – much the same concept as appears in the more widely appreciated work of Gause (1936)” (referred here as Gause 1934).
- 5.
“… if the species lay claim to the very same “niche”, and are more or less equivalent as concerns the utilization of the medium, then the coefficient α [in Lotka-Volterra’s equations] will approach unity” (Gause 1934: chap. III).
- 6.
“It appears that the properties of the corresponding [Lotka-Volterra] equation of the struggle for existence are such that if one species has any advantage over the other it will inevitably drive it out completely (Chapter III). It must be noted here that it is very difficult to verify these conclusions under natural conditions. (…) There being but a single niche in the conditions of the experiment it is very easy to investigate the course of the displacement of one species by another.” (Gause 1934: chap. V)
- 7.
By contrast, in France, L’Héritier and Teissier (1935), who carried out experiments on the coexistence of two species of Drosophila, came (in agreement with some experimental results of Gause 1934) to the conclusion that “two species sharing the same resource in an environment and using it in an apparently identical way may survive side by side in a state of approximate balance.” (see Gayon and Veuille 2001: 88). On the status of the competitive exclusion principle, seen as an a priori, and therefore irrefutable, principle, see Hardin (1960: 1293).
- 8.
The first formulation of the niche concept by Hutchinson is to be found in a footnote, in a paper in limnology (Hutchinson 1944: 20). Schoener (1989: 91) reports a very similar formulation (in french) in a book by Kostitzin (1935: 43): “Imagine a multi-dimensional symbolic space representing the vital factors: p = pressure, T = temperature, I = illumination, etc.. In this space every living creature at a given time occupies a point, a species may be represented by a set of points.”. Hutchinson (1978: 158, quoted in Schoener 1989: 91) acknowledged having been informed of Kostitzin’s work in the 1940s, without, however, remembering it when formulating his definition in 1944.
- 9.
Note that predation will also be set aside in the development of the neutral theory.
- 10.
The environmental and populational niches are however incommensurable if one holds the view that species make some ecological factors relevant that could not be suspected to be so before observing the species (that is, if species and niches are co-constituted, see e.g. similar views in Drake et al. 2007; Longo et al. 2012).
- 11.
The operational difficulties of Hutchinson’s concept come from the (binary) formalism of the set theory he used. They are already partly mentioned by Hutchinson (1957: 417) and discussed in length by Schoener (1989: 93).
All points of the fundamental niche represent the possibility of indefinite existence while all points outside the fundamental niche represent non-indefinite viability. Now, for the ecologist, the performance of a species cannot be reduced to a binary variable. (I thank François Munoz for an insightful comment on this point.) Despite this simplification, a major difficulty is to empirically determine the environmental states that allow the population to survive, because the viability of a population is difficult to assess – especially in the field. Similarly, it is physically impossible to measure the survival of a population at one point of the environmental values, and less precise measurements are likely to ignore the extent of the impact of competing species on the realized niche. Hutchinson (1978: 159, quoted in Schoener 1989: 93) proposed to use the average values instead, but this would lack both biological relevance (the same average can represent very different biological realities) and relevance for the limiting similarity (the niche width and overlap would not be represented).
Another difficulty concerns the nature of the environmental variables considered: strictly speaking, it is the occurrence of a factor (for example, the frequency of the seeds of a certain size) that is one axis of the niche, and not the measurement of this factor (seed size) (see Hutchinson 1957: 421, fig. 1 shown above: the axes are respectively “temperature” and “size of food”). This is because organisms compete, if any competition, for places in the biotope space, not for places in the niche space. This gets particularly clear if one considers possible biotopes where the places corresponding to the intersection of the two fundamental niches would be non-limiting. As Schoener (1989: 94) puts it: “Hutchinson’s formulation of niche overlap acts as if competing species are placed together in arenas having single values of such niche dimensions as food size or temperature. (…) But real arenas where populations interact are characterized by distributions of values over axes of resource availability, not by single values.”. A similar problem exists with the concept of utilization niche, as it also uses the measurement of a factor and not the measurement of its occurrence (see below).
- 12.
