Ecotone Hierarchies

Author(s): James R. Gosz

Source: Ecological Applications, Vol. 3, No. 3 (Aug., 1993), pp. 369-376
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1941905
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Ecological Applications,3(3), 1993, pp. 369-376

? 1993 by the Ecological Society of America

ECOTONE

HIERARCHIES1

JAMES R. GoSZ2
BiologyDepartment,
University
ofNewMexico,Albuquerque,
NewMexico87131 USA

Abstract. Ecological phenomena are evidentover a broad spectrumof space and time
scales. Ecotones, being definedas zones of transitionbetweenadjacent ecological systems,
also must occur over an equally broad spectrumof space and time scales. Scale-dependent
constraintsinfluenceecological phenomena and resultingzones of transition;however,in
traditionalecotone studies littletreatmenthas been given to these influences.This paper
addressesaspects of the ecotone conceptthatrelateto the strengthof interactionsbetween
adjacent systemsfor a hierarchyof ecotones in a biome transitionarea in centralNew
Mexico on the Sevilleta National Wildlife Refuge. Zones of transitionoccur at plant,
population,patch,landscape, and biome levels in thehierarchysuggestedhere.Constraints
differacross this hierarchy,primarilybecause of the different
scales at which these constraintsexerttheirinfluences.The basic strategyto understandthesecross-scaleinfluences
mustbe to performstudiesat different
scales, and a hierarchicalapproach identifiesthose
scales. This also is importantforidentifying
the appropriatetechnologiesthatfocusat the
scales where transitionzones between ecological systems/phenomenaare expressed. A
broad arrayof technologiesare available forintegrating
the pattern-processrelationships
that occur across the many scales in ecological systems.
Key words: biome;dynamics;ecotone;hierarchy;
remoteimaging;Sevilleta,New
interactions;
transition.
Mexico;threshold;
This paper will address aspects of the ecotone concept
thatrelateto the strengthof the interactionsbeEcotone: a zone of transitionbetweenadjacent ecotween
adjacent systems,fora hierarchyof ecotones in
logical systems,havinga set ofcharacteristics
uniquely
definedby space and time scales and by the strength a biome transitionarea. The example in this paper
space and time
oftheinteractionsbetweenadjacent ecological systems identifiesstudiesofecotonesat different
scales
that
are
used
to
translate
pattern-process
rela(Holland 1988). This definitionproposedby a working
tionships
for
this
biome
transition
zone.
groupin Paris in 1987 was recognizedas beinggeneral;
however, it also was agreed that it was a necessary
THE ECOTONE HIERARCHY
launchingpointfordevelopmentofthetheoreticalbase
Table 1 depictsa hierarchyofecotonesrangingfrom
necessaryforfuturediscussionsoftheecotoneconcept.
the
biome ecotone (i.e., biome transitionarea) to the
Ecotonal phenomenaare evidentover a broad specplant
ecotone. The traditionaltypes of ecotones intrum of space and time scales, and studies need to
adopt a multi-scaleperspectiveperformingstudies at cluded are those of the patch (Forman and Godron
several scales (Gosz 1991). An importantfocus must 1986) and biome (Gosz and Sharpe 1989, Gosz 1991,
be on linkagesbetweendomains of scale, as called for Neilson 1991). The hierarchyin Table 1 adds addiby Meentemeyerand Box (1987): a "science of scale." tional levels, down to the transitionthat occurs at a
Oftena hierarchyof scales is proposed; however,such
arrangementsdo not mean thatwe understandhow to
translatethe pattern-processrelationshipsassociated
withecotones across the nonlinearspaces betweendomains of scale (Gosz 1991). We are only likelyto recognize such linkages when we identifyhow different
hierarchicallevels constrainone another(Weins 1989)
as wellas thedifferent
typesofcontraintsthatdominate
at particularscales. These scale-dependentconstraints
are relatedto different
ofinteractionsbetween
strengths
adjacent systems.
INTRODUCTION

' Manuscriptreceived 5 March 1992; revised 1 November

1992; accepted 3 November 1992.
2 Presentaddress: Division Director,Division of EnvironmentalBiology,National Science Foundation, 1800 G Street,
N.W., Washington,D.C. 20550 USA.

