Vous êtes sur la page 1sur 23
The Role of Competition in Plant Communities in Arid and Semiarid Regions Author(s): Norma Fowler Reviewedy : Annual Reviews Stable URL: http://www.jstor.org/stable/2096990 . Accessed: 22/11/2011 04:40 Your use of the JSTOR archive indicates y our acce p tance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. Annual Reviews is collaborating with JSTOR to digitize, preserve and extend access to Annual Review of Ecology and Systematics. http://www.jstor.org " id="pdf-obj-0-2" src="pdf-obj-0-2.jpg">

The Role of Competition in Plant Communities in Arid and Semiarid Regions Author(s): Norma Fowler Reviewed work(s):

Source: Annual Review of Ecology and Systematics, Vol. 17 (1986), pp. 89-110

Published by: Annual Reviews

Accessed: 22/11/2011 04:40

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.

The Role of Competition in Plant Communities in Arid and Semiarid Regions Author(s): Norma Fowler Reviewedy : Annual Reviews Stable URL: http://www.jstor.org/stable/2096990 . Accessed: 22/11/2011 04:40 Your use of the JSTOR archive indicates y our acce p tance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. Annual Reviews is collaborating with JSTOR to digitize, preserve and extend access to Annual Review of Ecology and Systematics. http://www.jstor.org " id="pdf-obj-0-36" src="pdf-obj-0-36.jpg">

Annual Reviews is collaborating with JSTOR to digitize, preserve and extend access to Annual Review of Ecology and Systematics.

http://www.jstor.org

Ann. Rev. Ecol. Syst. 1986. 17:89-110 Copyright ? 1986 by Annual Reviews Inc. All rights reserved

THE ROLE OF COMPETITIONIN PLANT COMMUNITIESIN ARID AND SEMIARID REGIONS

Norma Fowler

Departmentof Botany, University of Texas, Austin, Texas 78712

INTRODUCTION

The importance,and even the existence, ecosystems has often been questioned.

of competitionamong plants in arid An influential statement of Shreve

(113) asserted that interspecific competition does not occur in deserts, and Went (145) denied that competition between desert plants occurs at all. Neither providedevidence for his assertions, althoughShreve pointed out the diversity of habits and phenologies found among desert species. He may have

been responding to the strong emphasis placed on competition by Clements and his followers (e.g. 27). The importanceof competition in naturalcom- munities has recently been debated (28, 109, 127). These reviews suggested that terrestrialplant communities are among the communitiesin which com- petition is relatively important.However, the majorityof studies upon which this conclusion is based were made in humid regions. Grime (53) suggested

that competition is less important in "high stress" habitats (in which he included dry habitats), but he presented little evidence from true arid or semiarid environments. This paper reviews the available evidence for competition in plant com- munities in aridand semiaridregions; as is demonstrated,competition certain- ly occurs in these communities and involves many different species. In several instances it appearsto be importantin the determinationof community structure.Competition may be less frequentin these communities,though not

less important on that account. This review also

addresses several other

questions concerningthe role of competitionin these communities, including:

the role of competition in determiningthe absence, or presence and abun- dance, of species in a community, and their spatial arrangement;which

89

0066-4162/86/1 120-0089$02.00

90 FOWLER

species are in competition with one another;and at which stages in the life cycle they experience competition.In addition,I review currentknowledge of the mechanismsof competition and the ways in which plants partitionniches in these communities, as well as facilitation of one plant by another, and succession. Finally, potential directions of future work are discussed.

DEFINITIONS

  • I have followed Bailey (6) in my use

of the terms aridand semiarid,especially

his Figures 3.2 and 3.3. The arid and semiarid regions of the world thus

defined are collectively nearly identical to Walter's (140) zonobiomes III

(subtropicalarid) and VI (aridtemperate with a

cold winter). This definitionis

somewhat broaderthan that used by Noy-Meir (93, 94) in previous articles in these volumes. Arid and semiaridregions are, from the viewpoint of a plant ecologist, those in which an insufficiency of water frequently limits or prevents plant growth or survival. Since water requirementsare partly a function of transpirationrates, which in turnare a function of temperature,as are rates of evaporationfrom the soil, the degree of insufficiency of water in an ecosystem is a function of temperatureas well as of rainfall (6). Arid regions are also generallycharacterized by very wide fluctuationsin precipita- tion between years (93, 140). The word competitionwill be used, unless otherwise noted, in the sense of negative interference,i.e. any direct or indirectnegative impact of one plant on another (58). Therefore, the use of the word does not imply that mech- anisms other than competition for resources (for example, allelopathy, or the harboringof predators)have been eliminated from consideration. There is some evidence that plants may facilitate each other's survival and growth in arid regions, i.e. positive interference, and this is also discussed.

EVIDENCE FOR THE OCCURRENCEOF COMPETITION Studies of Spatial Pattern

The majority of tests for the occurrence of plant competition in arid or semiaridregions have been studies of spatialpattern. In many cases evidence has been found that competitionoccurs and is important,at least in determin- ing spatial pattern. The hypothesis underlyingthese studies is that competition among neigh- boring plants will lead to density-dependentgrowth and survival, hence plants that are closer together will be smaller and more likely to die. Competition will therefore convert clumped (aggregated)distributions of plants into ran- dom ones, and randomdistributions into regular(i.e. overdispersed)ones. A

PLANTCOMPETITION IN ARIDREGIONS

91

variety of analytical methods have been used to determine whether plant

distributionsare clumped, random, or regular(26, 102, 103). Correlationsof

the distance between neighboringplants and their size have also been calcu-

lated (101, 102); positive correlations are considered to be the result of

competition. Some of the older studies use the variancevs block size methods

of Grieg-Smith (51).