Besides, niche theory was considered as inappropriate or of limited use by some botanists, who insisted on the fact that all autotrophic plants “need light, carbon dioxide, water and the same mineral nutrients” (Grubb 1977: 107) and that a substantial partitioning of these resources seems impossible (but see Sect. 3.4.3). Among them, Grubb pleaded for an extended definition of the niche, including notably the regeneration niche – that is, the way plants colonize the gaps arising in the environment (Grubb 1977: 119). Fagerström and Agren (1979) have used models to show how different regeneration properties (i.e. temporal average and variance, and phenology, of diaspore production) could enable coexistence.
- 13.
- 14.
- 15.
- 16.
These and other models are briefly reviewed in Schoener (1989: 98–99).
- 17.
The word comes from Chase and Leibold (2003: 11).
- 18.
- 19.
- 20.
On adaptation, See Grandcolas, and Downes, this volume (Ed. note)
- 21.
Stress: a factor having a negative impact on the organism and on which the organism has no impact (sensu Chase and Leibold 2003: 26, table 2).
- 22.
See Chase and Leibold (2003: 13–14) and references therein.
- 23.
- 24.
To be precise, Leibold (1995) and Chase and Leibold (2003: 15–61) refer to the union of the requirements of the organism and its impacts: “[the niche is] the joint description of the environmental conditions that allow a species to satisfy its minimum requirements so that the birth rate of a local population is equal to or greater than its death rate along with the set of per capita effects of that species on these environmental conditions” (p. 15). The generalization of the definition to the organism responses seems natural (see e.g. Meszéna et al. 2006).
- 25.
- 26.
See e.g. Chase and Leibold (2003: fig. 2.4 p. 27, p. 44).
- 27.
- 28.
Watt and Hogan (2000: 1427) give the following definition: “Although [the question of what a stem cell is] remains contentious after 30 years of debate (…) the prevailing view is that stem cells are cells with the capacity for unlimited or prolonged self-renewal that can produce at least one type of highly differentiated descendant. Usually, between the stem cell and its terminally differentiated progeny there is an intermediate population of committed progenitors with limited proliferative capacity and restricted differentiation potential, sometimes known as transit amplifying cells.” Laplane (2013) provides a thorough discussion of the stem cell concept.
- 29.
Though the stem cell niche concept has been later claimed to come by analogy with the ecological niche concept (e.g. Powell 2005: 268, see also Papayannopoulou and Scaddeb 2008), it does not seem to have been imported from the ecological literature by Schofield. I thank Lucie Laplane for drawing my attention to this point.
- 30.
It has been shown that tumoral cells can mobilize normal bone marrow cells, have them migrate to particular regions and change the local environment so that it attracts and supports the development of a metastasis (Steeg 2005).
- 31.
- 32.
We briefly discussed this point in Pocheville (2010: chap. III).
- 33.
See Delord, Chap. 25, this volume. (Ed. note)
- 34.
See Meszéna et al. (2006) for an examination of the structural stability (robustness of coexistence against changes of parameters) of models of stable coexistence.
- 35.
Here, we use the word “mechanism” in the – very broad – sense used in ecology: practically any form of generation of a pattern can be considered as a mechanism (e.g. Strong et al. 1984: 5&220, Bell 2000: 606, Hubbell 2001: 114, Leigh 2007: 2087; see the brief discussions in Turner et al. 2001: 53 and McGill et al. 2007: 1001). For example, the intensity of competition in a Lotka-Volterra model can be seen, in our view, as a mechanism for the exclusion of two species, while the consumption of the same resource by two species in a Tilman model can be seen as a mechanism, among other possible mechanisms, for the intensity of competition (Tilman 1982: 6, 1987: 769; Chesson 2000: 345). In this sense we say that a Tilman model is “more mechanistic” than a Lotka-Volterra model (e.g. Chase and Leibold 2003: 13), qualified as “more phenomenological” (see Mikkelson 2005: 561).
- 36.
We draw reader’s attention to the fact that here, fitness is not averaged over time but over all environmental states, e.g. the different values of resource availability (Chesson 2000: 346–7353) or the relative frequency of species (Adler et al. 2007: 96: fig. 1, 97: fig. 2). Last, we also speak of an average fitness in the sense of the per capita growth rate, averaged among individuals within a population.
