SECONDARY SUCCESSION AND THE FATE OF NATIVE SPECIES IN A CALIFORNIA COASTAL
PRAIRIE COMMUNITY
Author(s): Theodore C. Foin and Mary M. Hektner
Source: Madroño, Vol. 33, No. 3 (JULY 1986), pp. 189-206
Published by: California Botanical Society
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SECONDARY SUCCESSION AND THE FATE OF NATIVE
SPECIES IN A CALIFORNIA COASTAL
PRAIRIE COMMUNITY
Theodore C. Foin
Division of EnvironmentalStudies, Universityof California,
Davis 95616
Mary M. Hektner
U.S. National Park, Service, Redwood National Park,
Orick, CA 95555
Abstract
inthenorthern
coastal
California
informer
succession
sheeppastures
Secondary
ofsecondary
Studies
favors
coverdominance
grasses.
especially
prairie
byperennials,
coveris dominated
showthattherelative
SonomaCounty,
succession
atSeaRanch,
Native
anintroduced
odoratum,
species
perennial
grass.
increasingly
byAnthoxanthum
toregain
succession
andareunlikely
their
coverduring
atincreasing
arenotsuccessful
inthecoastalprairie.
coverdominance
Historically,largepartsof the Californiacoastal prairiewere used
forgrazingby domestic animals. One of the main effectsof grazing
in this grasslandis a change in species composition,accomplished
by selection against grazing-intolerantspecies. Grazing has been
implicatedas a major factorin thechangeofvegetationin theCentral
Valley (Burcham 1957), and is likelyto have been importantin the
coastal zone as well, although such changes have never been documented.
Secondarysuccession may be definedas the predictablesequence
of species replacementsoccurringaftera disturbancesuch as grazing
defoliation.It differsfromprimarysuccession only in thatthe latter
is assumed to start from a substratethat has not been occupied
previouslyby any vegetation.Secondary succession is the key ecologicalprocessgoverningchangein species compositiononce grazing
pressureis reducedor eliminated,and thussuccessional studiesmay
reveal whetheror not the natural recoveryprocess can ameliorate
the effectsof grazing(see Mcintosh 1980).
The classic expectationof the effectsof secondarysuccession are
based on earlyworkby Clements(1916), who arguedthatsuccession
would favor native species, which are well adapted to each other
and theirphysicalenvironment.Invaders and weedy species, however,would be crowded out eventuallyand recoveryto the original
climax vegetationwould be complete. More recently,Heady and
others(1977) have predictedthat "introduced plants will continue
Madroño,Vol.33,No. 3,pp. 189-206,1986
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190
MADROÑO
[Vol.33
to be abundant on many hectaresof coastal prairie,but succession
willapparentlymove rapidlytowarddominance ofperennialgrasses
if land managementpracticesare suitable." This is a modifiedClementsianview, which recognizesthat invading species may not be
so easily displaced. Heady refersspecificallyto the replacementof
introducedannuals by native perennialgrassesin thecoastal prairie.
AlthoughintroducedMediterraneanannuals are commonlytreated
as the dominant vegetationof interiorvalleys, they have been as
abundant in the coastal prairieflora,at least in range systems.
Studiesin secondarysuccession would appear to be usefulin Californiaprairiesfor several basic and applied reasons. There are a
number of succession models proposed (e.g., Connell and Slatyer
1977, Shugart1984) thatcould be testedusingCaliforniavegetation.
On the other hand, range and reserve managers could use such
informationin managementand restorationactivities.Despite the
obvious need, secondarysuccession in the coastal prairieis not well
known at present.Those few studies that exist suggestthat native
species may dominate the vegetationas succession proceeds. Huffakerand Kennett(1959) showed that perennialgrasses,especially
one native species (Danthonia californica)replaced an introduced
perennial( Hypericumperforatum)knownas Klamath weed, following successfulbiological controlby the Klamath weed beetle ( Chrysolina hypericï)in the coastal prairieof Humboldt County.