While some

studies of spatial pattern have reported regular plant dis-

tributionsin at least some sites, reportsof randomand clumped distributions

are much more common (Table 1). In contrast,positive correlationsbetween

plant size and the distance between plants (Table 2) have been found by most

of the workers who report testing for them (but see 55). Such positive

correlationsare not limited to pairs of conspecific plants, but also have been

found between neighboringplants of different species (157, 158).

Recently, the interpretationof regularspacing as evidence of competition

has been challenged. Ebert & McMaster (34) demonstratedthat failure to

distinguish individuals growing close together as separate individuals in-

troduces a bias towards regular dispersion and towards positive

correlations

between plant size and the distance between neighboringplants. While these

authors only addressed the conclusions of Woodell et al (153; see also 68),

their warning is potentially applicableto all the studies cited in Tables 1 and

  • 2. Furtherinvestigation of this problem is clearly needed, especially of how

common coalesced individuals are and how the phenomenon may be in-

corporated into tests of statistical significance. Nevertheless, it seems rash

and unnecessaryto dismiss all of the earlier findings of regulardistributions

on this basis.

A more fundamentalissue in the interpretationof studies of spatial pattern

is that, while a regular distributionof plants may reasonablybe ascribed to

competition, the absence of such a distributionis not evidence for the absence

of

competition. As many authorshave pointed out, both spatialheterogeneity

in

the environmentand restrictedseed dispersalcan overridethe tendency for

competitionto produceregular distributions of plants and positive correlations

 

between plant

size and distance apart. Measures of pattern are also scale-

dependentand hence depend upon choice of quadratsize and other sampling

decisions. Therefore, incorrectchoice of sampling units may result in failure

to detect existing patterns.Positive correlationsbetween the size of competing

plants and their distance apartmay also be absent simply because each plant

competes with many different neighboring plants (40,

117). In fact, the

frequency with which

significant correlationsare found in desert communi-

ties, as comparedwith more mesic ones (e.g. 139), indicatesthat desertplants

usually compete with relatively fewer neighborsthan do plants in more mesic

environments (40).