- 37.
Negative frequency-dependence : most frequent populations are disadvantaged. Negative density-dependence: for each population, the per capita growth rate decreases as density increases. While negative frequency-dependence can emerge from negative density-dependence (e.g., when each species has a specific niche which can support a given maximum density), density-dependence is not sufficient to generate frequency-dependence: each species must, in addition, reduce its own growth more than those of others (Chesson 2000: 348; Adler et al. 2007: 97). (Note that density-dependence is not necessary for frequency-dependence to occur: for instance rock-paper-scissors games can arise without any obvious link to underlying limiting conditions (e.g. Sinervo and Lively 1996).)
- 38.
In neutral theory, fitness equality is defined at the individual level (regardless of the species), which implies equality at the population level (the reverse is not true).
- 39.
We would like to draw once again the reader’s attention to the fact that these stabilizing feedbacks are not sufficient in themselves to ensure the stability of coexistence. Put in the graphical terms of Fig. 26.2 given here, niche partitioning will be expressed as a correlation between zero net growth isoclines and impact vectors, and equalizing mechanisms as a proximity of the intercepts of the zero net growth isoclines (Chase and Leibold 2003: 43).
- 40.
On predation and parasitism see also Chesson (2000: 356–357) and references therein.
- 41.
- 42.
A population is subject to an Allee effect when “the overall individual fitness, or one of its components, is positively related to population size or density” (Courchamp et al. 2008: 4, see also p. 10: box 1.1). This effect can be explained by difficulties in finding breeding partners, or by the need for a group to reach a critical mass to be able to exploit a resource or deal with predation (Courchamp et al. 2008: chap. 2).
- 43.
We draw reader’s attention to the fact that this stabilizing mechanism is different from the niche partitioning with respect to predation exposed above.
- 44.
To be precise, in this case we would speak of niche restriction rather than niche partitioning (e.g. Rohde 2005: 51–52).
- 45.
- 46.
See e.g. the discussion by Looijen (1998: chap. XIII).
- 47.
“I believe that community ecology will have to rethink completely the classical niche-assembly paradigm from first principles.” (Hubbell 2001: 320).
- 48.
For simplicity, we use in this section the term “niche theory” in a broad sense (equivalent to the niche-assembly perspective in Hubbell’s terms, 2001: 8), to mean the corpus of models that are based on the niche concept – and not, in the strict sense, the research program of MacArthur & Levins evoked in Sect. 1.4.
- 49.
To be precise, we already find the idea of neutral variation in Darwin (e.g. 1859: 46): “These facts [an inordinate amount of variation in some genera] seem to be very perplexing, for they seem to show that this kind of variability is independent of the conditions of life. I am inclined to suspect that we see in these polymorphic genera variations in points of structure which are of no service or disservice to the species, and which consequently have not been seized on and rendered definite by natural selection (…)”
- 50.
However, Hutchinson still considered the competitive exclusion principle as a starting point (Hutchinson 1961: 143), envisageing to explain unexpectedly high levels of diversity in functional terms, among others: non-equilibrium competitive dynamics (Hutchinson 1941, cited and deepened in Hutchinson 1961: 138), the mosaic nature of the environment (Hutchinson 1959: 154), and the supposed stability of more complex trophic relationships (Hutchinson 1959: 150).
- 51.
- 52.
Drift: variation in frequency (here, allelic frequency) due to a random sampling effect in the population: the offspring population of alleles represents a (finite) sample of the parental population. In virtue of the law of large numbers, the larger the sample, the more representative it is.
- 53.
On neutrality in population genetics, see Leigh (2007: 2076), and references therein.
- 54.
- 55.
Migration had already been studied in population genetics, but never had a central status as in Hubbell’s theory (Alonso et al. 2006: 452).
- 56.
See also Hubbell (1997).
- 57.
- 58.
