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The Role of Competition in Plant Communities in Arid and Semiarid Regions

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The Role of Competition in Plant Communities in Arid and Semiarid Regions
Author(s): Norma Fowler
Source: Annual Review of Ecology and Systematics, Vol. 17 (1986), pp. 89-110
Published by: Annual Reviews
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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, of competitionamong plants in arid
ecosystems has often been questioned. 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 pointedout the
diversity of habits and phenologies found amongdesert species. He may have
been respondingto the strong emphasis placed on competitionby 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 communitiesare 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-
munitiesin aridand semiaridregions;as is demonstrated,competitioncertain-
ly occurs in these communities and involves many different species. In
several instancesit appearsto be importantin the determinationof community
structure.Competitionmay be less frequentin these communities,thoughnot
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

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90 FOWLER

species are in competition with one another;and at which stages in the life
cycle they experiencecompetition.In addition,I review currentknowledge of
the mechanismsof competitionand 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 termsaridand 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 (aridtemperatewith a cold winter). This definitionis
somewhatbroaderthan that used by Noy-Meir (93, 94) in previous articlesin
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 functionof 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 generallycharacterizedby very wide fluctuationsin precipita-
tion between years (93, 140).
The word competitionwill be used, unless otherwisenoted, 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 competitionfor 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 competitionamong neigh-
boring plantswill lead to density-dependentgrowthand survival, hence plants
that are closer together will be smaller and more likely to die. Competition
will therefore convert clumped (aggregated)distributionsof plants into ran-
dom ones, and randomdistributionsinto regular(i.e. overdispersed)ones. A

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 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 clumpeddistributions
are much more common (Table 1). In contrast,positive correlationsbetween
plant size and the distancebetween 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 regulardispersion and towardspositive correlations
between plant size and the distancebetween neighboringplants. While these
authorsonly addressedthe conclusions of Woodell et al (153; see also 68),
their warning is potentiallyapplicableto all the studies cited in Tables 1 and
2. Furtherinvestigationof this problem is clearly needed, especially of how
common coalesced individuals are and how the phenomenon may be in-
corporatedinto tests of statistical significance. Nevertheless, it seems rash
and unnecessaryto dismiss all of the earlierfindings of regulardistributions
on this basis.
A more fundamentalissue in the interpretationof studies of spatialpattern
is that, while a regulardistributionof plants may reasonablybe ascribedto
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 tendencyfor
competitionto produceregulardistributionsof plantsandpositive 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 samplingunits 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), indicatesthatdesertplants
usually compete with relatively fewer neighborsthando plantsin more mesic
environments (40).

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92 FOWLER

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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

Atriplexpolycarpa 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 examinedthe directeffects 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 coworkersconducteda series of studies of competitionin
the Negev Desert. In one, seedlings of the shrubArtemisia herba-alba were
transplantedaroundthe codominantshrubZygophyllumdumosum(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 naturallyoccurringseedlings.
Few seedlings emerged underadult canopies, and their deathrate was higher
there, again indicating competition. Densities of naturallyoccurringannuals
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 varietieswere 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

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 IN ARIDREGIONS
PLANTCOMPETITION 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, anotherperennialgrass, increasedthe survival
of P. juliflora seedlings (48). Additionof E. lehmannianareducedthe density
of the native perennialgrasses, perhapspartlydue to selective grazing of the
natives (66). Removal of the subshrubAplopappustenuisectus(burroweed),
of T. californica, or of annual grasses as a group, demonstratedthat each
affected the growthof the othertwo, with the single exception thatthe annual
grasses did not appearto interferewith the growthof A. tenuisectus(19, 96).
In similar vegetation in southernAfrica, the removal of all herbaceousplants
increasedrates 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, removalof all shrubs(primarilyLarrea tridentata)in a Chihuahuan
desert site led to a significant increase in the cover of the perennial grass
Muhlenbergiaporteri (148). Clippingor removalof grasses improvedthe rate
of survival of seedlings and the growth and survivalof 2-yr-old plants of the
shrub Gutierreziamicrocephala (97, 98, 100). The removal of adults of this
species increasedthe survival and growthrates of conspecific 2-yr-old plants
(97, 100). Another species of Gutierreziaexcludes the forb Machaeranthera
canescens from some sites (99). Removal of all grasses aroundindividualsof
the grasses Stipa neomexicana and Aristida glauca increased recruitment,
survival, growth, and reproductionof both species (56).
Inouye (60) demonstrateddensity-dependentreduction,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) comparednaturallyoccurringstands 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. Experimentalremovalsof Larrea tridenta-

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96 FOWLER

ta and/orAmbrosiadumosa aroundtargetplantsof each species improvedthe

water potentialof 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
standsincreasedsoil waterpotentials,plantwaterstatus, andplant size (104).

