3.

_,~~ . __ .... _T~."

1.Toxostoma redi vivum sonomae 2.Toxostomar. reviv ium 4.Toxostoma .rpasadenens e

Specimans examined ... Published records

DISTRIBUTION MAP
lIUSE\.l!\( Of' YUlTI:mu.t't. ~y
1. ,. ..

: J:\CEPTS OF NICHES

10

CHAPTE R 2

"::ONCEPTS OF NICHES

11

12

CHAPTER 2

:ONCEPTS OF NICHES

13

14

CHAPTER 2

CONCEPTS OF NICHES

15

16

CHAPTER 2

~:)NCEPTS OF NICHES

17

sized the "requirements" view (Leibold 1995). However, since "Grinnellian niche" has already been used in applications to geographically oriented situations with nonlinked environmental variables, we retain it throughout this book. Of course, other extremes of niche meaning are possible and important; in particular, as we saw, niche concepts exist that are oriented toward communityecology questions, defined at local scales, and including models of resource consumption and impacts. We will refer to this scale and meaning as "Eltonian nihes." One of the best and most synthetic expositions of modem Eltonian niche theory is that by Chase and Leibold (2003). Ideally, to understand fully the geographic distributions of species, both Grinnellian and Eltonian niche elements are needed. Do EMPTY NICHES EXIST? The preceding discussion about niches as properties of the environment versus niches as properties of the species has led in the past to the question of whether such a thing as "empty niches" can exist (Colwell 1992). Unless rigorous de~nitions of terms are provided, of course, this question remains merely sernantic. Grinnellian niches, defined as subsets of scenopoetic environmental spaces, are entirely different entities in this sense than Eltonian niches, defined in terrns of zero-growth isoclines, impact vectors, and supply points. As such, the answer to the question depends, at the very least, on which of these two meanings is involved. It is clear that-strictly speaking-neither Grinnellian nor Eltonian niches can be operationalized without reference to a particular species. Indeed, in the Grinnellian case, niches represent subsets of elements of the E-space defined in terms of the effects of certain variables on population growth rates of the species in question. In the Eltonian case, on the other hand, it is also only in terrns of a given species in a given community context that resources can be specified, a consumption model proposed, isoclines obtained, impact vectors estimated, and the niche thus identified. In this sense, no "empty" Grinnellian or Eltonian niches can exist. However, it is easy to produce examples of realistic and biologically mean: :-:gful Ii-spaces using variables known to be important to distributions of large : .asses of species, without specific mention of any species in particular. Figure ~.2 is an example in which all combinations of three important bioclimatic :.:iables across the entire Western Hemisphere are presented. The Grinnel.. :n niche (defined in terms of these three variables) of any species living in the -.:nericas will be a subset of that E-space. Hence, in a very real sense, figure ~ displays the domain available for Grinnellian niches in the Americas for the = ~:iables considered; some sectors of this space (even if suitable) may indeed

-.'
'.

..

~ . ,~~~::
.

,~,

-" ., -~. ..

--

18

CHAPTER 2

be unoccupied by a species in the region for a variety of reasons (see the following and chapter 8). In contrast, it is very hard to illustrate a realistic zero-growth isocline Eltonian niche in resource space without reference to a particular species. (Presenting a hypothetical Eltonian niche, however, based only on verbal definitions of niches as roles or functions in a community, is not difficult.) This difference is a consequence of the fact that Grinnellian niches may also be regarded as an attribute (measured through scenopoetic variables, and certainly with heritable components related to the species' physiology) of the area occupied by the species across its geographic range, whereas Eltonian niches are attributes of interactions between a species and its local resources and other species. Hence, while it is intuitive and reasonable to refer to empty Grinnellian niches, it is probably meaningless to discuss empty Eltonian niches (Colwell 1992).
HUTCHINSONIAN IDEAS ApPLIED TO GRINNELLIAN AND ELTONIAN NICHES

Hutchinson (1957) defined two subtypes of his multidimensional niches-the fundamental and the realized. For Hutchinson, the "fundamental niche" was the set of "all the states of the environment which would permit the species S to exist" (Hutchinson 1957). The "realized niche," on the other hand, is the subset of the fundamental niche corresponding to environmental conditions under which species S is a superior competitor and can persist (Hutchinson 1957) in an interacting environment. Chase and Leibold (2003) provided rigorous definitions of Eltonian requirement-impact niches, and found that fundamental and realized niches can be defined in their terms. It is also possible and useful to analyze this dichotomy in relation to Grinnellian niches, as we will see in chapter 3. An important side note, however, is that other sorts of niches not considered by Hutchinson (1957) can be conceptualized for Grinnellian niches defined in terms of E-spaces and considering geographic extents. Hutchinson originally discussed niches in abstract and nonspatial ways, without providing actual examples and mostly ignoring the importance of scale and geography. Later, in his "What Is a Niche?" chapter, Hutchinson (1978) provided real-life examples of niches, but still refrained from outlining how consideration of broad spatial and temporal scales might introduce new possibilities. Jackson and Overpeck (2000), working with scenopoetic environmental spaces and following the main ideas of Hutchinson, defined the fundamental niche as the "subset of the environmental space defined by the n dimensions, consisting of the suite of combinations of variables that permit survival and reproduction of individuals." This definition is perfectly compatible with Hutchinson's ideas; however, it begs the question of how can one estimate "the suite of

