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12
C H A P T E R Community Properties

A
s we hike up a mountain in the central Rockies, we might start in a for-
est dominated by ponderosa pine (Pinus ponderosa), then soon find our-
selves among lodgepole pine (Pinus contorta), then move on to spruce-
fir forest (Picea engelmannii and Abies lasiocarpa) and end up in alpine tundra.
Each area contains a very different collection of species, yet some species are
found in many different areas, and boundaries can be hard to discern. In other
environments, such as grasslands with patches of serpentine soils (soils derived
from serpentine rock that are nutrient-poor and sometimes high in toxic ele-
ments; see Box 15A), differences in species composition between communities
can be abrupt and dramatic. Humans also create boundaries that define com-
munities. A vacant lot is a community, as is a park in the middle of an urban
landscape.
In previous chapters, we looked at how species interact with one another
and their environments; here, we explore the community-scale patterns cre-
ated by those interactions. What determines the boundaries of a community?
Are communities real entities with their own properties, or are they just hap-
penstance collections of individuals and populations? This chapter investigates
what a community is, and discusses methods for describing communities. The
following chapters will cover some additional processes that create commu-
nity patterns: disturbance, succession, dominance, and species invasions.

What Is a Community?
A community is a group of populations that coexist in space and time and inter-
act with one another directly or indirectly. By “interact” we mean that they
affect one another’s population dynamics. This definition of “community”
includes all plants, animals, fungi, bacteria, and other organisms living in an
area. It seems simple enough, but ecologists often use terminology inconsis-
tently, and unfortunately, traditional usage in plant ecology sometimes differs
from that in animal ecology. For example, we speak of “plant communities”
even though plants are only a subset of the community—we are ignoring the
decomposers, herbivores, diseases, pollinators, and many other organisms.
Fauth et al. (1996) have discussed some ways through this terminological
tangle (Box 12A).
Two closely related older terms that were formerly in wider use in plant
ecology have been incorporated recently into some modern vegetation classi-
234 Chapter 12
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BOX 12A
Communities, Taxa, Guilds, and Functional Groups
A very confusing array of terms has Geography
Resources
developed to describe communities.
Sometimes the same term is used in dif-
ferent ways by different ecologists,
while in other cases different terms are Local
given to the same concept. Here we Communities guilds
Guilds
describe a recently proposed scheme to
define these terms. Ensembles
This scheme defines groups of
species based on three criteria: geogra-
phy, phylogeny, and resource use, as Assemblages
shown in the accompanying figure.
Geography in this scheme defines com-
munities, sets of organisms living in
the same place at the same time. Phy- Taxa
logeny is the pattern of relationships A scheme for defining sets of species into
among species (or higher taxa) based communities, guilds and ensembles based
on evolutionary ancestry. Phylogeny on the combination of geographical loca-
tion, common ancestry, and shared
defines taxa, sets of organisms that Phylogeny resource use. (After Fauth et al. 1996.)
share a common ancestor. Resource use
defines guilds, sets of organisms that
use biotic or abiotic resources in a sim- ing in the same place. The intersection ed in the plant community. The plant
ilar way. The term “guild” is taken from of geography and resource use defines community could therefore be consid-
animal ecology, but has been used by local guilds. The trees in a forest in ered an assemblage, as it includes
plant ecologists as well. Ontario are an example of a local guild: species that use different resources. The
Plant ecologists often use the term they include distantly related species true community, of course, includes all
functional group to describe a concept such as sugar maple (Acer saccharum), species (e.g., animals, fungi, bacteria),
related to the guild. Functional groups an angiosperm, and eastern hemlock not just plants. Thus, the traditionally
can be defined in a variety of ways, (Tsuga canadensis), a conifer. The inter- defined plant community is actually a
depending on the application, but these section of all three sets defines ensem- subset of the full community and has
definitions are all based on a set of traits bles. The grass species living together properties of ensembles, local guilds,
that identify functionally similar in a prairie are an ensemble. The annu- and assemblages.
species. The traits used to identify func- al Asteraceae in Australia’s Great Sandy
tional groups can be chosen informal- Desert are another ensemble.
ly or based on formal mathematical Plant communities as traditionally
algorithms. Ecologists have used the defined are made up of a combination
concept of functional groups in a vari- of ensembles, local guilds, and assem-
ety of contexts, including studies of the blages. Typically, terrestrial plant com-
relationship between productivity and munities are defined as all the vascu-
diversity (see Chapter 14) and attempts lar plants living in a given space. Most
to reduce the number of types of plants species in this group are primary pro-
that must be taken into account in glob- ducers with similar resource require-
al climate modeling. Extensive recent ments. So, for example, all of the grass
reviews of the concept of plant func- species in a forest understory are one
tional groups and ecological applica- ensemble of that community. The com-
tions of this concept are provided by bination of all understory forbs and
Lavorel and Cramer (1999) and Wood- graminoids would constitute a local
ward and Cramer (1996). guild, as these species would include
Intersections of the sets described by more distantly related species of mono- Monotropa uniflora (Indian pipes, Ericaceae) is
an example of a flowering plant that is a sapro-
the terms above define more narrow cots, dicots, and possibly ferns. Some phyte rather than a primary producer, obtain-
distinctions. The intersection of geog- flowering plants are not primary pro- ing its nourishment not by photosynthesis but
raphy and phylogeny defines assem- ducers, but parasites or saprophytes; from dead or decaying organic matter. (Photo-
blages, groups of related organisms liv- however, these species are also includ- graph by S. Scheiner.)
 Community Properties 235
PAGE PROOF: 2ND PASS
fication schemes (see Table 16.3). An association was a middle ground between these viewpoints, and to a
defined as a particular community type, found in many large extent have moved beyond both of them.
places and with a certain physiognomy and species com- The Clementsian view was the majority view among
position; modern usage is not much different (e.g., Table English-speaking plant ecologists during the first half of
16.3 refers to the juniper-sage woodland association). the twentieth century. Clements saw plant communities
The term formation was originally used to denote a as highly organized entities made up of mutually inter-
regional climax community; modern usage is generally dependent species. In his view (Clements 1916), com-
more specific, referring to a physiognomic subtype. munities are superorganisms—the analogue of indi-
In practice, the boundaries of plant communities are vidual organisms—that are born, develop, grow, and
usually defined operationally, based on changes in the eventually die. Succession, in this
abundance of the dominant, or most common, species. view, is analogous to the process of de-
Sampling is then confined within those boundaries. A velopment and growth; its trajectory
stand is a local area, treated as a unit for the purpose of and end point are highly predictable
describing vegetation. Typically, a number of stands are (Clements 1937). Two of the hallmarks
used to sample the presence and abundances of species of the superorganism concept were the
as well as associated environmental variables. Based on presence of very tight linkages among
data from a number of stands, the community can be species within communities and coop-
characterized. eration among the species in a com-
Only in special cases (e.g., islands, ponds, forest pre- munity for the benefit of the function
serves surrounded by suburban development, vacant of the entire community.
lots) are community boundaries defined easily. Even Even at the height of Clements’s Frederic and
Edith Clements
then, the movement of organisms and the transport of influence, many ecologists held more
matter by wind and water make their boundaries fuzzy. moderate views. This version of Clementsian ecology
Ecologists, therefore, are often of two minds when deal- asserted only that species interactions such as competi-
ing with communities. On the one hand, we recognize tion, mutualism, and predation are important in deter-
that their boundaries are fuzzy; on the other hand, we mining community structure. Clements himself ac-
often need to define discrete entities for convenience of knowledged the effects of abiotic factors such as site
analysis. Typically, plant ecologists define a community history and soils in determining community composi-
based on the relative uniformity of the vegetation and tion. He focused on the idealized nature of communities,
use their knowledge of species biology to decide when and saw them as spatially distinct, with one superor-
they are moving from one community to another. ganism complex giving way to another with a very dif-
Ecologists based in different countries, educated in ferent collection of species. His major focus was on the
different historical traditions, tend to view communities nature and development of the community as a super-
in somewhat different ways. In particular, ecologists in organism, however, rather than on the boundaries
continental Europe were historically influenced by the between communities. Some ecologists of the day accept-
floristic-sociological approach, most extensively devel- ed a more moderate version of this view, which admits
oped by Josias Braun-Blanquet (see Chapter 16). This that communities are not entirely discrete, but still
approach emphasizes the discreteness of communities. divides them into nonarbitrary groups with recogniza-
In contrast, most ecologists in English-speaking coun- ble boundaries.
tries have been more strongly influenced by the history In striking contrast to Clements,
described in the following paragraphs; as a result, they Gleason believed that communities
tend to think of communities as blending continuously are the result of interactions between
into one another. These distinct ways of thinking are individual species and the environ-
becoming less prominent as a result of increased travel ment (biotic and abiotic factors) in
and communication among ecologists worldwide. combination with chance historical
events. Each species has its own envi-
The History of a Controversy ronmental tolerances and thus re-
Within plant ecology, there has been, and still is, a range sponds in its own way to environ-
of views on the nature of communities. The extremes are mental conditions (Gleason 1917,
sometimes labeled the Gleasonian and Clementsian 1926). The implication of this belief Henry A. Gleason
views, named after Henry A. Gleason and Frederic was that along an environmental gra-
Clements, their first major proponents in the English- dient, different species would have their boundaries at dif-
speaking world. These two extreme viewpoints differ in ferent places. Not only were communities not tightly
the importance they ascribe to biotic versus abiotic fac- linked superorganisms, but defining the collection of
tors and predictable versus random processes in shap- species living together in a particular place as a com-
ing community structure. Today, most ecologists hold munity was an arbitrary human construct.
 236 Chapter 12
PAGE PROOF: 2ND PASS
According to the Gleasonian view, within the range Salix nigra
(Black willow) Acer saccharum
of environmental conditions a species can tolerate, 150 (Sugar maple) Quercus rubra
chance events determine whether a species is actually (Red oak)
found in a given location. At the local scale, chance dic- Quercus alba