EVIDENCE FOR THE FREQUENCYOF 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 producefluctuatingresourcelevels, which
in turn could cause the size of populationsfrequentlyto decrease below the
level at which competition for resources would occur. Wiens (149) has
advancedthis argumentwith respectto birdcommunities,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 environmentthen
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 experimentalstudies 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 frequentphenomenonin these communities. However, two of the
studies were able to demonstratecompetitiononly in wateredplots (44, 69).
Since individualsof most of the species whose spatialpatternhas 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 demonstratedthatcompetitioncan 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

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 IN ARIDREGIONS
PLANTCOMPETITION 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 competitionrestrictedthe
distributionof S. mexicana. The forb Machaerantheracanescens is absentor
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 presentin 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 resultin the reduction,if not the regulation,
of populationsize. The natureand magnitudeof the effects of both intra-and
interspecific competition upon population sizes and populationdynamics in
arid and semiarid regions remain to be investigated.

STAGES OF THE LIFE CYCLE AFFECTEDBY

COMPETITION
Studies of patternsuggest that competition may affect a plant throughoutits
adult life, althoughperhapsonly intermittently.Andersonand 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 individualseliminatedfrom high density than from low
density phases"(5). In these studies, the failureto reacha regulardistribution
was ascribedto 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, althoughin no case did
the different size classes of a single populationdemonstrateall three types of
distribution. In Opuntia bigelovii (79), Tidestromiaoblongifolia (54), and
Eriogonum inflatum(155), living and dead individualswere more clumped
than living individualsalone, which suggests adultdensity-dependentmortal-
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-

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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
competitionupon a species should be looked for in all stages of the life cycle;
competitive effects should not be assumed a priorito 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 hypothesisis relativelylow, althougha large body of work has
demonstratedthat plants of arid and semiaridregions are often under water
stress (24). Wateringgenerally increases rates of growth and survival, con-
firming thatit 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 watercontentshiftedthe outcome
of competitionbetween Salsola kali andperennialgrasses in favor of the latter
(1) and led to an increase in the abundancesof warmseason 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,andregularlydistributedpopulationsof 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) presentedevidence that the water supply per unit of
"transpiringsurface" is relatively constant.

Competitionfor Minerals
Fertilizationwith nitrogenincreasedthe size of winter annualsin the Mojave
desert, but phosphorusdid not (152). Nitrogen addition also increased the
biomass of most species in a dry grassland,where its principaleffect was to
magnify the results of differentwateringtreatments(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 shrubArtemisia tridentata took up much more phosphorus
from the root space it shared with Agropyron spicatum than from the root
space it sharedwith Agropyrondesertorum,and thatA. desertorumtook up
more phosphorusthandid A. spicatumwhen competingwithA. tridentata.As
the authorswere carefulto point out, these results do not imply that phospho-
rus is the only, or even the most important,resource for these plants.

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 IN ARIDREGIONS
PLANTCOMPETITION 99