Elliott and Wehausen (1974) analyzed cover in threeplots with
different
grazingpressurein Point Reyes National Seashore. Their
data predictthatthe coastal shrubBaccharispilularisand the native
perennialbunchgrassDeschampsia holciformiswould increase during succession. Lathrop and Gogan (1985) surveyedthe Tule Elk
Range at Tómales Point,immediatelyto thenorthofthearea studied
by Elliottand Wehausen. They arguedthatshrubswould dominate
secondary succession in wind-protectedareas, but that perennial
grasseswould also increase (particularlythe native perennialStipa
pulchraHitchcock).The invasion ofprairiebynortherncoastal scrub
is indicated by their data, and is more stronglysuggestedby the
work of McBride and Heady (1968) and McBride (1974). It is not
clear,however,thatthe coastal prairieis a serai stagewhose climax
is ultimatelyshrubor forestdominated.
Despite the evidence suggestingthe successional superiorityof
native species over introducedspecies, it is farfromconclusive. A
numberofcounterexamples,demonstrating
thetenacityofinvading
species, also exists; perhaps the best example is that of Bromus
tectorumin theGreat Basin (Harris 1967, Mack 1981). The literature
pertainingto the emergenceof natives during succession in Californiaprairiesalso does not supportthetheory.White(1967) claimed
thatStipa pulchrashould be one of the climax dominantsin the oak
woodland at Hastings Reservationin Carmel Valley, and Burcham
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1986]
FOIN& HEKTNER:SUCCESSIONIN COASTALPRAIRIE
191
(1957) hypothesizedthatthe same species dominated the primeval
CentralValley prairies.On the otherhand, Wester(1981) has used
historicalrecordsto argue that perennialgrasses did not dominate
the inland prairiesnow dominated by introducedannuals, exceptin
wet places. Bartolome and Gemmili (1981) showed experimentally
that Stipa pulchra was competitivelyinferiorto Bromus mollis at
highdensities.Their evidence makes it clear thattheremay be great
between species, and that being "native" does not necdifferences
essarilyconfera competitiveadvantage over introducedspecies.
Even the theoreticalexpectationis fragilebecause it was based on
a number of assumptions that have been proven erroneous, the
largestof whichwas the assumptionof evolutionaryadvantage in a
highlycoevolved plant community(for a good discussion of the
contributionof Clements to modern plant ecology see McMahon
1980). These argumentsmay be used to challenge the importance
of the communityin succession and to emphasize the importance
of the population dynamics of "key" species in governingthe outcome of succession (Foin and Jain 1977, Westoby 1979, Mcintosh
1981).
To evaluate the dominance of native versus introducedplants in
secondary succession of coastal prairie, we conducted chronosequential studies at Sea Ranch, Sonoma Co., California (38°40'N,
123°24'W) in 1974. An earlierpaper (Hektnerand Foin 1977) exareas of the Sea
amined differencesin the vegetationin different
Ranch coastal prairieusingthedata obtained in thesummerof 1974.
The presentpaper reportstheresultsof successionbased on the 1974
census and additional samples taken in 1975-1978. Detailed information on siteconditions,species present,and land use historymay
be foundin the 1977 paper.
Study Areas
Sea Ranch is a recreationalsubdivision 180 km north of San
Francisco. It occupies 16 km of the coastal terracefromStewarts
Point northwardto the Mendocino Co. line. Homesites have been
developed such that large areas of meadow and foresthave been
preservedas permanentopen space. Sea Ranch includestwo uplifted
marineterraceswithsandyloams (Baywood and Rohnervilleseries:
U.S. Dept. Agr. 1972) overlyingsandstone and basalt. All study
areas reportedin thispaper are located in the commons areas of the
firstterrace(that terraceadjacent to the coast) and were protected
fromgrazing,fire,mowing, and other overt management for the
durationofthestudy.Disturbanceofthevegetationwas notexcluded
completelybecause soil disturbanceby gophersand moles, grazing
by insects,rabbits,deer, and mice, and limitedtramplingby hikers
wereall present.Three studyareas weresampled,thenames ofwhich
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192
MADROÑO
[Vol.33
inJune1984.Thephotograph
ofLoneTreeMeadowtaken
Fig.la. Photograph
mark
stakes
Thewhite
tothenorthwest.
from
thetopoftheslopelooking
wastaken
inthe
Thelight-colored
stations.
ofsoilmoisture
thelocation
vegetation
monitoring
it is thewetbehind
is an annual
patch.Thevegetation
-Rytidosperma
foreground
totheleftsideis mostly
). Thevegetation
type(Deschampsia-Holcus-Anthoxanthum
odoratum.