Importance value
tates whether a seed happens to get to a particular spot. Ulmus americana (White oak)
On larger scales, the chance events of history play a 100 (American elm)
strong role. For example, species in the family Cactaceae
(the cacti) are found in the desert communities of the Ulmus rubra
Americas because the family originated in this region, (Slippery elm)
50
while deserts elsewhere have no cacti (except where they
have been recently introduced by humans). Further-
more, the mix of species changes from place to place as
one moves across the landscape. The Gleasonian view-
0 Wet Wet-mesic Mesic Dry-mesic Dry
point posits gradual changes in community composition
Environmental gradient
as opposed to abrupt boundaries between communities
unless there are abrupt and large environmental bound- Figure 12.1
aries. A more moderate viewpoint was that some iden- Change in the importance of various tree species along a
tifiable community types exist, but that these tend to moisture gradient in Wisconsin. Importance was measured
as the sum of the relative cover, relative density, and relative
blend into other community types. frequency of a species in a community. (After Curtis 1959.)
Clementsian ecologists were not receptive to Glea-
son’s ideas. While Gleason’s work was well known, it
failed to influence many plant ecologists until after
Clements’s death in 1945. Why Clements’s views held
sway for so long is not fully understood. Clements was matic turnover from one community type to another as
said to have an extremely strong personality that could altitude increases. Whittaker realized that if he could
dominate scientific meetings. Given the very small num- demonstrate that even along such a gradient, species
ber of plant ecologists active during the first half of the turnover was gradual, this would provide very power-
twentieth century, perhaps this alone was sufficient. ful evidence in support of Gleason’s ideas and in con-
Gleason, finding little interest in his ideas, abandoned tradiction to Clements’s superorganism model. Whit-
his work in plant ecology by 1930 and spent the rest of taker did just that. He showed that forest communities
his career as a taxonomist. along an elevational gradient in the Great Smoky Moun-
Not until the 1950s did the separate but almost tains of Tennessee changed gradually in species com-
simultaneous work of John T. Curtis and Robert H. position without abrupt boundaries (Figure 12.2). He
Whittaker convince many ecologists that Gleason’s then repeated this work in other areas, including the
views were largely correct. Curtis and his students Siskiyou Mountains of Oregon and the Santa Catalina
mapped the vegetation of Wisconsin and looked at how Mountains in southern Arizona.
species’ optima and ranges were distributed along envi- Another important line of evidence that strongly
ronmental gradients (Curtis 1959). The Clementsian the- affected many ecologists’ views of communities was a
ory predicts that species optima and ranges should show series of studies conducted since the 1970s on the distri-
distinct groupings, whereas the Gleasonian theory pre- butions of plant species during and after the most recent
dicts independence of optima and ranges. The re- glaciation (see Chapter 21). Many of these studies looked
searchers found just such independence; each species at fossil pollen from lake bottoms. Some of the earliest
had a different set of environmental tolerances (Figure and most influential of this research was carried out by
12.1). A key innovation that con- Margaret Davis. She showed that many species that co-
tributed to this study was the devel- occur today did not always do so during glacial periods;
opment of ordination, a set of tech- rather, species were distributed among communities in
niques for describing patterns in the past in very different combinations from those that
complex vegetation; we discuss ordi- are found today (Davis 1981). For example, Pinus strobus
nation in detail in Chapter 16. (eastern white pine, Pinaceae), Tsuga canadensis (eastern
Whittaker (1956) also demon- hemlock, Pinaceae), Castanea dentata (American chestnut,
strated that Gleason was right about Fagaceae), and Acer spp. (maples, Sapindaceae; maple
the nature of boundaries between pollen cannot be identified to species) are often found
communities. One of the most strik- together in the eastern deciduous forest of North Amer-
ing patterns one encounters going up ica today. However, these forest tree species were not
Robert H. Whittaker an elevational gradient is the dra- always associated in the past. White pine and eastern
 Community Properties 237
PAGE PROOF: 2ND PASS
40 Tsuga canadensis
Liriodendron tulipifera (Canadian hemlock) Halesia monticola
30 (Mountain silverbell)
(Tulip poplar)
20

10
0

40 Acer spicatum
Acer saccharum (Mountain maple,
30 Tilia heterophylla (Sugar maple) moosewood)
(White basswood)
20
Percent of stand

10
0

40 Fraxinus americana
Betula allegheniensis
30 Carpinus caroliniana (Yellow birch) (White ash)
(American hornbeam) Aesculus octandra
20 (Yellow buckeye)
10
0