Allelopathy
Aqueous extractsof Partheniumincanum(13, 14), Enceliafarinosa (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. Extractsof L. tridentatawere
found to have no deleteriouseffects on its own germinationor early growth
(8, 70). The relevance of the toxicity of aqueousextractsto 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 concludedthat
the productionof toxic substancesis relevantonly in greenhousesandperhaps
crowded nurseries. Muller & Muller (86) found no correlationbetween the
degree of toxicity of the aqueousextractfrom a shrubspecies and patternsof
herb species' occurrence under those shrub species; they concluded that
"toxins are ineffectual as factors in competitionbetween plants" in deserts.
Herbivory
The harboringof predatorsor pathogensis 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 strongerthanintraspecificcompetitionin an experimentalstudyof
Larrea tridentata and Ambrosia dumosa, although other factors were more
importantin determining these species' abundancesand distributions(38,
39). The limited evidence from studies of patternon this point, however,
supportsthe opposite generalization:Interspecificcompetitionis weakerthan
intraspecific (157, 158). The degree of reciprocity of competitive rela-
tionships would also be of interest, if relevant data existed.
The presence and relative intensityor absence, of interspecificcompetition
among different component species of a community is also of interest,
because these cast considerablelight uponthe 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 distancesbetween plants and their sizes, between species pairs.
In the Mojave desert (157) competitionoccurredamong all threepairs of two
Opuntia species and a Yucca, apparentlyat equal intensity. However, for
individualsof a given size, pairsof plantsof differentspecies of Opuntiawere

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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 additionto 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 survivalof
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 phenologicalpatternsand 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 intensityof competitionamong them;it may therebypromotetheir
coexistence. Niche separation among plants primarily takes the form of
separationof resource use in space and/orin time. The plant species of arid
and semiaridregions representa very wide range of adaptationsthat tend to
separate their use of water (113, 114). Many of these adaptationshave
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 Sonorandesert, there may be two separatesets of
ephemerals (119, 144). Furthermore,the relative abundancesof 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).

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 IN ARIDREGIONS
PLANTCOMPETITION 101

Some perennials, including most perennialgrasses (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 theiruse of small rainfalls(106, 107). Otherperennialsmaintain
leaves, photosynthesize, and absorband transpirewater even duringvery dry
periods. Both groupsof perennialsmay furtherseparatetheirperiodsof 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 aridand 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 amountsof sand and clay in the soil (119), are of particular
importancein 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 othersucculents
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 plantstend to root more deeply thangrasses, and the
resulting separationof wateruse can be sufficientto permitcoexistence (121,
137, 138).
The extent to which the soil is fully occupied with roots is usually con-
sideredto 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 individualsextend much furtherthan their cano-
pies; the apparentseparationof plants is very misleading.

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102 FOWLER

FACILITATION
The apparentfacilitationof the establishmentor growthof other plant species
by woody perennial shrubs in arid regions has been observed by numberof
authors (e.g. 112, 113). Osborn et al (95) noted that annuals were most
abundantin the mounds of sand around the bases of Atriplex vesicaria in
Australia. Went (143) described a complex set of associations between an-
nuals and shrubsin 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 annualswith woody shrubshas been ascribedto higher
soil organic content, shading (which could cause lower rates of evaporation
and transpiration),the trappingof windblown seeds, bird dispersalof 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 moundof organicmatterunderneathby
trapping wind-blown material to add to their own dead shoots. The shrub-
dependentannualswere 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 undershrubswith 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 transplantexperimentsof
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 associatedwith an increasein 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 relationshipcould produce oscillations in population

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 PLANTCOMPETITION
IN ARIDREGIONS 103

size (131), but the long life of the cactus and the importanceof occasional,
lethal freezes in determiningpopulationsizes of this species (125, 126) make
such oscillations unlikely. Based upon observationsof 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 providingprotectionfor 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 Gutierreziamicrocephalaare 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 vegetationfollowing 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 alreadypresentin 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 competitionfrom 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 mineralsby 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 grazingand 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,

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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 strongenough to be readilydetected, 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 communitiescan now be taken as given.
Studies should now be directedtowardsdeterminingthe circumstancesunder
which competition occurs and its effects upon plant populations.
The existing studies are too few to warrantgeneralizationsconcerningmost
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 questionsis, of course, uniqueto plantcommunitiesof aridand

semiaridregions, but the particulardearthof relevantstudies in these regions
warrantsdirecting attentionto them.
Anothercategory of unanswered,and at this point unanswerable,questions
addresses the similarities and differences among different regions. What
generalizationscan be made aboutthe natureand effects of plant competition
among plant communities of different temperatearid or semiaridregions of
similar climate? Between plant communitiesof temperateand tropicalaridor
semiarid regions? Between arid and humid regions? With regardto the last
question, the relative commonness and importanceof the facilitation of one
plant by another seems to me to be particularlyinteresting.

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 IN ARIDREGIONS
PLANTCOMPETITION 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.

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