Anthoxanthum
8 Jul1975.
ofBuckMeadowlooking
Fig.lb. Aerialphotograph
east-northeast,
The
to theright.
ofMonterey
Thereis a prominent
cypress)
(composed
hedgerow
andseveral
tracks
Sims.Onesetofvehicle
arboreus
dotsarescattered
Lupinus
light
inthephotograph.
areevident
the1974vegetation
trails
madeduring
survey
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1986]
FOIN& HEKTNER:SUCCESSIONIN COASTALPRAIRIE
193
are traditionalones used by the previous owner of the ranch (the
late E. J. Ohlson).
Lone Tree Meadow. This studyarea is a 160 x 80 m rectangular
slope, approximately300 m from
gridlocated on a northwest-facing
the ocean to its westernboundary(Fig. la). The site is unprotected
fromthe prevailingnorthwesterly
winds. It was the one area of the
formersheep ranchthatwas used most intensively,particularlyfor
year-roundherds of ewes and their lambs. Estimates of stocking
rates are not available.
Buck Meadow. This studyarea is a 200 x 300 m rectanglewith
the seaward edge 100 m fromthe coast bluff(Fig. lb). This site is
flat,withno pronouncedslope. It was named forits grazinghistory:
it was used as the ram (buck) pasture. Ohlson noted that ram densitieswerealwayslow, particularlyin lateryearswhenstockdensities
were fallingrapidly.
Stable Meadow. This site is the only studyarea available on Sea
and used as a pasture
Ranch thatis stillgrazed.The area is fenced-off
and paddock forhorses. Use rates are low but continuous,on the
order of 2 animals per ha. Vegetative biomass is low, making the
site visually distinctive,and there appears to be at least as much
damage fromhooves breakingthe soil surfaceas fromdefoliation.
The area sampled (50 x 50 m) is approximately120 m fromthe
coastal bluff,in the centerof the pasture,with a gentlewest-facing
slope towardsthe ocean. Otherthan its presentuse, Stable Meadow
is verysimilarin slope and aspect to Buck Meadow.
The Sea Ranch lands were used forgrazingpurposesfromthe late
19th centuryto 1968. Small parts were used experimentallyfor
but those experimentswereunsuccessful.In 1965, when
agriculture,
the ranch was sold fordevelopment,sheep were removed progressively fromsouth to north,until the last sheep were sold in 1968.
Lone Tree and Buck meadows remained under grazinguntil 1968.
Methods
Sampling procedure.The samplingmethodologyfollowedin this
studyused line transectsestablished forthe initial surveyin 1974.
Complete details ofthe methodologyare foundin Hektnerand Foin
(1977). In brief,line transectswere establishedrandomly,one transect each 10 m, and sample locations were taken randomly,one for
each 10 m of transectlength.Cover estimationswere made using
theDomin index,as modifiedbyMajor fromEvans and Dahl (1 955),
using 0.25 m2 circularquadrats.
Stable Meadow was sampled using30 quadratsin 1978 to establish
a frameof referenceforthe othertwo sites. Lone Tree Meadow was
sampled threetimes (1974, 1976, 1978) at 143 quadrat locations.
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194
MADROÑO
[Vol.33
Buck Meadow was sampled twice (1975, 1978) at 242 quadrat locations.
Two possibleproblemsexistin themethodologyused. First,placementofthequadrat samplingringwas notcenteredon thepermanent
markingstake (which was on the transectline), but was offsetto
avoid sampling pathways. This procedure did not permit precise
relocationof the quadrat in subsequent censuses. Second, all sites
were sampled late in the summer(August and September).This is
an artifactof the firstcensus (1974) because all later censuses were
made at the same time to maximize year-to-yearcomparability.