80
Fagus grandifolia
60 (American beech)

40

20
0
500 700 900 1100 1300 1500 1700
Elevation (m)
Figure 12.2
Changes in plant species frequencies along an altitudinal gradient in the Great
Smoky Mountains of Tennessee. (After Whittaker 1956.)

hemlock survived the Wisconsin glaciation (about 75,000 plowed at some time in the past. Thus, current envi-
to 12,000 years before present) in a region east of the ronmental boundaries do not always match past bound-
Appalachian Mountains, while during the same period, aries. Ecologists still disagree as to the relative impor-
chestnut and maples were found near the mouth of the tance of biotic and abiotic processes and chance events
Mississippi River (Davis 1981), more than a thousand in determining community structure.
kilometers to the southwest. Echoes of the controversy between the Clementsian
Today most plant ecologists take a middle position and Gleasonian views of community among English-
between Clements’s and Gleason’s views, and in many speaking ecologists of North America and the United
ways have diverged from both of their views. There is Kingdom continue to influence ecologists schooled in
wide agreement that species are distributed individual- those traditions. Related issues were debated among
istically, and that community composition typically European and Russian ecologists, but not to the extent
changes gradually along environmental gradients. that they were argued in American and British journals;
Abrupt changes are most likely to be found where there the ecologists of continental Europe and Russia had dif-
are abrupt changes in the environment. However, abi- ferent concerns. Heavily influenced by Braun-Blanquet,
otic boundaries and community boundaries do not they focused primarily on systems of community clas-
always match. Because of processes such as dispersal sification. We will return to their traditions in Chapter
from one habitat into another (see Chapter 17), a popu- 16. The key difference is that the European tradition was
lation may extend partway into an unfavorable envi- more concerned with describing patterns than with ana-
ronment. Abrupt changes may also reflect past events, lyzing processes, mainly sidestepping the arguments
such as the edge of a fire or a part of a forest that was described here and in the next chapter.
238 Chapter 12
PAGE PROOF: 2ND PASS
A Modern Perspective on the Issues in Contention documenting patterns (as Curtis, Whittaker, and others
The primary issues surrounding the nature of plant com- did), understanding the processes responsible for creat-
munities divide roughly into those of pattern and those ing those patterns, and posing plausible explanations—
of process. Underlying these issues is theory: the effort elucidating the mechanisms causing those patterns and
to explain the patterns and processes, which includes a processes.
search for the mechanisms responsible (see Chapter 1). Whether communities can be considered to have
The issues of pattern focus on how species and com- emergent properties depends in part on the relative
munities are distributed over the landscape. Are bound- importance of biotic and abiotic processes in shaping
aries among communities abrupt or gradual? How pre- community structure, including how strongly species
dictable are the patterns? Questions of pattern are critical interact with one another within communities. (We view
because they set the stage for the rest of the debate. Once the issues in somewhat different terms today than either
patterns are identified, theories can be constructed to Clements or Gleason viewed them, as we will see in
explain them. In this chapter we examine ways of meas- Chapter 13.) Emergent properties are those that come
uring patterns within communities; among-community about through interactions, such as competition, preda-
patterns are dealt with in Chapters 16 and 17. tion, and mutualism, that occur among the populations
The issues of process focus on what processes actu- in a community (Box 12B). If these processes play a
ally function in natural communities and which of these major role in shaping communities, then communities
are most important in determining the observed pat- have at least some emergent properties. But if commu-
terns. We know of many processes that influence com- nities are mainly structured by the tolerances of indi-
munity composition, including the physiological toler- vidual species for abiotic factors (such as minimum tem-
ances of species, competition, herbivory, biogeography, peratures) in the environment, then community
historical contingency, and random factors influencing properties are largely aggregates of the individual
colonization. All can be shown to operate at some times species’ properties. It is important to note that one can
in some places; the question is their relative importance. recognize that emergent properties are important with-
Do some processes predominate in determining pat- out accepting Clements’s superorganism view, by dis-
terns? Is the relative importance of these processes carding the ideas that communities consist of species
always the same, or are some more important some of adapted for one another’s benefit and that all or most
the time, or in certain situations, or in certain types of species in a community are tightly interlinked. Among
communities? One fundamental issue is whether com- modern ecologists, for example, E. P. Odum, H. T.
munities are primarily static, exist at some sort of Odum, R. V. O’Neill, and their co-workers have strong-
dynamic equilibrium, or are always changing. We will ly emphasized the idea that communities and ecosys-
explore this issue in detail in Chapters 13 and 14. Final- tems have emergent properties, but none of them
ly, pattern and process are tied together by theories embrace the superorganism view (E. P. Odum 1971; H.
regarding the nature of communities—such as T. Odum 1983, 1988; O’Neill et al. 1986).
Clements’s superorganism theory—which seek to On the whole, ecologists have shifted their attention
explain the mechanisms responsible for the patterns and to working out the mechanisms behind the patterns
processes that are documented. found within particular communities. This shift from the
An overarching issue is the problem of scale. Recent- study of emergent properties to the study of mechanisms
ly, ecologists have paid increasing attention to the real- is emblematic of the seesawing between reductionist
ity that different patterns and processes may exist and (mechanistic) and holistic (emergent) approaches that
function at different scales. The processes that are impor- has characterized community ecology for the past cen-
tant within communities at the scale of meters may dif- tury. Ecologists in the 1980s and 1990s tended toward
fer from the processes that are important across biomes reductionist approaches. Now there is some movement
at the scale of thousands of kilometers. Throughout this back in the direction of holistic studies under the man-
book, we examine how scale affects both the patterns tle of macroecology (Brown 1995, 1999; Lawton 1999).
and the processes one finds, as well as the answers one
reaches to questions of the relative importance of dif- Are Communities Real?
ferent causes and mechanisms. As we have seen, the extent to which communities are
In Clements’s original superorganism theory, the “real” has been a contentious issue among plant ecolo-
community was an organic entity: a distinct unit with gists for much of the twentieth century. The heart of
strong emergent properties. Species within the com- the debate has been philosophical: What types of enti-
munity were tightly linked and interdependent. In con- ties are real, and what types are just mental constructs?
trast, in Gleason’s view, any community-level proper- It is clear that populations and species are real entities,
ties were simply the sum of the properties of individual but are communities real entities as well, or are they
species. Determining the truth of the matter requires merely convenient but arbitrary human inventions? In
 Community Properties 239
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BOX 12B
A Deeper Look at Some Definitions:
Abiotic Factors and Emergent Properties
Ecologists have distinguished between Chapter 22) is affected by living things. property of the biomass of each indi-
the effects of abiotic (nonliving) and We do not have a good substitute to vidual: We can determine community
biotic (living) factors in the environ- suggest for the term “abiotic,” so while biomass by just adding up the biomass
ment for a very long time. Typical biot- we use it here to mean “things like cli- of all of the species in the community.
ic factors include competition and pre- mate and soils,” we recognize that these But we can always measure communi-
dation; typical abiotic factors include may have major biotic components. ty biomass simply by weighing all of
soil nutrients, microclimate, weather, Another term that bears closer ex- the individuals; community biomass is
and general climatic influences. The amination is “emergent properties.” A not an emergent property.
problem with this terminology is that central issue in the argument over the In contrast, consider canopy photo-
the distinction between abiotic and nature of communities is the question synthesis: we cannot measure the pho-
biotic factors may be far less clear than of whether emergent properties exist. tosynthetic rate of an entire forest
we might think. As we emphasize in An emergent property is one that is canopy just by measuring the photo-
Chapters 4 and 15, soils are a product found at a certain level of organization synthetic capabilities of the individual
of organisms and their interactions due to properties, structures, and pro- plants. Canopy photosynthetic rates
with the environment. The nitrogen cesses that are unique to that level of depend on how individuals interact in
available to the roots of a plant, for organization. Emergent properties can several ways, including shading one
instance, depends on the actions of be contrasted with properties that are another, interfering with wind, and tak-
many different kinds of soil organisms merely aggregates of properties at a ing up CO2. Canopy photosynthetic
and the interactions of the plant with lower hierarchical level. For example, rate is an emergent property of the
those organisms. Likewise, the micro- the total biomass of all of the plants in community.
climate—and even global climate (see a community is merely an aggregate