It is in particular the case when two species are exactly similar (for instance, if they have exactly the same genes and allelic frequencies as for the functional aspects) and are only inter-sterile: there would be intraspecific, but not interspecific, competition. Hubbell (2006) proposed (without, however, stating it explicitly) such a mechanism to explain the evolution of neutrality at the interspecific level. (A similar result would probably be obtained assuming no limitation on (epi)mutations at the intraspecific level.) Chave (2004: 249) quicky discusses how restrictive the assumption of individual equivalence is.
- 59.
Bell (2000: 613) proposed a different – and compatible – definition: “Even the notion of ecological equivalence is rather vague; I shall take it to refer to a set of species for each member of which no interaction with another member is positive. If community structure is determined to some extent by competition, then at least one interaction for each member is negative; the neutral model is the limiting case in which all interactions are negative and equal.”
- 60.
Neutral theory considers fitness equivalence at the individual level (e.g. Hubbell 2001: 6), which implies fitness equivalence at the population level.
- 61.
On the use of these terms, see e.g. Hubbell (2001: 6, 2005: 166, 2006), and the discussion in Clark (2009: 9). For instance Hubbell’s following statement shows a slippage between demographic and functional equivalence: “These life history trade-offs equalize the per capita relative fitness of species in the community, which set the stage for ecological drift.” (Hubbell 2001: 346, briefly discussed in Alonso et al. 2006: 455, similar statements can be found elsewhere in the literature, see e.g. Kraft et al. 2008: 582: note 11). Notice, however, that a full ecological drift would in addition require the absence of any stabilizing mechanisms (an absence that seems to be implicitly hypothesized by Hubbell 2001: 327–328). The word trade-off itself is ambiguous, as trade-offs can theoretically produce both equalizing and/or stabilizing effects (Chesson 2000: 346–347), be they trophic (e.g. Clark et al. 2003) or life-history trade-offs (e.g. Clark et al. 2004). Chase and Leibold (2003), as for them, seem to use trade-offs (here in niche use) as explanantes of stabilization in their whole book: “That is, Hubbell’s hypothetical species show no niche differences or trade-offs.” (p. 42, note the contrast with Hubbell’s quote above). Clark (2009: 9) shows, using Lotka-Volterra equations, how species can have identical parameters (demographic equivalence) while displaying stable coexistence, in particular if there are trade-offs that entail that each species negatively impacts itself more than it impacts the other (functional differences). (Functional equivalence would in this case be represented by an equivalence of the intra- and inter-specific competition terms for each, and all, species. Notice that, still, it would not imply that species be ecologically equivalent, as Lotka-Volterra parameters can be ecologically multiply realized (see Clark 2009: fig. 1).)
- 62.
That is, complete overlap of responses and impacts to environmental factors in Chase’s and Leibold’s (2003: 23) account. Note that with this concept, two species having exactly the same niche behave neutrally, and the only “competitive exclusion” occuring is mere drift.
- 63.
- 64.
- 65.
- 66.
- 67.
As we have seen, Darwin (1859: 46, quoted above) already aknowledged the possibility of neutral differences in phenotypes; he however supposed that the abundances of species in an ecosystem could not be explained by chance, but by the struggle between kinds: “When we look at the plants and bushes clothing an entangled bank, we are tempted to attribute their proportional numbers and kinds to what we call chance. But how false a view is this! Every one has heard that when an American forest is cut down, a very different vegetation springs up; but it has been observed that the trees now growing on the ancient Indian mounds, in the Southern United States, display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. What a struggle between the several kinds of trees must here have gone on during long centuries, each annually scattering its seeds by the thousand; (…)” (Darwin 1859: 74–75).
- 68.
- 69.
Overyielding: positive correlation between the productivity and the diversity of a community.
- 70.
Hubbell (2006: 1395) argues that he found no evidence for overyielding in the tropical forest on Barro Colorado Island.
- 71.
See Beeravolu et al. (2009: 2607).
- 72.
Munoz et al. (2007) have proposed an approach that relaxes the speciation modalities and do not imply any estimation of the speciation parameter. The estimation of the speciation parameter seems generally highly unreliable, contrary to the estimation of the migration parameter, that seems more robust (on parameter estimation, see also Beeravolu et al. 2009). I thank François Munoz for an insightful comment on this point.
- 73.
Unless explicitly stated, this part draws on the remarkable review by Bell et al. (2006).