Samplingearlierin the summer,when the grass species were clearly
identifiable,but annuals were more evidentin the field,would have
been preferable.Althoughannuals, especially forbs,were undoubtedly underrepresentedin our results,it is the comparative aspect
and trendin time that we required. These two aspects of the data
are least sensitiveto absolute bias in cover estimates.
Treatmentofcoverdata. The Domin value of each species in each
quadrat was converted firstfrom its index value, each of which
covers a range of percentcover, to the midpoint of the range for
that value. This follows the procedure used by Hektnerand Foin
(1977). The convertedDomin values were then calculated to yield
fourindices: absolutecover,the sum of convertedDomin values for
each species over all quadrats; relativecover, the percentrepresentationof a given species over all quadrats as a fractionof the cover
attributableto all species; mean cover, the mean absolute cover per
quadrat; and adjusted mean cover, the mean absolute cover calculated only forthose quadrats in which the species occurred.
Errorbars weretakenas ± 2 s.e. (standarderrors)on mean cover.
Nonoverlappingerrorbars were used as tests of significantdifferences between means with a = 0.05. This procedureis more conservative,i.e., it has smallerType I error,than standardparametric
confidencelimitsunless df < 5 (see Steele and Torrie 1960). None
of the comparisonswas based on so fewdegreesof freedom.
Assignmentsof lifeformweremade usingReed et al. (1963). Five
classes were used: annual forbs,perennialforbs,annual grasses,perennialgrasses,and "otherlifeforms".The last categoryis composed
primarilyofthreespecies:Pteridiumaquilinumvar.pubescens, Rubus
ursinus,and Linum bienne.All nomenclaturefollowsMunz (1973),
except forDeschampsia holciformis(Crampton 1974) and Rytidospermumpilosum ( =Danthonia pilosa).
Results
Successional Changes in Life Form Composition
Species data fromthe threecensuses of Lone Tree Meadow are
aggregatedinto life formgroups and plotted in Fig. 2. This plot
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1986]
FOIN& HEKTNER:SUCCESSIONIN COASTALPRAIRIE
195
inStableMeadow
Fig.2. Meancovertrends
(MC2)forlifeforms
(leftward
points)
Error
barsare±2 standard
errors.
andLoneTreeMeadow,1974-1978.
shows mean cover for each of the major life formsexcept "other
life forms". The threecensuses in Lone Tree Meadow are plotted
with the data from Stable Meadow displayed at the extremeleft.
The Stable Meadow data are included to provide an indication of
the structureof the presuccessional coastal prairie vegetation.Because these data are only an indication of life form mean cover
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196
MADROÑO
[Vol.33
values for Lone Tree Meadow, these points are connected to the
1974 data with dashed lines. In any case, the extrapolationsmust
be viewed cautiously,because environmentaldifferencesbetween
sitesand differences
betweensheep and horse grazingcould be very
important(Harper 1977). Note thatannual forbscomprise so little
of thetotalcover thatthe ordinatein Fig. 2 mustbe brokenin order
to include them.Two s.e. confidenceintervalsdo not overlap in the
followingcomparisons: 1) the mean cover of annual forbsdecreased
from 1974-1978; 2) annual grasses increased from 1974 to 1976
and decreased from 1976-1978; 3) perennialforbsincreased from
1974 to 1976, and decreased from1976-1978 and from1974-1978;
and 4) perennialgrassesincreasedin cover at each samplingperiod.
Lone Tree Meadow had less cover forannual grassesand perennial
forbsthan Stable Meadow forall threesampling periods, and had
greatercover in perennialgrasses for 1976 and 1978.
Figure3 shows the cover data forBuck Meadow arrangedin the
same fashion as the data in Fig. 2. Using the same comparisons,
annual grassesshowed a decrease in cover,whereasperennialgrasses
showedan increaseover theperiod 1975 to 1978. Neitherforbgroup
showed a significant
change over this period. Comparison to Stable
Meadow data shows (1) no significantdifferencesforannual forbs;
(2) significantly
highercover forannual grassesin Buck Meadow in
lower cover in perennialforbsforboth years;
1978; (3) significantly
and (4) a significantdifference
forperennialgrasses by 1978.