92

FOWLER

cn

N

O

~~~~~0000

-

C~~~~~~~~~~:

m

m

CZmm,

C~~~~~~~~C CZ

C"

CZ

Z

e

cE

;

;

e

r

E

H

E

t

X

s

?

s

s,,~~~~~~~~~~~~~:3

;:

E

r

z

r

r

E rECZ

E

E

Y

Y;;

E=~~~C

s,

U

u

9C9

c S ! S9

H

v:

f

fo:

o:

e

o:o:

E

tr

o:

t

o:o:

E

w

o:

8

m

X

T

i~~~~~~~~C

t

X

X

X>

PLANT

COMPETITION

IN ARID REGIONS

06,06,

93

00

C6

C6

0

CN

0

Nt

00

CZ

;:I

CZ

CZ

C6

+--

C6

+--

0

>

CZ

CZ

CZ

CZ CZ

CZ

CZ

CZ

CZ -

Cl.

-

;:I

-

;:3

-

CZ

CZ 0

;:I

-t:)

CZ

CZ

CZ

.-

+--

CZ

CZ

CZ

> 0

CZ

CZ

=

>

CZ

>

CZ

CZ

CZ

CZ

CZ

-

CZ

>

0

u

40-

1j

zr

Q)

to

tz

to

Q.

Q.

to

to

94

FOWLER

Table 2

Summaryof studies that have found positive correlationsbetween plant size and the

distances between pairs of individuals

 

Species

Habit

Location(s)

Reference(s)

Ambrosia dumosa

shrub

Sonoran, Mojave

Deserts

101

Atriplex polycarpa

shrub

Mojave Desert

101

Calandrinia arenaria

forb

Chile

17

Carnegiea gigantea

cactus

Sonoran Desert

158

Chrysothamnuspaniculatus

shrub

Mojave Desert

101

Croton menthodarus

shrub

Ecuador

118

Encelia

farinosa

shrub

Sonoran Desert

35

Eriogonum inflatum

forb

Mojave Desert

155

Fouquieria

splendens

shrub

Sonoran Desert

158

Franseria deltoidea

shrub

Sonoran Desert

158

Hilaria rigida

grass

Sonoran Desert

91

Larrea tridentata

shrub

Sonoran, Mojave

Deserts

101, 108, 158

Opuntia acanthocarpa

cactus

Mojave Desert

157

Opuntiafulgida

cactus

Sonoran Desert

158

Opuntia ramosissima

cactus

Sonoran, Mojave

Deserts

101, 157

Yucca schidigera

succulent

Mojave Desert

157

Studies of the Direct Effects of Competition

A relatively small number of studies of competition in arid and semiarid

regions have

examined the direct effects of competitionupon individualplants

and plant populations. Most of these studies involved manipulatingplant

densities by the removal of individuals, and each found some evidence of

competition.

Friedmanand his coworkersconducted a

series of studies of competitionin

the Negev Desert. In one, seedlings of the

shrubArtemisia herba-alba were

transplantedaround the codominantshrub Zygophyllum dumosum (42). Both

survival and growth of seedlings were lower where seedlings were planted

closer to adult shrubs, indicating that competition occurred. Competition

between seedlings of A. herba-alba and adults of the same species was

examined in another study (45) by following naturallyoccurring seedlings.

Few seedlings emerged under adult canopies, and their death rate was higher

there, again indicating competition. Densities of naturallyoccurring annuals

were lower and mortality rates higher near adult A. herba-alba than in the

open, but not near adults of Z. dumosum.However, numbersand biomass of

annuals increased following the removal of either shrub species (46). Two

varieties of the annualMedicago laciniata

had more fruits per plant when all

nearbyplants were removed (44), but only when both varieties were watered.

In the greenhouse the

variety that "lost" in intervarietalcompetition used

water more efficiently and had a higher seed set (43).

Another series of studies was conducted in a southern Arizona desert

PLANTCOMPETITION IN ARIDREGIONS 95

grassland, at the Santa Rita Experimental Range. Removal of Prosopis

juliflora

(mesquite) led to an increase in annual and perennial grasses, es-

pecially

in Trichachne californica (cottongrass), Eragrostis lehmanniana,

and species of Bouteloua (20,

66, 96). Removal of B. eriopoda, T. californi-

ca, or Muhlenbergiaporteri, anotherperennial grass, increased the survival

of P. juliflora seedlings (48). Additionof E. lehmannianareduced the density

of the native perennialgrasses, perhapspartly due to selective grazing of the

natives (66). Removal of the subshrubAplopappus tenuisectus (burroweed),

of T. californica, or of annual grasses as a group, demonstratedthat each

affected the growth of the other two, with the single exception that the annual

grasses did not appearto interferewith the growth of A. tenuisectus(19, 96).

In similar vegetation in southernAfrica, the

removal of all

herbaceousplants

increased rates of establishmentand growth of two Acacia

species but not of

two other woody species; the removal of the latter two species increased

herbaceous cover (71).

Sagebrush (Artemisia tridentata) interferes with the growth of

perennial

grasses in the American intermontanedesert; sagebrushremovals led to an

increase in the dry weight of individuals of several native grasses (105).

Similarly, removal of all shrubs(primarily Larrea tridentata)in a Chihuahuan

desert site led to a significant increase in the cover of the perennial grass

Muhlenbergiaporteri (148). Clippingor removal of grasses improvedthe rate

of survival of seedlings and the growth and survival of 2-yr-old plants of the

shrub Gutierreziamicrocephala (97, 98, 100). The removal of adults of this

species increasedthe survival and growth rates of conspecific 2-yr-old plants

(97, 100).

Another species of Gutierreziaexcludes the forb Machaeranthera

canescens

from some sites (99). Removal of all grasses aroundindividuals of

the grasses Stipa

neomexicana and Aristida glauca increased recruitment,

survival, growth, and reproductionof both species (56).

Inouye (60) demonstrateddensity-dependent reduction, not only of survival

and growth but also of germination in desert annuals, by experimentally

thinning stands of annuals in the Sonoran desert, and by observing natural

stands of different densities. In at least one of two sites, the interactionwas

primarilyintraspecific, involving a single dominantspecies. In anotherstudy

(61), the effects of thinning were reported to involve only growth and

fecundity, not survival. Klikoff (69) comparednaturally occurring stands of

different densities of the annual Plantago insularis in the Sonoran desert.

Stands of lower initial density had higher survival rates when watered mod-

erately. Removal of all other species increasedthe numberand size of plants

of the annual Salsola inermis (87).

Some studies have examined the effects of competition upon the water

status of the affected plants or upon soil water content, as well as upon

measuresof plant size or survival. Experimentalremovals of Larrea tridenta-

96 FOWLER

ta and/orAmbrosia dumosa aroundtarget plants of each species improvedthe

water potential of individualsof the other species (38, 39). In a monospecific

stand of Enceliafarinosa, the removal of all neighboringplants improvedthe