- 74.
- 75.
- 76.
- 77.
See e.g. Hubbell (2001: 320–321), or this interview of Hubbell by Baker (2002): “Look, I think the biggest question to come out of the neutral theory is: “Why does it work so well?” I’m as puzzled as the next person. But one idea is these trade-offs.” (Notice that here Hubbell still seeks to explain neutrality in functional terms, while a possibly more neutral explanation would be that environmental variations in space and time are such that the environment is not selective, as for instance with fractal perturbations; a case briefly discussed in Pocheville 2010: 85–86).
- 78.
- 79.
The controversy about SADs draws back to Fisher et al. (1943) and Preston (1948). According to Fisher et al. (1943) the expected number N of species having n individuals in a sample can be described by a log-serie: N = αn /n, where α (a parameter now known as Fisher’s α) is a measure of species diversity. According to Preston (1948), the log-serie lacked the bell-shape he observed in his data on bird abundances, a phenomenon he attributed to the presence of trully rare species that are hardly detectable in small samples (a concept now known as Preston’s veil line). Preston (1948) remarked that, by contrast, a log-normal distribution fitted his data. See Hubbell (2001: 31–37) and McGill et al. (2007: 998–999,1004–1005) for short historical introductions, emphasizing respectively the theoretical and empirical sides.
- 80.
The maximum entropy technique consists in describing the microscopic degrees of freedom of a system (e.g. the species abundances) by the probability distribution that maximizes the Shannon entropy, under a set of macroscopic constraints (such as bounded mean abundance). On entropy maximization in ecology, see also Banavar and Maritan (2007), Banavar et al. (2010), Dewar and Porté (2008) and the controversy between Shipley et al. (2006) and Shipley (2009), and Haegeman and Loreau (2008, 2009). Haegeman and Loreau (2008) provide a nice and critical introduction to the technique.
- 81.
- 82.
- 83.
Note that qualitative patterns (e.g. Bell et al. 2001: 133) could be an insufficient method to detect selective processes.
- 84.
- 85.
See McGill et al. (2006: 1414). Such a question is already mentionned by MacArthur (1972: 21), and is repeated, in a less general form, in Chesson and Huntly (1997: 520), quoted in Hubbell (2001: 9–10). Leigh (2007: 2080) raises, in passing, a similar question. Hubbell (1997: S9) interprets the niche assembly perspective of ecologists (vs the dispersal assembly perspective of biogeographers) as a mark of the different processes occuring on the respective scales of these disciplines. See also the three-levels spatially implicit neutral model of Munoz et al. (2008: 117)
- 86.
A major difficulty of this research program is to separate the effects of the environmental variability (on fitness) from the effects of physical/biological distances (on dispersal), for there is a covariation between environment similarity and distance in natural landscapes: environmental variability tends to increase with the geographic distance, and the biologically perceived distance tends to increase with environmental variability (due to barriers to dispersal for instance – such barriers need not be, of course, purely “neutral”, i.e., equivalent for all species). Fort short discussions of this issue, see Bell (2006: 1382), Chave (2008: 21–23). Borcard et al. (1992, see also Legendre and Legendre 2012) proposed a method to statistically partition environment from distance, implemented in Gilbert and Lechowicz (2004: 7653) who found “strong evidence of niche-structuring but almost no support for neutral predictions” (2004: 7651). Jeliazkov (2013: Chap. III) performed an implementation in a similar vein, finding that the environment explained a major part of the community variation only when it was joined to a spatial component. On dispersal as a non-neutral phenomenon see Clark (2009: 12).
- 87.
In other terms, while the composition of a neutral community does not show any equilibrium nor resilience, it is not the case for the caracteristics of this composition (species number, relative frequencies, etc.).
- 88.
A similar counterargument has been opposed by Hubbell (2001: 330–331) to the conclusions reached by Terborgh et al. (1996) on floodplain forests and Pandolfi (1996) on a paleo-reconstruction of coral reefs. Leigh (2007: 2082) points to the fact that Hubbell’s (2001: 331) and Volkov et al.’s (2004) arguments rely on “the fictitious concept of a panmictic source pool”, a fiction that contrasts with a – desirable – approach studying the long-range correlations produced by local dispersal alone (as hypothesized by Bell et al. 2006: 1382). As another step in the controversy, Dornelas et al. (2006) have shown that Indo-Pacific coral communities exhibit far more variable, and lower on average, community similarities than expected by neutrality.