the past, these questions often focused on the problem them if it is not. The debate as to whether communities
of boundaries—of identifying where a community are real entities or imaginary constructs is more than just
begins and ends. Many studies have shown that, except an academic exercise. For example, The Nature Con-
where there are abrupt physical discontinuities, plant servancy (TNC), based in the United States, makes deci-
communities tend not to have discrete boundaries. At sions about land acquisition and restrictions on land use
one extreme, this observation might be interpreted as based on the classification of the communities present.
implying that there are no community-level processes In collaboration with the Natural Heritage network,
worth studying. TNC has devoted substantial effort to describing and
This conclusion would be wrong; in fact, the debate classifying communities under the U.S. Natural Vegeta-
is miscast. Instead of focusing on patterns, let’s shift tion Classification (USNVC) system (see Table 16.3; May-
the focus to processes and rephrase the question by ask- bury 1999). Classifications of communities are even
ing whether community-level processes are important incorporated into law in a number of places. In south-
in structuring the living world. We have already dis- ern California, for example, land development is regu-
cussed several processes responsible for interactions lated quite differently for “coastal sage scrub” than for
among species: competition (see Chapter 10), herbivory “chaparral” communities, although by any definition
(see Chapter 11), and mutualisms (such as those dis- the two include many of the same species, and different
cussed in Chapter 4). These are all community-level scientists may have used one name or the other to cate-
processes that occur among the component parts of com- gorize the same place.
munities (e.g., populations). If such processes are sig-
nificant factors in the structuring of a particular system, Describing Communities
then we can regard that system as a community with
unique properties. Such an outlook eliminates the need Determining which processes are most important in
to worry as much about the existence of clear bound- shaping communities requires that we be able to
aries between communities. We can recognize the exis- describe and compare communities. What kinds of prop-
tence of communities if it is useful to do so, and ignore erties do plant communities have, and how can they be
240 Chapter 12
PAGE PROOF: 2ND PASS
characterized? One set of community properties encom- (A)
passes the numbers of species present and their rela-
14
tive abundances: species richness and diversity. A sec-

Number of species
ond set encompasses the physical structure of the plant 12
community: its physiognomy. 10
8
Species Richness 6
One way to describe a community is by a list of the 4
species in it. Species richness is the number of species 2
on such a list. Because we often have information about
the biology and ecology of those species, the list can 0 1 1 1 1 1 2
16 8 4 2
point to many other types of information, such as the Quadrat area (m2)
numbers of trees or herbs present, or the numbers of
species in different taxonomic groups. (B)
How would one gather such a list? A simple and 1m
widely used method is to establish the boundaries of the
community and then walk through it, identifying all of
the plants and listing them. Such a survey should be
done several times during the year because some species
may be visible only during a single season. Spring
ephemerals, for example, are perennial plants common
in temperate deciduous forests whose aboveground
parts are present for only one to two months in the
spring; during the rest of the year they exist only under-
ground as dormant corms or rhizomes.
While simply finding and listing all species is useful,
this method has limitations. Most importantly, if we wish
to compare communities, we need comparable sam-
ples—otherwise we might find that two communities are
“different” simply because we sampled one more inten-
sively than the other. The area sampled can have strong Figure 12.3
effects on the number of species found. This sort of sam- (A) A hypothetical species-area curve. (B) A system of nested
pling effect is best dealt with by using plot-based meth- quadrats used to determine a species-area curve.
ods, in which a series of sample plots, or quadrats, are
marked out in a community and a list of species is col-
lected for each. The quadrats may be any shape, such as
square, rectangular, or round. They may be nested, con- curve reaches a plateau. Of course, if the area becomes
tiguous, spaced along a line, placed in a grid, or placed large enough to encompass new environments, the
randomly. These different arrangements can be used to curve begins to rise again.
ask different types of questions or to control for variation The best mathematical model for the shape of the
that occurs on different spatial scales (Krebs 1989). species-area curve has been debated over the years.
By looking at how the total number of species found Three candidates have been proposed: the exponential
increases as quadrats are combined, one can examine the curve,
effects of area on species richness. The species-area
S = Z ln(A) + C
curve (Arrhenius 1921; Gleason 1922; Archibald 1949)
describes the increase in the number of species found as the power curve,
the area sampled increases (Figure 12.3A; see also Fig-
S = CAZ
ure 17.2). This increase comes about for two reasons:
first, as more individuals are sampled, the chance of and the logistic curve,
encountering a new species increases; and second, a larg- B
S=
er area is more environmentally heterogeneous. For a C + A−Z
given community, if it has a relatively uniform envi-
ronment, the number of new species found for each where S is the number of species, A is the area, and B, C,
increment in sampling area decreases. Eventually, very and Z are constants (Figure 12.4). The three equations
few or no new species are found, and the species-area are actually members of the same mathematical family,
 Community Properties 241
PAGE PROOF: 2ND PASS
such that the exponential function is a special case of the An alternative method of estimating species rich-
power function—that is, they are equivalent when ness is to ask how many species are found in a sample
Z ln( A) of some number of individuals. For some functional
C= groups, such as trees, in which individuals are easy to
AZ − 1
identify, this approach can be useful. Mathematical tech-
and the power function, in turn, is a special case of the niques, called rarefaction, exist for standardizing esti-
logistic function—that is, the power and logistic func- mates among samples that include different numbers of
tions are equivalent when individuals and for estimating the number of very rare
species that were missed in the sample (Krebs 1989).
ln (B − C )/ C 2 