Comparison of Figs. 2 and 3 illustratesthat trendsin life forms
in each site were similar.Annual forbswere not common in either
site and did not have significantly
highercover in Stable Meadow.
This situation,however,is likelyto be an artifactof the timingof
sampling,because adjacent sheep meadows can supporta varietyof
annual forbs,especially when heavily grazed. Both annual grasses
and perennialforbsdecreased duringsuccession, which is particularlyevident in the Lone Tree Meadow data. Perennial grasses increased significantly
to similarcover values in both sites. Lone Tree
and Buck Meadows showed decreasingcover in perennialforbsand,
withtime, more perennialgrassesthan Stable Meadow.
This trendcan be seen more easily in a plot of relativecover (Fig.
4). These data are separateestimatesforLone Tree and Buck Meadows, exceptfor 1978, which is the arithmeticaverage forboth sites.
The figureshows a decrease in relativecover forall categoriesand
forbare ground,except forperennialgrassesand "otherlifeforms".
Bare ground never exceeded 5% relative cover at Sea Ranch, and
decreasedto less than 1% of the relativecover in the 1978 censuses.
Successional Changes in Species Composition
Principalserai dominantsduringsuccession.The majorityof the
species encounteredin the quadrats were infrequentand low in
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1986]
FOIN& HEKTNER:SUCCESSIONIN COASTALPRAIRIE
197
Fig.3. Meancovertrends
forStableMeadow
toBuckMeadow,
1975compared
1978.Arrangement
ofthisfigure
tothatofFig.2.
corresponds
cover. We used two criteriato singleout those species thathad high
cover values at one census or another: 1) any species that occurred
in fiveor more quadrats at any one of the threesites in one or more
census periods,and 2) thatalso had 1% relativecover in the site for
which criterion(1) was met. Application of these criteriaexcludes
all annual forbsand reduces the list to 19 species (Table 1).
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198
MADROÑO
[Vol.33
Fig.4. Relative
coverassigned
toalllifeforms
andtobareground
forLoneTree
andBuckMeadows;1978is theaverage
forbothsites.Relative
coverforStable
Meadowis givenattheleftendoftheabscissa,
as doneinFigs.2 and3.
The species in Table 1 are arrangedby lifeform.Absolute cover
(AC) and frequencyin sample quadrats are tabulated for all 19
species meetingthe criteriaoutlined above for each surveyof the
threesitessampled. There is sufficient
consistencyamong theresults
shown in Table 1 to suggestsuccessional patternsin the species as
well as the lifeforms.The species thathad highfrequencyalso had
high absolute cover. The dominants of early and later stages in
succession are readily identifiedfromthis table, assuming Stable
Meadow representsthe startingpoint. Cynosurusechinatus is an
example ofan annual thatshowedhighcover and frequencyin Stable
Meadow and in earlysurveysof the othertwo sites. Otherexamples
ofcharacteristicspecies ofearliersuccessionalstages,whichfollowed
the terminationof grazing, include Plantago lanceolata, Linum
bienne, Lolium perenne,and Rytidospermapilosum.
Changes in species composition may be identifiedby considering
significant
changesin mean cover (Table 2). All but one of the early
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1986]
FOIN& HEKTNER:SUCCESSIONIN COASTALPRAIRIE
199
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This content downloaded from 169.237.27.245 on Sun, 5 Jan 2014 15:23:56 PM
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1986]
FOIN& HEKTNER:SUCCESSIONIN COASTALPRAIRIE
201
dominants (the exception being Rytidospermapilosum) decreased
(p < 0.05) in Buck and Lone Tree Meadows by 1978.
significantly
The dominantspecies in the latestsurveyswere mostlyperennial
grasses.Anthoxanthumodoratum and Holcus lanatus were two introducedspecies absentfromStable Meadow thatshowed significant
increasesin mean cover in the othersites.Anthoxanthumodoratum
had highercover values, greaterdispersion over the successional
sites,and higherrates of increase than most otherspecies. In Lone
Tree Meadow, Anthoxanthumodoratumreached 69% oftherelative
cover (Fig. 5a) of the perennial grasses in 1978, occupied 91% of
the sample quadrats, and had increased 266% in actual cover. In
Buck Meadow, the correspondingfigureswere 54% (Fig. 5b), 80%,
and 59%. Holcus lanatus increased significantly
in both meadows
and showed higherpercentrates of increase than did A. odoratum.