water status of

the remainingplants, as well as plant size and seed set (35).

Survival rates of transplantedseedlings of the grass Bromus setifolius in-

creased away from shrubs where soil water content was greater (122). The

removal of all vegetation around the grass Hilaria rigida in monospecific

stands increasedsoil waterpotentials, plant water status, and plant size (104).

EVIDENCE FOR THE FREQUENCY OF COMPETITION

The extremely variable climates characteristicof arid and semiarid regions

would lead one to expect that competition would be a relatively infrequent

event there. Variableclimates will producefluctuating resource levels, which

in turn could cause the size of populationsfrequently to decrease below the

level at which competition for resources would occur. Wiens (149) has

advancedthis argumentwith respect to bird communities, and I found that, in

a subhumidbut water-limitedgrassland, a perennialgrass populationwas so

reduced

by a moderate drought that density-dependenteffects were greatly

reduced or eliminated (41). Perhapsonly after populationsincrease during a

series

of "good" years, and the carrying capacity of the environment then

drops in a "bad"year, will competition for resources become intense. (It is

importantto bear in mind, however, that competitionmay be infrequentand

yet play an importantrole in structuringcommunities and regulating pop-

ulations.)

Despite its

plausibility, the existing evidence does not

supportthis hypoth-

esis. Almost

all of the experimental studies just described were short-term

(<5 yr, usually <3 yr), and all found some evidence of competitionoccurring

during the course of the experiment. This implies that competition is a

relatively frequent phenomenonin these communities. However, two of the

studies were able to demonstratecompetition only in watered plots (44, 69).

Since individualsof most of the species whose spatialpattern has been studied

are long-lived, the existence of regular spatial patternssays little about the

frequency of competition involving these species.

THE EFFECTS OF COMPETITIONON COMMUNITY COMPOSITION

A few studies have demonstratedthat competitioncan restrictthe

distribution

of a species. Gurevitch (56) showed

that the distributionof the grass Stipa

mexicana is restricted to ridgecrests in a site in Arizona by interspecific

competition. Because this species respondedmore positively to the removals

PLANTCOMPETITION

IN ARIDREGIONS

97

of competitorslower down the slope than it did to removals on the ridgecrest

(to the extent that estimated population growth rates after removals were

almost equal in all locations), she concluded that competition restrictedthe

distributionof S. mexicana.

The forb Machaerantheracanescens is absent or

rarein all but disturbedsites

due to competitionfrom Gutierreziamicrocepha-

la (via herbivory;99 and see below). Competitionmay restricttwo varieties

of the annual Medicago laciniata to north and south slopes in the Negev

Desert, respectively. Relative fecundities were greaterfor each variety on its

usual slope, but only in undisturbedvegetation (44).

The effects of competitionupon the abundancesof species present in a site

are also little known. Few studies have looked directly at the impact of

competition upon populationsizes (19, 20, 66, 96, 148). Other studies have

only measured the effects of competition on individuals, although one can

infer that these effects must often result in the reduction,if not the regulation,

of populationsize. The natureand magnitudeof

interspecific competition upon population sizes

the effects of both intra-and

and population dynamics in

arid and semiarid regions remain to be investigated.

STAGES OF THE LIFE CYCLE AFFECTED BY COMPETITION

Studies of patternsuggest that competition may affect a plant throughoutits

adult life, althoughperhaps only intermittently.Anderson and

coworkers (5,

74) and Grieg-Smith & Chadwick (52) found that smaller (hence, younger)

plants had a clumped distribution,whereas older plants had a more random

one. This they interpretedas evidence for competition among young plants,

with "relativelymore individualseliminated from high density than from low

density phases" (5). In these studies, the failure to reach a regulardistribution

was

ascribed to environmentalheterogeneity, not

to the absence of competi-

tion among older plants. Phillips & MacMahon(101) divided populationsof

Larrea tridentata, Ambrosia dumosa, and several other shrubs into size

classes, and found that 20 of 22 were consistentwith the expected trend, from

small to large plants, of clumped-*random-->regular, although in no case did

the different size classes of a single populationdemonstrate all three types of

distribution. In Opuntia bigelovii (79), Tidestromiaoblongifolia (54), and

Eriogonum inflatum (155), living and dead individuals were more clumped

than living individualsalone, which suggests adultdensity-dependent mortal-

ity.

Measures of the direct effects of competition demonstratedcompetition

between adult perennials (35, 38, 39, 56), from adult perennials against

seedling perennials (42, 45, 97, 98, 100) and against seedling annuals (46),

among seedling annuals(60, 61, 144), and even among seeds (60). Competi-

98

FOWLER

tion may affect survival, growth, or reproduction(see above). There are too

few studies to generalize, but the existing results suggest that the effects of

competition upon a

species should be looked for in

all stages of the life cycle;

competitive effects

should not be assumed a priori to be absent at any stage.

MECHANISMS OF COMPETITION Competitionfor Water

Most plant ecologists working in arid or semiaridregions have assumed that

the principal form of competition among plants is competition for water.

Perhaps because it appears to be obvious, the number of studies directly

supportingthis hypothesis is relatively low, althougha large body of work has

demonstratedthat plants of arid and semiarid regions are often under water

stress (24). Watering generally increases rates of growth and survival, con-

firming that it is a limiting resource(e.g. 44, 61, 69, 73). A numberof studies

have found that a reduction in the intensity of competition, in addition to

increasing plant

size, survival, or fecundity, also is associated with an im-

provementin plant water status (35, 38, 39, 104) or an increase in soil water

content (48, 87, 96, 104, 122).

Raising soil water content shifted the outcome

of competitionbetween Salsola