Z=   These techniques were developed first and are used
ln( A) most often in animal ecology, although they are being
used increasingly by plant ecologists. Because animals
Which of the three curves best fits a given data set? This often move, area-based measures are less useful to ani-
depends on the area being sampled and the scale of envi- mal ecologists. Plant ecologists, on the other hand, tend
ronmental variation. For small or intermediate-sized to emphasize area-based techniques because the clonal
areas, a logistic curve often fits bests, while for large nature of many plants can make it difficult to distinguish
areas a power curve is often the best fit. individuals (see Chapter 8). In plant ecology, dry weight
These three models make different assumptions (the weight of a sample after it has lost all its moisture
about the relationship of species number to area. Both and is at a constant weight), or biomass, is often used
the exponential and power functions result in curves instead of number of individuals.
that continue to rise indefinitely. The logistic function, How is the number of species in a community actu-
on the other hand, eventually reaches a plateau. This dis- ally determined? One can use species-area curves to
tinction is important if one wishes to answer the ques- determine the total area required for standardized sam-
tion, “How many species are in the community?” If the pling. By using equal areas in different stands within a
species-area curve rises indefinitely with increasing area, particular community, we avoid differences due to sam-
then the answer depends entirely on the area of the com- pling intensity. We may also wish to compare different
munity. However, the “area” of a community is often community types in a study using the same methods. For
arbitrary, making the question meaningless. Instead, we example, we might sample a beech-maple community
can rephrase the question as, “How many species are and an oak-hickory community using the same
contained in an area of size X?” This value is known as approach. For very different kinds of plant communities,
species density. If the species-area relationship is best however, such as grasslands and forests, we might need
described with a logistic curve, then this question has to use different sampling areas and somewhat different
meaning, because the curve has a definite plateau or techniques to best determine the characteristics of each.
asymptote, calculated as B/C. The issue of the shape of How large an area should be sampled? We want a
the species-area curve will become important when we sample that is large enough to contain most of the species
explore comparisons of diversity among communities and which will minimize differences due to random sam-
in Chapter 16. pling effects, such as where plots happen to be placed.
On the other hand, for practical purposes, we do not
want the area to be bigger than necessary. With respect
to a species-area curve, we want to sample a total area
for a given community that is at least as large as the point
Power
at which the curve begins to level off at a plateau, or, if
there is no plateau, at which the rate of increase with
Number of species

Logistic
increasing area is very small. We will return to the par-
ticular methods used for sampling vegetation shortly.

Diversity, Evenness, and Dominance

Species richness is only one aspect of diversity. Not all
Exponential species exist in equal numbers: some are rare, some are
common but not numerous, and others are very abun-
dant. Imagine two forest stands, both of which contain
ln (Area) a total of 100 individuals (or 100 kg of biomass) belong-
Figure 12.4 ing to 5 different species. In one forest stand, there are
Examples of three differently shaped species-area curves: 20 individuals of each species. In the other, one species
exponential, power, and logistic. has 60 individuals, while each of the other 4 species has
242 Chapter 12
PAGE PROOF: 2ND PASS
Community A Community B There are various measures and meth-
ods for expressing species diversity (Table
12.1). Which one is most appropriate in a
given situation depends on what aspects of
a community one wishes to highlight; each
has different assumptions, advantages, and
limitations. Two commonly used measures
of species diversity (which combine the
effects of species richness and evenness)
are the Shannon-Weiner index and the
inverse Simpson’s index.
The Shannon-Weiner index is comput-
ed as
s
H′ = ∑ (pi ln pi )
i =l

where s is the number of species, pi is the

proportion of individuals found in the ith
species (pi = ni/N), ni is the number of indi-
viduals of species i in the sample, and N
Figure 12.5 is the total number of individuals sampled.
Samples from two different communities. In the sample on This index assumes that individuals were sampled from
the left all the species have equal numbers of individuals; this
a very large population and that all species are repre-
sample has greater evenness than the sample on the right.
sented in the sample.
Simpson’s index measures the chance that two indi-
viduals chosen at random from the same community
10 individuals. These two samples differ in a property belong to the same species:
called evenness (Figure 12.5). The first, in which the s
∑ pi
species are represented by the same number of indi- 2
L=
viduals (or the same amount of biomass), is more even, i =l
and thus has one of the essential elements of being more
diverse than the second. The species diversity of a com- where the summation is over all the species. Both of these
munity depends on both its richness and its evenness: indices are commonly transformed so that they range
higher species numbers, with the individuals (or bio- from a minimum of 1 to a maximum of S, the total num-
mass) more evenly distributed among them, contribute ber of species in the sample, when all species are equal-
to higher community diversity. A way to think about ly common. For the Shannon-Weiner index, this trans-
evenness as a contributor to diversity is to consider the formation is the exponential (eH′), and for Simpson’s
following thought experiment: Pick two plants at ran- index it is the inverse (1/L = D). The ratio J′ = H′/ln S,
dom from a community. Are they members of the same called the Shannon evenness index, provides a measure
species or different species? In a highly diverse com- of evenness.
munity, it is more likely that they will belong to differ- If L is the probability that two randomly chosen indi-
ent species. viduals are from the same species, then (1 – L) is the prob-
In this example, the species that has by far the great- ability that they are from different species. This number
est number of individuals, or biomass, in the second is also known as the Gini coefficient. While the Gini coef-
community is the dominant species in that communi- ficient has been used as a measure of evenness among
ty. The first community, with greater evenness, does not individuals in a population, it could also be used as a
have any dominant species. The greater the numerical measure of species evenness in a community (Lande
preponderance of one or a few species, the lower the 1996).
diversity of the community tends to be. Of course, a Which of these indices is more useful? This question
community may have one strongly dominant species has been the topic of much discussion, which has led to
and a large total number of species, giving it a high the development of numerous ways of correcting for
diversity value, but this is actually fairly unusual. Other sampling bias (departure of the estimated value for the
ways of characterizing variation among species in abun- sample from the true value due to systematic, non-
dance are considered in Chapter 14. chance causes). Lande (1996) showed that Simpson’s
 Community Properties 243
Table 12.1 Some species diversity and evenness indices

Usual
Index symbol Formula Emphasizes

Species number indices

S
Species density None area sampled

Margalef ’s index DMg S −1

ln N

Menhinick’s index DMn S

N
Proportional abundance indices
Shannon-Weiner index H′ Σ
– [p ln (p )]
i i Rare species
Inverse Simpson’s index D 1/Σpi2 Common species
 
log 2  N ! 
Pielou’s index HP  ∑ n i ! Rare species
N

Brillouin index HB ln(N !) − ∑ ln(n i !) Rare species

N
McIntosh’s U index U ∑ n i2 Rare species

∑ni
2
N−
McIntosh’s D index D Common species
N− N
N max
Berger-Parker index d Common species
N

∑ n i − NS
Cuba’s index DC S +1− Common species
2N
1
2
n R1 + ∑ n r + 21 n R2
Q statistic Q Rare species
R 
log  2 
 R1 
Evenness indices
H'
Shannon evenness J ln(S )
HB
 