CONCEPTS OF NICHES

19

combinations" of favorable environments that composes the fundamental Grinnellian niche. This fundamental niche can, after solving several technical difficulties, be estimated via physiological experiments or biophysical first principles (Sutherst 2003, Kearney and Porter 2004, Crozier and Dwyer 2006, Buckley 2008). The fundamental niche therefore is an expression ofthe physiology and behavior of an organism, and its definition can be obtained independently of the localities where a species is observed. An important contribution to Grinnellian niche theory is the realization by Jackson and Overpeck (2000) that the fundamental niche may include combinations of environmental variables currently missing in the existing E-space, which is constrained by geography, and changes continuously across evolutionary time periods (Manning et al. 2009). As a consequence, E-space may include large regions of biologically sensible, but currently lacking, combinations of variables. For example, in figure 2.2, we can observe a large gap corresponding to high precipitation and intermediate temperatures-areas holding these conditions simply do not presently occur in the Western Hemispher~. However, nothing is absurd in considering that a species may have physiological tolerances that permit maintenance of populations under some of these nonexistent environmental combinations. Consider, for example, species distributed in archipelagos of small islands with limited sets of environments represented, even though the species' distributional possibilities are much broader. Therefore, it makes sense to define a "potential niche" (Jackson and Overpeck 2000) that is quite simply the intersection of the existing E-space (or the "existing environmental space") with the fundamental niche-in other words, the portion of the fundamental niche that actually exists somewhere in the study region at the time of analysis. As we will discuss later, the term "potential" is somewhat .mfortunate in this context, but the concept is very useful, so in chapter 3 we rechristen this concept as the scenopoetic existing fundamental niche. Also, as discussed in chapter 3, other perfectly sensible Grinnellian niches not considered by Hutchinson can be defined by considering dispersal and movement in spatially explicit analyses.

ESTIMATING GRINNELLIAN NICHES: PRACTICALITIES ~ey to our exposition of many ideas in this book is a relationship between en.ronmental and geographic spaces. By defining a geographic space (G-space), ~~ a given resolution, and at a particular point in time, and by intersecting that :.:ea with digital data layers of environmental data, it is possible to extract sub: ets of the existing E-space that correspond to different regions of G-space.

5.

7,500 7,000 6,500 6,000 5,500

0 0

a a

E
.

5,000 c 4,500 0

" "

.'3 4,000 '0. 'o 3,500 l!! Q . 3,000

'"

::l

2,500

-c 2,000
1,500 1,000 500 0
I I

-15

-10

-5

10

15

20

25

Mean t emper ature (OC)

=ONCEPTS OF NICHES

21

-=-onversely, GIS operations can be used to map in G-space all the regions that .orrespond to a given subset of E-space. Note that, for every point in G-space, .me and only one point exists in E-space, whereas for each point in E-space more than one point may exist in G-space. This dual-space correspondence is .llustrated in figure 2.3. E-space is changing all the time (Jackson and Over?eck 2000) according to the dynamics of global geology, climate, and physical environment in general. The dramatic recent improvements in availability of data have enabled the work of pioneers like Grinnell, Hutchinson, and Austin to develop into a thriving area of macroecology and biogeography. In particular, as mentioned earlier this field includes what has been termed "species distribution modeling" (SDM; Guisan and Zimmermann 2000, Hirzel et al. 2002, Araujo and Guisan 2006), as well as the related (but by no means equivalent) endeavor named "ecological niche modeling" (ENM; Sober6n and Peterson 2005, Soberon 2007). These fields-the subject of this book=-consist of application of niche theory to ques.ions about the observed and potential spatial distributions of species in t~e oast, present, and future. In a very real sense, the availability of large quantities of data, technological developments like GIS, and several computational tools are enabling a multitude of applications that are not only of biological imporranee, but also can often be of extreme practical utility.

SUMMARY The general idea of "niche" refers to the ecological conditions that a species requires to maintain populations in a given region, together with the impacts .hat the species has on its resources, other interacting species, habitat, and environment. The different emphases that various schools of thought have placed on different components of this central idea have led to a variety of particular niche concepts. For understanding of geographic distributions of species, two .nterpretations of the idea are important. The first, with roots in the work of Joseph Grinnell, emphasizes noninteractive unlinked variables measured mostly

:=IGURE 2.3.

Correspondence of geographic and environmental spaces, showing ariation in mean monthly temperature and annual precipitation across the Ameri:35 in geographic and environmental dimensions. Cell resolution is 0.2SO, or ~)ughly 27 km at the Equator. The climatic data are drawn from the WorldClim .ataset (Hijmans et al. 2005). An example of environments showing moderate .emperature and high precipitation is shown in black in both spaces.

22

CHAPTER 2

at coarse spatial resolutions. These dimensions are the scenopoetic variables of G. Evelyn Hutchinson and are very important in determining the broad aspects of species' distributions. Here, niches and distributions are estimated and visualized in associated geographic (G) and environmental (E) spaces. These niche dimensions (the Grinnellian niche, in effect) can be measured using data available in large quantities and estimated operationally. Another important perspective on the niche concept emphasizes impacts of a species on its environment, via linked variables and phenomena that are generally measured at finer spatial resolutions, which can be termed the "Eltonian niche." In this book, we focus on Grinnellian niches, as our interests lie primarily at geographic extents and resolutions.