kali and perennialgrasses in

favor of the

latter

(1) and led to an increase in the abundancesof warm season

grasses and forbs

and a decrease in succulents in a dry grassland (73).

It has been suggested that competitionfor water is most intense in deserts

with an intermediatelevel of rainfall;this theory is based upon the occurrence

of

clumped, random, and regularlydistributed populations of Larrea tridenta-

ta

(67, 153; but see 9). However, Anderson(4) pointed out that since density

decreases as rainfall does, the water available per plant does not necessarily

decrease. Walter (140) presented evidence that the water supply

per unit of

"transpiringsurface" is relatively constant.

Competitionfor Minerals

Fertilizationwith nitrogenincreased the size of winter annualsin the Mojave

desert, but phosphorus did not (152). Nitrogen addition also increased the

biomass of most species in a dry grassland, where its principaleffect was to

magnify the results of differentwatering treatments (73). Nitrogen levels had

no effect on the outcome of competition between the perennial grasses

Bouteloua gracilis and Agropyron smithii (15). Caldwell et al (21) demon-

strated that the shrub Artemisia tridentata took up much more phosphorus

from the root space it shared with Agropyron spicatum than from the root

space it shared with Agropyrondesertorum, and that A. desertorumtook up

more phosphorusthan did A. spicatumwhen competingwith A. tridentata.As

the authorswere careful to point out, these results do not imply that phospho-

rus is the only, or even the most important,resource for these plants.

Allelopathy

PLANTCOMPETITION

IN ARIDREGIONS

99

Aqueous extracts of Partheniumincanum (13, 14), Encelia farinosa (13, 49,

50, 86), Ambrosia dumosa (85, 86), Thamnosmamontana (86), Artemisia

herba-alba (46), and Larrea tridentata(70) have been shown to have detri-

mental effects on one or more plant species. Extracts of L. tridentatawere

found to have no deleterious effects on

its own germinationor early growth

(8, 70). The relevance of the

toxicity of

aqueous extractsto plant growth and

survival in the field has been doubted. Bonner (13) failed to find any toxicity

in the soils of fields in

which P. incanumhad been grown; he concluded that

the productionof toxic substancesis relevantonly in greenhousesand perhaps

crowded nurseries. Muller & Muller (86) found no correlationbetween the

degree of toxicity of the aqueous extractfrom a shrubspecies and patternsof

herb species' occurrence under those shrub species; they concluded that

"toxins are ineffectual as factors

in competition between plants" in deserts.

Herbivory

The harboringof predatorsor pathogens is anotherway in which plants may

interfere with one another; hence it is a form of competition in the broad

sense. The composite shrub Gutierrezia sarothrae excludes the composite

forb Machaerantheracanescens from some

sites by harboringa grasshopper

that eats both species; transplantsof the forb survivedonly in exclosures (99).

WHICH PAIRS AND GROUPS OF SPECIES COMPETE?

The relative strengthsof intra- and interspecific competition are relevant to

the problems of species coexistence and stability (28), for example, to

identifying competitively dominant species. Interspecific competition was

found to be strongerthan intraspecificcompetition in

an experimentalstudy of

Larrea tridentata and Ambrosia dumosa, although other factors were more

important in determining these species' abundances and distributions(38,

39). The limited evidence from studies of pattern on this point, however,

supportsthe opposite generalization:Interspecific competition is weaker than

intraspecific (157,

158). The degree of reciprocity of competitive rela-

tionships would also be of interest, if relevant data existed.

The presence and relative intensity or absence, of interspecificcompetition

among different component species of a community is also of interest,

because these cast considerablelight upon the niche structureof the communi-

ty. Few studies of competition, however,

have included several species from

one community. Yeaton and coworkers (157, 158) comparedthe degree of

correlationof distances between plants and their sizes, between species pairs.

In the Mojave desert (157) competitionoccurred among all three pairs of two

Opuntia species and a Yucca, apparentlyat equal intensity. However, for

individualsof a given size, pairs of plants of differentspecies of Opuntiawere

100

FOWLER

closer together than conspecific pairs, which Yeaton et al interpretedas

suggesting some differences in root systems and hence niche separation. In

the Sonoran desert (158), four of the nine interspecific pairs tested had

significant positive correlationsbetween size and distance, in addition to all

five correlationsbetween conspecific pairs.

The experiments conducted at the

Santa Rita ExperimentalRange also

involved several species from one community. The shrub Aplopappus

tenuisectus, the perennialgrass Trichachnecalifornica, and the annualgrass-

es as a group,

were all found to compete with one another, except that the

annual grasses

did not affect T. californica (19). These relationshipsamong

  • A. tenuisectus and the

grasses were judged consistent with observed pheno-

logical patterns. Prosopis juliflora reduced the growth and abundance of

several grasses (20, 66, 96), and in turnthese grasses reducedthe survival of

  • P. juliflora (48). However, in similar African vegetation, competition be-

tween herbaceous and woody species was apparentlynot reciprocal (71).

Here again, too few studies have been made to generalizewith confidence,

but those to date

generally support the competitive relationshipsthat have

been inferredfrom phenological patternsand root location. They also

suggest

that despite these niche differences, competitionamong many or even

most of

the species pairs of a community probably occurs.

NICHE SEPARATION

The separation or differentiation of niches among species is expected to

reduce the intensity of competitionamong them; it may therebypromote their

coexistence. Niche separation among plants primarily takes the form of

separationof resource use in space and/or in time. The plant species of arid

and semiarid regions representa very wide range of adaptationsthat tend to

separate their use of water (113,

114). Many of

these adaptations have

received detailed study by physiological ecologists (24, 120). Only a brief

outline of potential mechanisms of niche separationcan be given here.

Phenology