ln  n!
Brillouin evenness EB
1

N  [ N ]!S − r {( N + 1)!}r 
 S S 

McIntosh evenness EU
N− ∑ n i2
N− N
S
Definitions of symbols
S = the number of species in the sample
pi = the proportion of individuals in the ith species (pi = ni/N)
ni = the number of individuals of species i in the sample
N = the total number of individuals sampled
Nmax = number of individuals of the most abundant species
nr = the total number of species with abundance R
R1 and R2 are the 25% and 75% quartiles
nR1 = the number of individuals in the class where R1 falls
nR2 = the number of individuals in the class where R2 falls
Note: While these indices were originally defined with regard to numbers of individuals, other measures, such as
cover or biomass, can be used. For more information on diversity indices, see Magurran (1988).
244 Chapter 12
PAGE PROOF: 2ND PASS
index can be estimated without bias, but that estimating There are other methods besides the use of indices
the Shannon-Weiner index requires actually knowing for quantifying diversity and comparing diversity
the true number of species. The unbiased estimate of D among communities. These methods include graphical
is (1 – N)/(1 – NL). This lack of bias is an advantage for approaches based on comparisons of dominance-diver-
Simpson’s index over the Shannon-Weiner index. Which sity curves (discussed more fully in Chapter 14). Other
index is preferable in a given study depends in part on approaches have been used, but they go beyond the
what you wish to emphasize: the Shannon-Weiner index scope of this book.
is sensitive to changes in the proportions of rare species,
while the inverse Simpson’s index is sensitive to changes Sampling Methods and Parameters for Describing
in the proportions of common species (Table 12.2). If one Community Composition
had hypothesized, for example, that some environmen- Several techniques can be used to sample a communi-
tal factor were having a negative effect on community ty. Which to use in a given case will depend on the type
diversity over time by disproportionately affecting rare of vegetation being sampled and the goal of the survey.
species, reducing their abundance or eliminating them, Plant ecologists influenced by the Zurich-Montepellier
the Shannon-Weiner index might pick up such effects school (the Braun-Blanquet approach; see Chapter 16)
more readily. typically sample and compare multiple communities by
Just documenting that one community has a higher placing a single large sample plot—called a relevé—in
diversity than another community is not enough to con- each stand. The relevé is located subjectively, with an
clude that the two communities are truly different. The attempt to place it within a uniform patch of vegeta-
communities may actually be identical, and the differ- tion that is representative of the community. How large
ences in diversity found might be due to chance sam- should the relevé be? Ecologists determine the appro-
pling events. One more piece of information, a meas- priate relevé size for a given type of community by using
ure of the precision of our diversity parameter, is needed a set of nested relevés in one or a few communities,
before comparisons can be made. Methods for estimat- beginning with a small one and increasing the area sam-
ing precision are described in the Appendix. A formula pled around it in a series of steps (see Figure 12.3B). As
to estimate the variance for Simpson’s index is given in the total area is increased, a species-area curve is con-
Lande (1996), and the variance for the Gini coefficient structed until the curve reaches a plateau. That size
can be estimated with methods in Lande (1996) or Dixon relevé is then used for the remainder of the communi-
(2001). More detailed information on such topics as sam- ties sampled (if they have similar enough characteristics
pling effects, effects of quadrat size and shape, and sen- and structure). Because European botanists carried out
sitivities to various patterns of abundance can be found many of the early vegetation surveys in the Tropics,
in Greig-Smith (1964), Pielou (1975), Magurran (1988), often the only available quantitative descriptions of these
and Krebs (1989). areas have been done using the relevé method.
Ecologists in English-speaking countries
have long used the approach of sampling a
given stand or community using some num-
Table 12.2 A comparison of three species diversity measures as applied to ber of smaller quadrats, or sample plots,
six communities, each containing five species making the size and number of quadrats the
Community same across stands or communities. These
quadrats can be placed either randomly with-
1 2 3 4 5 6
in the community or regularly along a grid.
Species A 20 30 40 50 60 960 The advantage of using a number of smaller
Species B 20 30 30 20 10 10 quadrats, rather than one large relevé, is that
Species C 20 20 10 10 10 10 any local patchiness will even out across the
Species D 20 10 10 10 10 10 entire sample. This method also provides a
Species E 20 10 10 10 10 10 measure of local heterogeneity. The issue of
Sample size 100 100 100 100 100 1000 regular (systematic) versus random place-
Species richness 5 5 5 5 5 5 ment of the quadrats has been heatedly
eH′ 5 2.84 2.07 1.95 1.36 1.04 debated, and involves considerations beyond
D 5 4.17 3.57 3.13 2.50 1.08 the scope of this book (interested readers
J 1 0.65 0.45 0.42 0.19 0.02 should consult the methodological references
at the end of this chapter).
Note: The exponential Shannon-Weiner index and the inverse Simpson’s index differ in how
they change as evenness changes from sample 1 to sample 6. The numbers refer to numbers The size and shape of the quadrats is
of individuals, but could also represent biomass. also an issue. Forests in North America are
 Community Properties 245
PAGE PROOF: 2ND PASS
often sampled using 0.1-hectare quadrats. The total area Quadrats and transects can be used to estimate sev-
sampled should generally be determined by the species- eral parameters, based on species’ presence or abun-
area curve, as we saw above. As for quadrat shape, dance in the samples. Frequency is the percentage of
square quadrats are typically used, especially for small quadrats in which a species appears. Such a measure has
areas (< 4 m2). Usually the quadrat is marked with a the advantage that it can be done quickly, even by a sin-
rigid sampling frame, formerly made of wood or alu- gle person. In particular, it is not necessary to be able to
minum, now typically made of PVC pipe. Larger determine how many individuals of a given species are
quadrats can also be set up using posts and measuring in a quadrat. The limitation of this method is that it is
tapes in the field, particularly in forests. For the largest inaccurate for very rare species or species with clumped
areas, circular quadrats may be more efficient. In this distributions and uninformative for very common
case, a post is placed at the center of the sampling area species (those that appear in every quadrat).
with a rope or tape measure attached to a swivel point Another quantitative measure is cover: the percent-
at the top. The desired radius is marked on the rope, and age of the ground covered by a given species. Cover may
a circular area is swept out. This method has the advan- be measured as basal cover or basal area—the area
tages of avoiding the problem of precisely determining occupied by the base of a plant, such as a tussock grass
90° corners for a square or rectangular plot and of hav- or a tree with a definable base—or as canopy cover.
ing a lower area-to-circumference ratio. Cover values can tally to more than 100% if there are sev-
Ecological conditions may also dictate the shape of eral layers of vegetation (e.g., in a forest, trees may cover
the quadrat. For example, on a steep hillside, if you 70% of the ground, herbs 50%, and shrubs 20%). Cover
wanted a given sample to be within a particular eleva- can be estimated in several ways.
tional contour, a narrow rectangular plot would be most The most precise, but most laborious, method of
appropriate. A narrow rectangle might also be prefer- estimating cover is to map the vegetation. Efficient map-
able in a patchy habitat if one wanted to include more ping techniques exist for large individuals—shrubs and
patches in the samples. With a narrow rectangle, how- trees. Alternatively, for herbaceous vegetation, if it is not
ever, there is a greater chance of miscounting because too dense and the species are distinctive, a photograph
more plants are near the edge of the quadrat. The shape can be taken from above. Areas of cover are then deter-
and placement of quadrats are known to affect the num- mined using image analysis software. The point-inter-
ber of species found. Depending on the pattern of envi- cept or point-quadrat method for quantifying cover uses
ronmental heterogeneity, rectangular quadrats may con- a sampling frame with a grid of strings or a frame that
tain more species than do square or circular quadrats is used to drop pins at precise locations. The identity of
of the same size. the plant at each point where the strings cross or the pin
A related approach to sampling vegetation is the use is dropped is then determined. The total number of
of transects: very long, narrow sampling areas along “hits” (points occupied) by a species is an estimate of its
which species abundances are determined. Often tran- cover, given that enough points are sampled. In other
sects are designed to run either across or parallel to gra- words, if a species were found at 12% of the points, its
dients of environmental change, depending on the pur- estimated cover would be 12%.
pose of the sample. Transects are particularly useful in Photographic and point-intercept techniques are
sampling very large areas. One kind of transect is just a intensive and time-consuming, and are best for small
line. In the line-intercept method, a line of a set length sampling areas. For larger areas or less intensive efforts,
(e.g., 100 m) is laid through the community, and the pro- visual estimates are typically used. Humans are able to
portion of the length along the line occupied by each discern differences of about 10% by eye, so cover can be
intercepted species is determined. In a forest, a tape estimated to approximately the nearest 10% in this way.
might be suspended at some height, and the cover of Nonequal cover classes can also be used (Table 12.3). The
overstory species above the tape and understory species use of cover classes is an established part of relevé sam-
below the tape recorded along the length of the line. pling.
Another method is the belt transect. As with the line- A further quantitative measure is density, the num-
intercept method, a line of a set length is laid through a ber of individuals of a species per unit area. Density can
community. Now, all individuals lying within a belt of be measured only for species with distinct individuals
some set width on either side of the line (e.g., 0.5 m) are (e.g., trees) or where ramets are treated as individuals.
counted (belt transects can also be used to measure For small plants, quadrats are used and all of the indi-
cover). A belt transect, in other words, is a long, thin viduals are counted. For larger individuals, especially
quadrat. A long belt transect may encompass more topo- trees, plotless methods, such as the point-centered quar-
graphic or soil variation, and thus may include more ter method, can be used. To implement this method, a
species, than a square grid for a given total area sampled. series of random points are located in the community,
246 Chapter 12
PAGE PROOF: 2ND PASS