Its frequencyvalues werelower,reflecting
highpatchinessand locally
high cover. Rytidospermapilosum was the only perennialgrass to
show differences
between sites. In Lone Tree Meadow, R. pilosum
disappeared fromhalf the quadrats in which it occurred and lost
47% ofitscover. In Buck Meadow, R. pilosumincreasedsignificantly
(71%), despite a reduction in frequency,because its mean cover
increased in those plots occupied at the beginningof the study in
1975. Deschampsia holciformis
, a large native bunchgrasscharacteristicof wettersites,did not show significantchange in time and
may representa relict species that survives grazingratherthan a
later successional dominant. Other species that showed significant
increasesin cover include Vulpia bromoides( =Festuca dertonensis)
and Rubus ursinus.Rubus ursinusis characteristicof heavy stands
of perennialgrass,especially in wetterareas. Vulpia bromoideswas
unique in that its frequencydecreased, whereas cover increased in
the remainingquadrats. This patternof locally patchy occurrence
in perennialstands persiststo the presenttime.
Representationof nativespecies in secondarysuccession. Table 3
shows thatnative species are presentat all stagesof succession,but
at low cover values. There is no significanttemporalincrease in the
ofnativespecies,withtheexceptionofRubus ursinus.
representation
We see little evidence suggestingthat a major increase in native
species, at least in the herbaceous vegetation,is to be expected in
the future.Deschampsia holciformis
, however,is one native species
thatcould be a successional dominant,but it has a staticpopulation
at Sea Ranch with no evidence suggestingdominance at climax,
contraryto what Elliott and Wehausen (1974) suggestfor Point
Reyes.
Two other well-knownnatives are relativelyrare and likely to
remain so. Stipa pulchra is uncommon at Sea Ranch. It tends to be
concentratedalong roadsidesand in relictannual-dominatedpatches.
Our observationson this species at Sea Ranch supportthe conclu-
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202
MADROÑO
[Vol.33
inLoneTreeMeadow(Fig.5a) and
coverforperennial
Fig.5. Relative
grasses
BuckMeadow(Fig.5b).
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1986]
FOIN& HEKTNER:SUCCESSIONIN COASTALPRAIRIE
203
inthe
Table 3. Representation
of NativeSpeciesinSecondarySuccession
areabsolute
cover(AC)assigned
tothe19
Sea RanchCoastal Prairie.Statistics
ofTable1.
andintroduced
native
species
AC
nativespecies
Siteandyear
StableMeadow
119
1978
LoneTreeMeadow
2714
1974
1976
3243
3882
1978
BuckMeadow
1150
1975
1294
1978
AC
allspecies
of
Proportion
natives
3663
0.032
10,761
15,527
16,014
0.252
0.209
0.242
12,338
27,273
0.093
0.047
sions drawn by Bartolome and Gemmili (1981). Danthonia californica is a low-growing,native, perennialthat is restrictedto the low
vegetationfoundon exposed bluffs(see Hektnerand Foin 1977). It
has never been found inland in the tallervegetation.
Discussion
Secondary succession in the California coastal prairie. Our data
show that secondarysuccession in the Sea Ranch coastal prairieis
characterizedby rapid replacementof annuals by perennials and
subsequentlytheconcentrationofdominance withina small number
of mostlyintroducedspecies of perennialgrasses. The replacement
of annuals by perennials is most easily explained as a decline in
yearlyestablishmentby annuals because perennialsutilize most of
the soil surface.Peart and Foin (1985) have shown that establishment is stronglyretardedby biomass at Sea Ranch. As succession
has progressed,biomass has accumulated (much of it standingdead
material and litter).The cause of the replacementof annuals and
perennialforbsby perennialgrassesis more speculativeat thistime,
but may be a functionof competitionfor lightand soil moisture.