Plants may separate their use of water by being physiologically active at

different times (113, 119, 120). Ephemerals, including annuals, algae, and

lichens, grow only when water conditions are favorable. In areas with two

rainy seasons, such as the Sonoran desert, there may be two separatesets of

ephemerals (119, 144). Furthermore,the relative abundances of different

annual species will vary from year to year depending upon the amount and

timing of rainfall (12, 16, 62, 92, 113, 144). Tidestromiaoblongifolia seems

to be an 'ephemeralperennial,' growing rapidlywith little waterconservation

and then dying when the soil water in a temporarywash channel is exhausted

(54).

 

PLANTCOMPETITION IN ARIDREGIONS

101

Some perennials, including most perennial grasses (19, 121),

grow only

during relatively

favorable seasons.

Leaves die back or are shed

during dry

periods, and this reduces transpirationrates and water uptake. These species

may differ in their use of small rainfalls(106, 107). Otherperennials maintain

leaves, photosynthesize, and absorband transpirewater even duringvery dry

periods. Both groupsof perennialsmay furtherseparate their periods of active

growth by being warm-seasonor cool-season species, often (but not always)

following a C4

VS

C3

grouping (12,

15, 33,

56,

64,

65,

80, 81).

With stored reserves of water, succulents fall into none of these groups.

Their primary period of water uptake is limited to the time immediately

following rains, while their period of active growthmay extend much longer.

Finally, along watercourses some perennials (phreatophytes)depend upon

water supplies that are more or less continuously abundant. They may be

evergreen or deciduous, with a variety of phenological patterns (89).

Separation in Horizontal Space

It is outside the limits of this article to review the many studies of plant

distributionin relation to local environment. Like other plant communities,

those of arid and semiaridregions are characterizedby the separationof plants

according to microtopographicand other environmentalvariation. Washes

and

other drainage features (108), dunes (32), existing shrubs (see below),

and the relative amounts of sand and clay in the soil (119), are of particular

importance in controlling local distributionsof

species.

Rooting Zones

Excavations of roots of a number of species

have demonstratedthat the

species of arid and semiarid regions are characterizedby several different

patternsof root distribution(23, 25, 84, 119, 158).

Cacti and other succulents

typically have shallow rooting zones. The roots of perennialsoccupy rooting

zones that are both wider and deeper than those

of the annuals, and different

perennials may root at different depths (e.g. 158). Phreatophytesare often

very deeprooted. Woody plants tend to root more deeply than grasses, and the

resulting separationof water use can be sufficient to permitcoexistence (121,

137, 138).

The extent to which the soil is fully occupied with roots is usually con-

sidered to be an indicatorof the importanceof competition. Gulmonet al (55)

found that cactus roots of adjacent individuals met; Bustamente et al (18)

made the same observationof a Chilean shrub. Otherinvestigators, however,

have reportedfinding space between adjacentroot systems (23, 25). It is clear

that the roots of almost all individuals extend much furtherthan their cano-

pies; the apparentseparation of plants is very misleading.

102

FOWLER

FACILITATION

The apparentfacilitation of the establishmentor growthof other plant species

by woody perennial shrubs in arid regions has been observed by number of

authors (e.g.

112, 113). Osborn et al (95) noted that annuals were most

abundant in the mounds of sand around the bases of Atriplex vesicaria in

Australia. Went (143) described a complex set of associations between an-

nuals and shrubs in the Mojave and Sonorandeserts, which Muller (85) later

simplified to two groups of annuals-those that are shrub-independentand

those that are shrub-dependent.In contrast, competition, not facilitation,

between shrubs and annuals occurs in the Negev desert (46).

The association of

annuals with woody shrubshas been ascribedto higher

soil organic content, shading (which could cause lower rates of evaporation

and transpiration),the trappingof windblown seeds, bird dispersal of seeds,

and protectionof seeds or seedlings from predation(86, 95, 112). Muller (85,

86) determinedthat the shrub-dependentannuals grow abundantlyonly be-

neath shrubspecies that accumulatea mound of organic matterunderneath by

trapping wind-blown material to add

to their own dead shoots. The shrub-

dependent annuals were also found to

grow abundantlyin areas with tempo-

rarily high

levels of organic matterbut without shrubs. Halvorson & Patten

(57) found that the total biomass, but not the density, of annualswas greater

under shrubsthan in the

open, especially undershrubs with

a relatively high,

open canopy; this implies that ameliorationof the physical environment,not

seed dispersal, was the cause of the relationship.Plant litter has been shown

to aid the establishmentof several annual species (37).

Associations between woody shrubsand cacti have also been noted. Shreve

(112) reportedthat Carnegiea gigantea (saguarocactus) often grows beneath

various trees and shrubs. This

association has since been described by a

numberof authors(78, 88, 123, 124, 125, 126, 129, 130). Shrubshave been

described as providing young C. gigantea with protection from grazing,

trampling,high temperatures,freezing, and drought;rocks also performall of

these functions (88, 123, 124, 125, 129, 130). The transplantexperiments of

Turner et al (129, 130) demonstratedthat both shade and protection from

rodents are necessary for seedlings of C. gigantea to survive. Using a model

that predictedtissue temperature,Nobel (90) confirmedthat nurse plants can

protect cactus plants <2m in height from freezing.

However, the presence of C. gigantea is associated with an increase in the

proportionof dead branchesin at least one shrubspecies; this suggests thatthe

relationshipis one of facilitationfor the cactus but competitionfor the shrub

(75). This relationship should, therefore, produce a constantly changing

mosaic, as cacti replace shrubsand then die, to be replacedin turnby shrubs.

Theoretically, such a relationship could produce oscillations in population

PLANTCOMPETITION IN ARIDREGIONS 103

size (131), but the long life of the cactus and the importanceof occasional,