Table 12.3 Three different systems for estimating cover by categorizing

estimates into a limited number of classes
Braun-Blanquet Domin-Krajina Daubenmire

Class Cover range Class Cover range Class Cover range

5 75–100 10 100 6 95–100
4 50–75 9 75–99 5 75–95
3 25–50 8 50–75 4 50–75
2 5–25 7 33–50 3 25–50
1 1–5 6 25–33 2 5–25
+ <1 5 10–25 1 0–5
r <<1 4 5–10
3 1–5
2 <1
1 <<1
+ <<<1
Source: Mueller-Dombois and Ellenberg 1974.

often along a transect line. The area around each of the usually determined by harvesting samples of the vege-
points is divided into 90° quarters, usually along com- tation, but can sometimes be estimated nondestructive-
pass headings. Then, in each quarter, the distance of the ly. For trees, a typical measure of biomass is diameter
nearest individual from the central point is measured. at breast height (DBH). DBH is measured at a standard
These distance measures are then converted into densi- height above the ground, often by measuring circum-
ties. Other methods, such as nearest individual, nearest ference using a special tape that converts circumference
neighbor, and random pairs, are variants of this tech- to diameter units. For some tree species, especially those
nique. Greig-Smith (1964) and Krebs (1989) discuss the that are commercially important, the relationship
details of using these methods, none of which are per- between DBH and total biomass is known. The investi-
fect. For example, clumping of individuals can result in gator can also determine this relationship by harvesting
incorrect estimates of density, depending on the details a sample of individuals, if necessary.
of how sampling is carried out.
Because different species may be represented more Physiognomy
heavily by one or another of the above measures of Physiognomy is the form, structure, or appearance of a
abundance, it is sometimes useful to combine them in a plant community. Knowledge of the physiognomy of a
weighted measure called an importance value (IV). Cur- community may tell us about the adaptations of its dom-
tis and McIntosh (1951) first defined the importance inant species to environmental conditions. The Danish
value of a species as the sum of the relative cover (i.e., ecologist Christen Raunkaier developed one widely
cover of that species divided by total cover of all species used system for describing community physiognomy.
present), relative density, and relative frequency of the He classified species based on their growth form and on
species in a community. Importance values can also be the location of their overwintering organs (those that can
defined by combining other measurements of relative survive an unfavorable season), such as meristems and
abundance, depending on what is most relevant in a underground bulbs (see Figure 8.2). According to this
given study. system, a tropical forest is made up almost exclusively
Biomass is generally correlated with cover. Cover, of phanerophytes (Table 12.4). A desert community is
however, is limited to two dimensions, whereas biomass dominated by shrubs—chamaephytes—and annual
is correlated with three-dimensional properties because herbs. A prairie is dominated by grasses and perennial
plant parts fill space. In some ways, it is a better meas- forbs—hemicryptophytes and cryptophytes. We can use
ure of abundance than cover. For many species, biomass this system both to classify general vegetation types
is a more useful measure than number of individuals (Raunkaier’s original intent) and to begin to make infer-
because individuals within a population of the same ences about the environmental factors that shape these
species can often differ in size by several orders of mag- communities. The prairie, for example, is dominated by
nitude (consider a seedling and a full-grown oak tree), species whose meristems are at or below ground level,
and the effect of an individual within its community typ- suggesting that meristems above ground level are vul-
ically depends on its size (see Chapter 10). Biomass is nerable to damage. In this case the critical environmen-
 Community Properties 247
PAGE PROOF: 2ND PASS
(A) Figure 12.6
40 (A) A bisect of a tropical rainforest in Trinidad,
British West Indies. (After Beard 1946.) (B) A
35 diagrammatic bisect of a temperate woodland.
The letters in the key can be used as “short-
30 hand” to provide information about the life
form, leaf shape, function, and leaf texture of
25 the species. For example, Tmdh(v)zi would
Height (m)

indicate a deciduous tree of medium stature

20
with large, compound membranous leaves and
15 discontinuous coverage. (After Dansereau
1951.)
10

0 10 20 30 40 50 60 70 80
Horizontal distance (m)
tal factors are fire and grazing (see Chapters
13 and 19).
(B) An important feature of plant commu-
nities is vertical structure. A forest commu-
nity, for example, may consist of canopy
trees, understory trees, shrubs, and herbs. A
10
tropical rainforest may have as many as six
different vegetation layers, or strata. Grass-
lands can consist of a mixture of ground-
Meters

hugging forbs and tall grasses. These prop-

erties can be quantified in several ways. The
5 bisect is a visual technique consisting of a
scale drawing of the vegetation along a tran-
sect (Figure 12.6A). A more diagrammatic
2 form of such a drawing is also possible (Fig-
0.5 ure 12.6B). Data from bisects or other sur-
0 veys can be used to determine the vertical
profile of the community. Along with den-
Life form Leaf shape and size sity, cover, or basal area, the height of each
individual is measured. Each species will
T Trees n Needle occupy some range of heights; often seed-
F Shrubs g Graminoid lings or saplings of canopy tree species are
treated separately from adults. By combin-
H Herbs a Small ing species that occupy the same stratum,
M Bryophytes h Large, broad one can produce a graph that summarizes
the distribution of vegetation in different
E Epiphytes v Compound vertical strata (Figure 12.7). These data can
L Lianas q Thalloid also be used to statistically compare verti-
cal structure among sites or community
Function Leaf texture types.
d Deciduous f Filmy
s Semideciduous z Membranous Long-Term Studies
e Evergreen x Sclerophyll
j Evergreen-succulent, leafless k Succulent or fungoid The measurements described above are like
snapshots of a community at a single point
Size Coverage
in time. But ecological processes, such as
t = Tall (T = to 25 m; F = 2–8 m; H = 2 m) b = Barren
m = Medium (T = 10–25 m; F, H = 0.5–2 m) i = Discontinous succession (see Chapter 13) or response to
l = Low (T = 8–10 m; F, H = to 50 cm) p = Tufts, groups global change (see Chapter 22), can take
c = Continuous many years, so these measurements may
be inadequate to describe a community.
Changes in a community may be gradual,
as the environment changes from year to
248 Chapter 12
PAGE PROOF: 2ND PASS

Table 12.4 Distribution (percentage) of growth forms for various representative communities based on
the Raunkaier systema
Phanerophytes Chamaephytes Hemicryptophytes Cryptophytes Annuals
World average 46 9 26 6 13
Tropical rainforest 96 2 0 2 0
Subtropical forest 65 17 2 5 10
Warm-temperate forest 54 9 24 9 4
Cold-temperate forest 10 17 54 12 7
Tundra 1 22 60 15 2
Mid-temperate 34 8 33 23 2
mesophytic forest
Oak woodland 30 23 36 5 6
Dry grassland 1 12 63 10 14
Semidesert 0 59 14 0 27
Desert 0 4 17 6 73
Source: Whittaker 1975, Table 3.2.
a
These terms are defined in the legend of Figure 8.2.