Peart (1982) measured lightlevels beneath the grass canopy using a
active spectrum.
lightmetermeasuringonly the photosynthetically
He foundthat lightvalues beneath the grass canopy were typically
much less thanat the floorof nearbyredwood forests.Because most
of the forbsat Sea Ranch are low-growing,it is easy to imagine that
theywould be readilyovertoppedand shaded out.
The replacementof some perennialgrassesby othersat Sea Ranch
is exemplifiedmoststrikingly
bytheincreasein Anthoxanthumodoratum. This species has continuedto increase;at present,it has even
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204
MADROÑO
[Vol.33
largercover values than those presentedin this paper (the informal
estimatefor 1985 in Lone Tree Meadow is greaterthan 70%). Explanation of the success of this species is stillincomplete,although
thereare suggestionsin some of our currentworkthatsoil moisture
is an important,perhapsthemost important,explanation.Peartand
Foin (1985) showedin experimentalinvasion studiesthatAnthoxanthum odoratum was the best at colonizing the stands of annualperennialforb-Rytidospermathatwere presentwiththe sheep. Differencesin colonizingabilityare adequate to explain the changesin
secondarysuccession reportedin this paper, but newer data (Foin,
unpublished) strengthenthe argumentby showing the role of soil
moisturein the dynamicsof the dominant perennialspecies.
Succession and the nativevegetation.Our researchat Sea Ranch
demonstratesthat native species cannot be expected to recover in
the coastal prairiein the face of competitionfroma small number
of well-adapted introducedspecies. Even with intensive management, species like Anthoxanthumodoratum and Holcus lanatus
probablycannot be eliminatedfromthe prairiecommunity.This is
because the coastal climate is mild enough in the summerto permit
the survivalof perennials,and because thesetwo introducedspecies
are opportunisticand adapted to a wide varietyof environmental
conditions. The researchreportedhere suggeststhat, under mesic
conditionson the northerncoast, these two introducedspecies are
the prairiedominantsand likelyto remain so. Barryand Schlinger
(1977) have shown that introducedperennial grasses dominate at
Inglenookfen,and recentlySaenz and Sawyer(1986) have drawn a
similarconclusion about introducedspecies in Humboldt County.
The Sea Ranch data also are in harmonywith the replacementof
natives by introducedannuals in the interiorvalleys. Thus, Heady
and others(1977) were correctin theirpredictionthat introduced
species would continueto persistin the coastal prairie,but theydid
not foreseethat persistencealso would mean even greaterdominance.
Acknowledgments
forthiswork
wasprovided
Science
Foundation
Support
bytheNational
(DEB 76andtheOpportunity
forEcological
Foundation,
82503),theRockefeller
Program
ofCalifornia.
Wearegrateful
tothemembers
andstaff
ofThe
Research,
University
SeaRanchAssociation,
thelateLarry
RobWickstead,
Marshall,
particularly
George
ertKirkwood,
andRuthRiordan,
fortheir
assistance
overtheyears.We
generous
alsothank
HaroldHeadyandCarlaScheidlinger
fortheir
extensive
comments
on
themanuscript.
LiteratureCited
Fen.Dept.Parks
andRecreation,
Barry,W.J.andE.I. Schlinger.1977. Inglenook
StateofCalifornia.
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1986]
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Ecology
20 Apr1986.)
6 Sep 1985;revision
accepted
(Received
ANNOUNCEMENT
Symposiumto Honor G. Ledyard Stebbins
AnInternational
willbe heldin Davis,California,
on
Symposium
inthe
12-14September
Professor
G. Ledyard
Stebbins
1986,tohonor
will
Invited
talksbyleading
plantbiologists
yearofhis80thbirthday.
andecological
andnuinclude
topicsinpopulation
genetics,
organelle
andplantdevelopment,
and
clearmolecular
morphogenesis
genetics,
information
Dr.
evolution
andsystematics.
Forfurther
pleasecontact:
orDr.S. K. Jain,
L. D. Gottlieb,
ofGenetics,
Department
Department
ofCalifornia,
ofAgronomy
andRangeScience,
Davis,CA
University
95616.
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