lethal freezes in determiningpopulation sizes of this species (125, 126) make

such oscillations unlikely. Based upon observations of the distributionand

"vigor" of individuals, Yeaton (156) postulated that Opuntia leptocaulis

preferentiallyestablished underLarrea tridentata, thus reducing that plant's

vigor and eventually replacing it, until finally Opuntia leptocaulis itself

succumbed to soil erosion and rodent burrowing.

In one instance, a cactus, Opuntiafulgida, was shown to be a sheltering

plant, the bed of spine-coveredjoints beneath it providingprotection for two

smaller species of cacti by reducing predationon them (76). Finally, plants

other than annuals and cacti may be facilitated. The seedlings of the semi-

shrub Gutierreziamicrocephala are protectedfrom predationby neighboring

adults of the same species; their rate of survival decreased when the adults

were removed (97). Seedlings of the small tree Cercidium microphyllum

suffered less herbivory-causedmortality under other perennials than in the

open (77).

SUCCESSION

There has been considerabledebate as to whether succession occurs in arid

and semiaridregions. If by succession one

means an orderly, naturalseries of

changes in vegetation following disturbance,then succession certainlyoccurs

(e.g. 2, 30, 63, 72, 83, 111, 132, 133, 134, 135, 141, 142). In most places,

however, areas of disturbanceare colonized by species alreadypresent in the

community, often growing in washes or other small disturbances(111, 113,

134, 142). Only the relative abundancesof the species are altered. Therefore,

if a strict definition of succession is used, it may be said not to occur in arid

and semiarid regions (83).

Some authorshave assumed that competition from later successional spe-

cies reduces or eliminates populationsof early successional species. This has

been demonstrated in the case of the early successional summer annual

Salsola inermis (87). Succession may involve the facilitationof later species

by earlier ones (sensu 29), as it does, for example, in the Mojave lake beds,

where "the accumulationof soil and the redistributionof minerals by plants

permit a primaryplant succession to occur" (135). Or the plants may them-

selves neither promote nor retardvegetational change (e.g. 63, 83).

Grazing is sometimes considered a disturbance, and the changes which

follow its cessation, succession. The effects of grazing and of the cessation of

grazing have been documented for many plant communities of arid and

semiaridregions (e.g. 22, 31, 32, 36, 47, 59, 115, 116, 146, 147, 150, 151).

Studies conducted at the Santa Rita ExperimentalRange, described above,

104

FOWLER

suggest that competition between palatable, "decreaser"species (e.g. most

grasses) and unpalatable,"increaser" species (e.g. Prosopis) plays an impor-

tant role in this form of succession.

CONCLUSIONS AND FUTURE DIRECTIONS

The importanceof competition among plants growing in arid and semiarid

regions has been doubted, but studies suggest that competitionamong plants

is both common and strong enough to be readily detected, both in deserts and

in dry grasslands. The number of such studies is not negligible, even if

allowances are made for the reluctance of authors and journals to report

negative results. It is thereforemy opinion that the occurrenceand potential

importanceof competition in these communities can now be taken as given.

Studies should now be directedtowards determining the circumstancesunder

which competition occurs and its effects upon plant populations.

The existing studies are too few to warrantgeneralizations concerning most

aspects of the nature of competition among plants or its effects on plant

communities. The available evidence has been discussed above; I now

summarizeby outlining four questions that appearto be particularlyinterest-

ing and critical to our understandingof the role of competition in these

communities.

  • 1. How frequently does competition occur?

  • 2. Does competition determine community composition, and, if so, when and how?

  • 3. At which stage(s) of the life cycle are plant populations affected by competition?

  • 4. Which groups of species compete, and how is competitionavoided among co-occurring species?

None of these questions is, of course, uniqueto plantcommunities of arid and

semiaridregions, but the particulardearth of relevant studies in these regions

warrantsdirecting attention to them.

Anothercategory of unanswered,and at this point unanswerable,questions

addresses the similarities and differences among different regions. What

generalizationscan be made about the natureand effects of plant competition

among plant communities of different temperatearid or semiarid regions of

similar climate? Between plant communitiesof temperateand tropicalarid or

semiarid regions? Between arid and humid regions? With regard to the last

question, the relative commonness and importanceof the facilitation of one

plant by another seems to me to be particularlyinteresting.

PLANTCOMPETITION

IN ARIDREGIONS

105

ACKNOWLEDGMENTS

Supportwas provided by NSF grant 8118968. I thank P. Fonteyn, J. Gure-

vich, J. Russell Holman, W. Lauenroth,G. Montenegro, C. H. Muller, M.

Price, 0. Sala, K. Schwaegerle, I. Serey, and N. Waser for comments and

references.

Literature Cited

  • 1. Allen, E. B. 1982. Water and nutrient competition between Salsola kali and 63:732-41

shrubs on a desert plain in Ethiopia. Ecology 49:744-46

two native grass species (Agropyron smithii and Bouteloua gracilis). Ecology

  • 12. Beatley, J. C. 1974. Phenologicalevents and their environmentaltriggers in Mo- jave Desert ecosystems. Ecology

  • 2. Allen, M. F. 1980. Nat-

Allen, E. B.,

ural re-establishment of vesicular- arbuscularmycorrhizae following strip- mine reclamationin Wyoming. J. Appl.

55:856-63

  • 13. Bonner, J. 1950. The role of toxic sub- stances in the interactions of higher plants. Bot. Rev. 16:51-65

Ecol. 17:139-47

  • 14. Gaiston, A. W. 1944. Toxic

Bonner, J.,

  • 3. Anderson, D. J. 1967. Studies on

struc-

substances from the culture media of

Gaz. 106:185-98

ture in plant communities. V. Patternin Atriplex vesicaria communitiesin south-

guayule which may inhibit growth. Bot.

eastern Australia. Aust. J. Bot. 15:451-

  • 15. Boryslawski, Z., Bentley, B. L. 1985.

  • 58 The effect of nitrogen and clipping on

  • 4. Anderson, D. J. 1971. Patternin desert

perennials. J.

Ecol. 59:555-60

  • 5. Jacobs, S.

Anderson, D.

J.,

W.

L.,

Malik, A. R. 1969. Studies on structure