(A) Eastern deciduous forest

year, or dramatic, as after a hurricane or major fire.
Therefore, a sample taken in a single year may not be
30
very meaningful; in any case, its value is greatly Canopy trees
enhanced by other samples taken at other times. If we
want to study the effects of environmental variation, we
Height (m)

20
need to sample that variation adequately; even if we are
interested only in knowing what happens on average, a
single year’s data may not be “average.” 10 Understory trees
Some of our most important ecological questions
can be answered only with long-term studies (Leigh and Shrubs
Johnston 1994). A good example of such a question is the 0 Forbs and graminoids
effect on global warming. The changes predicted to 25 50 75 100
occur in temperature and precipitation will, in some Cover (%)
cases, take decades to be manifested (see Chapter 22).
From one year to the next, these changes are expected to (B) Boreal coniferous forest
be small relative to normal year-to-year variation. By
monitoring the same communities and individuals over 30
long periods of time, we may be able to most precisely
detect the effects of long-term climate change.
Height (m)

Within long-term study areas, quadrats or individ- 20

uals are permanently marked. Repeated measure- Canopy trees
ments—perhaps of cover, productivity, diversity, phe-
nology, or other population, community, or ecosystem 10
features—can then be taken according to a set schedule,
often yearly but possibly at shorter or longer intervals. Shrubs
In such situations, nondestructive sampling methods are 0 Forbs and graminoids
25 50 75 100
usually preferable to destructive ones. If destructive
Cover (%)
methods must be used, they need to be done in prede-
termined locations so that, for example, the destructive Figure 12.7
sampling does not affect other sampling areas in the Vertical profiles of two different forest types. Each bar repre-
future. sents a different stratum of vegetation. The average height
range of a stratum is given by the width of the bar. The total
Ecology is a relatively young science, so most stud- cover by that stratum is given by the length of the bar. (A)
ies that we call “long-term” are less than a few decades Typical eastern deciduous forest. (B) Typical boreal conifer-
old. However, there are several studies that have been ous forest.
 Community Properties 249
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BOX 12C
The Long-Term Ecological Research Network
The Long-Term Ecological Research deciduous forest (Coweeta, Georgia), ing on international networking in long-
(LTER) network is a collaborative effort old fields (Kellogg Biological Station, term ecological research. Representa-
involving more than 1100 scientists and Michigan), and the desert-urban inter- tives of scientific programs and net-
students, with the goal of investigating face (Phoenix, Arizona). Besides pro- works that focus on ecological research
ecological processes that operate at long moting long-term studies, these sites over long temporal and large spatial
time scales. The network promotes syn- also encourage studies that are inte- scales decided to form the Internation-
thesis and comparative research across grated across a range of disciplines: al Long-Term Ecological Research
sites and ecosystems and among relat- physiological ecology, population biol- (ILTER) network. They recommended
ed national and international research ogy, community ecology, and ecosys- developing a worldwide program to
programs. The National Science Foun- tem science. facilitate communication and the shar-
dation established the program in 1980 Global scientific interest in develop- ing of data. As of May 2000, 21 countries
to support research on long-term eco- ing long-term ecological research pro- had formed national LTER programs
logical phenomena in the United States. grams is expanding rapidly, reflecting and joined the ILTER network. Ten
The network now consists of 24 sites the increased appreciation of their more are actively pursuing the estab-
representing diverse ecosystems and importance in assessing and resolving lishment of national networks, and
research emphases, including arctic complex environmental issues. In 1993, many others have expressed interest in
tundra (Toolik Lake, Alaska), southern the U.S. LTER network hosted a meet- the model.

going on for much longer. The oldest experiment in ecol- processes responsible for community structure. One
ogy is the “Park Grass Experiment” of nutrient addition extreme view is that communities are highly predictable
and mowing on pasture communities at the Rothamst- entities in which species are tightly interlinked. An
ed Experimental Station (now the Institute for Crops opposing view is that communities are chance group-
Research-Rothamsted) in Hertfordshire in the United ings of species that are shaped by abiotic factors and
Kingdom, which as been ongoing since the 1850s. Stud- historical contingency. The current consensus lies
ies on the Swedish island of Öland, led by Eddy van der between these views. Deterministic and chance events
Maarel and conducted since the 1960s, have included and biotic and abiotic processes are all considered
both descriptions of vegetation dynamics and experi- important.
mental studies of factors maintaining species diversity. Communities can be described by several measures.
Forest dynamics have been studied at Hubbard Brook The simplest of these is species richness, the number of
Experimental Forest in the White Mountain National species in a community. Other indices of species diver-
Forest, near Woodstock, New Hampshire, since 1963. sity take into account both the number of species and the
Hubbard Brook is now part of the Long-Term Ecologi- relative abundance of each species. Relative abundance
cal Research network that was initiated in 1980 (Box can be measured as numbers of individuals, frequency,
12C). cover, or biomass, using a variety of techniques. Phys-
iognomy, the general form of the vegetation, is another
community property that can be measured. These meas-
Summary ures are used to describe community patterns in time
A community is a group of organisms that coexist in and space. Describing patterns is a first step in deter-
space and time and interact with one another. Ecologists mining which processes are responsible for shaping the
have long debated the relative importance of the plant communities that we see around us.
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Additional Readings
Classical References Contemporary Research
Clements, F. E. 1916. Succession. Carnegie Institution of Washington, Hoagland, B. W. and S. L. Collins. 1997. Gradient models, gradient analy-
Washington, DC. sis, and hierarchical structure in plant communities. Oikos 78: 23–30.
Curtis, J. T. 1959. The Vegetation of Wisconsin. University of Wisconsin Lande, R. 1996. Statistics and partitioning of species diversity, and simi-
Press, Madison, WI. larity among multiple communities. Oikos 76: 5–13.
Gleason, H. A. 1926. The individualistic concept of the plant association. Lawton, J. H. 1999. Are there general laws in ecology? Oikos 84: 177–192.
Bull. Torrey Bot. Club 53: 7–26.
Mueller-Dombois, D. and H. Ellenberg. 1974. Aims and Methods of Veg- Additional Resources
etation Ecology. John Wiley & Sons, New York.
Krebs, C. J. 1989. Ecological Methodology. Harper Collins, New York.
Magurran, A. E. 1988. Ecological Diversity and Its Measurement. Prince-
ton University Press, Princeton, NJ.