- 89.
- 90.
E.g. Hubbell (2001: 220), McGill et al. (2005: 16706), Bell et al. (2006: 1379), Gewin (2006: 1309), Daleo et al. (2009: 547). These terms are not new, as in the 1970s Lewontin for instance could write: “Genetic variation is removed from populations by both random and deterministic forces” (Lewontin 1974: 192).
- 91.
See Malaterre & Merlin, Chap. 17, this volume (Ed. note)
- 92.
We mean here by “directionnal” a direction in the composition dynamics (of alleles or species frequencies for instance) or in spatial patterns of distributions. Drift, by contrast, can be considered as a noise: it “explains” to what extent we cannot know the direction. (This, of course, does not hold for parameters that are explananda of neutral theory, such as the number of alleles/species, mentionned in the preceding section.)
- 93.
This notion of epistemic randomness is, to our knowledge, the most common notion of randomness in ecology (e.g. Clark 2009: 10: “First, there is no evidence for stochasticity in nature at observable scales. Stochasticity is an attribute of models”). To be precise, random terms could also be considered to reflect deterministically random phenomena, as in classical physics, or intrinsically random phenomena, as in quantum physics. Other concepts of randomness could be developed for ecology. The distinction between direction/dispersion proposed here holds for epitemic randomness.
- 94.
This explanandum is significant in, for instance, the review by Lavergne et al. (2010).
- 95.
Huneman (2012) questions in the same vein the conception of causation (counterfactual vs statistical) required to make sense of natural selection (by contrast with drift) in evolutionary biology.
- 96.
Neutral theory has not always been perceived as a null hypothesis. Bell (2001: 2418) distinguishes two versions of the theory: “The weak version recognizes that the NCM [neutral community model] is capable of generating patterns that resemble those arising from survey data, without acknowledging that it correctly identifies the underlying mechanism responsible for generating these patterns. The role of the NCM is then restricted to providing the appropriate null hypothesis when evaluating patterns of abundance and diversity. (…) The strong version is that the NCM is so successful precisely because it has correctly identified the principal mechanism underlying patterns of abundance and diversity. This has much more revolutionary consequences, because it involves accepting that neutral theory will provide a new conceptual foundation for community ecology and therefore for its applied arm, conservation biology.”
- 97.
A similar argument would hold for implicitly spatial models (involving limited dispersal without necessarily defining a distance between communities): dispersal, even symmetric, is not “null” regarding the niche.
- 98.
The a priori principle belongs to the same familiy than the strong adaptationist principle, that can be formulated as, for example: “every trait is an adaptation to a selection pressure, even if this pressure is not shown”, or: “it is the fittest who survives, even if fitness is not shown”. (On adaptationism, see Orzack and Sober 2001, in particular the chapter by Godfrey-Smith.)
- 99.
I am indebted to Philippe Huneman for having drawn my attention to this point.
- 100.
The author wish to thank Frédéric Bouchard, Antoine Collin, Régis Ferrière, Jean Gayon, Philippe Huneman, Maël Montévil, Michel Morange, François Munoz, Aurélien Pocheville and Marc Silberstein, whose suggestions enabled to greatly improve previous versions of the manuscript. Sylvie Beaud and Robert Pocheville were of considerable help for the translation of the french version. This work consists in a partial update of a previous work in French (Pocheville 2009), realized while the author was benefiting from a funding from the Frontiers in Life Sciences PhD Program and from the Liliane Bettencourt Doctoral Program. This update was realized while the author was benefiting from a Postdoctoral Fellowship from the Center for Philosophy of Science, University of Pittsburgh.
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Pocheville, A. (2015). The Ecological Niche: History and Recent Controversies. In: Heams, T., Huneman, P., Lecointre, G., Silberstein, M. (eds) Handbook of Evolutionary Thinking in the Sciences. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-9014-7_26
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