in plant communities. VI. The signifi-

cance of

patternevaluation in some Aus-

tralian dry-land vegetation types. Aust. J. Bot. 17:315-22

  • 6. Bailey, H. P. 1979. Semi-arid climates: Their definition and distribution. In Agriculture in Semi-AridEnvironments, ed. A. E. Hall, G. H. Cannell, H. W. Lawton, pp. 73-97. Berlin: Springer- Verlag

  • 7. Barbour, M. G. 1969. Age and space distribution of the desert shrub Larrea divaricata. Ecology 50:679-85

  • 8. Barbour, M. G. 1973. Desert dogma re- examined: Root/shoot productivity and plant spacing. Am. Midl. Nat. 89:41-57

interferencebetween C3 and C4 grasses. J. Ecol. 73:113-21

  • 16. Brennan, H., Cisse, A. M., Djiteye, M. A., Elberse, W. Th. 1979/1980. Pasture dynamics and forage availability in the Sahel. lsr. J. Bot. 28:227-51

  • 17. Bustamente, R., Serey, I., Guerrero,I.

    • 1978. Competencia intraespecifica en

plantas de las dunas de Quintero. I.

Calandriniaarenaria Cham. An. Museo Hist. Nat. Valparaiso 11:55-60

  • 18. Bustamente,R., Serey, I., Leighton, G.

    • 1981. Estructuraespacial y competencia

intraespecifica en arbustos de desierto:

Alona carnosa Lind. An. Museo Hist. Nat. Valparaiso 14:115-18

  • 19. Cable, D. R. 1969. Competition in the semidesert grass-shrub type as in- fluenced by root systems, growth habits, and soil moisture extraction. Ecology

9.

Barbour,M. G., Diaz, D. V. 1973. Lar-

moisture gradients in the

United States

and Argentina. Vegetatio 28:335-52

50:27-38

Cable, D. R., Tschirley, F. H. 1961.

 

rea plant communities on bajada and

20.

Responses of native and introduced grasses following aerial sprayingof vel-

10.

Barbour, M.

G.,

MacMahon, J. A.,

 

vet mesquite in southern Arizona. J.

Caldwell, M. M., Eissenstat, D. M.,

 

Bamberg, S. A., Ludwig, J. A. 1977.

The structureand distributionof Larrea communities. In Creosote Bush: Biology

21.

Range Manage. 14:155-59

Richards, J. H., Allen, M. F. 1985.

 

and Chemistryof Larrea in New World

 

Competition for phosphorus: Differen-

 

Deserts, ed. T. J.

Mabry, J.

H. Hunzik-

er, D. R. DiFeo, pp. 227-51. Strouds- berg, Penn: Dowden, Hutchinson, &

 

tial uptake from dual-isotope-labeled soil interspaces between shrub and grass. Science 229:384-86

 

Ross

   

22.

Canfield, R. H. 1957. Reproductionand

106

FOWLER

southernArizona. J. Range Manag. 10:

experimental analysis of structurein a

199-203

desert plant community. J. Ecol. 69:

  • 23. Cannon, W. 1911. The Root Habits of Carnegie Inst.

883-96

Desert Plants. Publ. Carnegie Inst. WashingtonNo. 131, Washington, DC:

  • 40. Fowler, N. L. 1984. The role of germination date, spatial arrangement, and neighbourhoodeffects in competi-

  • 24. Chabot, B. F., Mooney, M. A., ed.

tive

interactions in Linum J. Ecol. 72:

1985. Physiological Ecology of North

307-18

American Plant Communities. New York: Chapman& Hall

  • 25. Chew, R. M., Chew, A. E. 1965. The primary productivity of a desert shrub (Larrea tridentata) community. Ecol. Monogr. 35:355-75

  • 26. Clark, P. J., Evans, F. C. 1954. Dis- tance to nearest neighbor as a measure of spatial relationships in populations. Ecology 35:445-53

  • 27. Clements, F. E., Weaver, J. E., Han- son, H. C. 1929. Plant Competition. Carnegie Inst. Washington Publ. 398 Washington, DC: Carnegie Inst.

  • 28. Connell, J. H. 1983. On the prevalence and relative importance of interspecific competition: Evidence from field ex- periments. Am. Nat. 122:661-96

  • 41. Fowler, N. L. 1986. Density-dependent population regulation in a Texas grass- land community. Ecology 67:545-54

  • 42. Friedman,J. 1971. The effect of compe- tition by adult Zygophyllum dumosum Boiss. on seedlings of Artemisia herba- alba Asso in the Negev desert of Israel. J. Ecol. 59:775-82

  • 43. Friedman, J., Elberse, W. Th. 1976. Competition between two desert vari- eties of Medicago laciniata (L.) Mill. under controlled conditions. Oecologia 22:321-39

  • 44. Friedman, J., Orshan, G. 1974. Allopatricdistribution of two varietiesof Medicago laciniata (L.) Mill. in the Negev desert. J. Ecol. 62:107-14

  • 45. Friedman,J., Orshan,G. 1975. The dis-

  • 29. Slatyer, R. 0. 1977.

Connell, J. H.,

tribution, emergence and survival of

in the Negev desert of Israel in relation

Mechanisms of succession in natural communities and their role in communi-

seedlings of Artemisia herba-alba Asso

ty stability and organization. Am. Nat.

to

distance from the adultplant. J. Ecol.

111:1119-44

63:627-32

  • 30. Costello, D. F. 1944. Naturalrevegeta- tion of abandoned plowed land in the mixed prairieassociation of northeastern Colorado. Ecology 25:312-26

  • 46. Friedman, J., Orshan, G., Ziger-Cfir,

    • Y. 1977.

Suppression of annuals by

Artemisia herba-alba in the Negev Des-

ert of Israel. J. Ecol. 65:413-26

  • 31. Crisp, M. D. 1978. Demography and

  • 47. Glendening, G. E. 1952. Some quantita-

survival under grazing of three Austra-

tive data on the increase of mesquite and

lian semi-desert shrubs. Oikos 30:

cactus on a desert range in

southernAri-

520-28

zona. Ecology 33:319-28

  • 32. Danin, A. 1978. Plant species diversity

  • 48. Glendening, G. E., Paulsen, H. A.

and plant succession in a sandy area in

1955.

Reproduction and establishment

the NorthernNegev. Flora 167:409-22

of velvet mesquite as related to invasion

  • 33. Dickinson, C. E., Dodd, J. L. 1976.

of semidesert grasslands. USDA Tech.

Phenological pattern in

the short grass

Bull. No. 1127. Washington, DC:

prairie. Am. Midl. Nat.

96:367-78

USGPO

34.

Ebert, T. A., McMaster, G. S. 1981. Regular patternof desert shrubs:A sam- pling artefact?J. Ecol. 69:559-64

49.

Gray, R., Bonner, J. 1948. An inhibitor of plant growth from the leaves of En- celia farinosa. Am. J. Bot. 35:52-57

35.

36.

Ehleringer, J. R. 1984. Intraspecific competitive effects on water relations, growth, and reproduction in Encelia farinosa. Oecologia 63:153-58

50.

Gray, R., Bonner, J. 1948. Structure determinationand synthesis of a plant growth inhibitor, 3-acetyl-6-methoxy benzaldehyde,found in the leaves of En-

celia farinosa. J. Am. Chem. Soc. 70:

 

Ellison, L. 1960. Influence of grazing on plant succession of rangelands.Bot.

Evans, R. A., Young, J. A. 1970. Plant

 

1249-53

37.

Rev. 26:1-78

51.

Grieg-Smith, P. 1983. Quantitative

Plant Ecology. Berkeley: Univ. Calif.

litter and establishment of alien annual

 

Press. 3rd ed.

 

weed sp.cies in rangelandcommunities. Weed Sci. 18:697-703

52.

Grieg-Smith,P., Chadwick,M. J. 1965. Data on pattern within plant communi-

  • 38. Fonteyn, P. J., Mahall