FIG. 1. Hypothetical
biometransition
zonereflecting
the
mosaicpatternacrossthezone separating
the two biomes
(biomeecotone).The patternchangesfromrelatively
large
patchesinthecoreareaofeachbiometo smallpatchesat the
ecotone.Environmental
changesthatresultin biomemovementshouldcausea displacement
ofthemosaicpattern.

EcologicalApplications

JAMES R. GOSZ

370

A.

Vol. 3, No. 3

GreatPlainsSteppe
B. eripoda

Deep Well

30B.

L. tridentata

Larrea tridentata
>

U,

20

5-Points

5-Points
0 5-Points

10-

,L

010

B. gracilis

20

30

40

50

60

70

80

90

100 110

60

70

80

90

100 110

301

>20

La Joya

0)
LL

Joy

10

10

ChihuahuanDesert

La Joya

20

30

40

50

Patch Widthat Midpoint(m)

of theextentof
of threespeciesin theecotone.(A) Schematicrepresentation
FIG. 2. Ecotonediagramand occurrence
area,and thelocation
in theSevilleta,New Mexico,biometrainsition
rangeofthethreestudyspeciesalonga 10-kmtransect
forthethreedominantspeciesin this
and degreeofmovement
ofthethreestudysites.The arrowsindicaterelativedirection
Boutelouaeripoda,andB. gracilis.Notethe
biomeecotone.(B), (C), and (D) Patchsizeand abundanceofLarreatridentata,
(Deep Wellstudy
end ofthetransect
at thenorthern
graphs.Larreawas notencountered
scalechangesin thesepatch-size
(La Joyastudyplot).
endofthetransect
at thesouthern
plot),and B. graciliswas notencountered
plant edge. The reason forthese additional levels is to
increasethe numberof scales of studyand the ability
to extrapolatebetween them. For example, the landscape ecotone relatesto a transitionin the mosaic patternthat is characteristicof an ecotone between two
biomes. The mosaic patternis made up of variations
in the numberand sizes of patches that representthe
two biome types(Fig. 1). There are many patch ecotones in a biome ecotone, and it is likelythat thereis
a spectrumof patch-interactionprocesses depending
on thejuxtaposition,size, substratepatterns,and distances betweenpatch types.A studyof one patch eco-

tone cannotcompletelycharacterizeor be extrapolated

to the biome ecotone.
At the other end of the hierarchyis the plant-edge
ecotone. Measurementsat the 1-iM2 scale oftencharacterize plant edges. At this scale the two ecological
microclimates,soil chemissystemsare the different
tries,and soil fauna and florathatoccur undera plant
plant cancanopy vs. in the open or under a different
opy. The level betweenthepatchand theplantecotone
is identifiedas a population ecotone. This represents
thespatialpatternofthepopulationofindividualplants
of a certainspecies and is similar conceptuallyto the

August1993

ECOTONE HIERARCHIES

C.

70-

D.
D
Boutelouaeripoda

60-

371

70

Bouteloua gracilis

50C 40C 30 !1
U 300-

>.0

C 40

Deep Well
12?

242326

46

Deep Well

(D _30

20100-

10

2 4 6 8 1Q12 14222430424696

100

42 _

194

70-

8 10 12 14 16 18 20 22 24 26 28

70-

6050-

I
C

50

40

L40O

30~

5-Points

~~~5-Points

201
0

2 4 6 8 10 12 14222430424696

2 4 6 8 10 12 14 16 18 20 22 24 26 2

142
100 194

Patch Width at Midpoint

70-

(in)

602-01

10

10
50C400

Joya
~~La

Cr 30~

LL201I

1oE
24

8 10 121422

24 3042 4696

142
100 194

Patch Widthat Midpoint(in)

FIG. 2. Continued.

spatialpattern(mosaic) of patchesin the landscape

ECOTONECONSTRAINTS
ecotone.Wherelandscapepatchesare dominatedby
one plantspecies,thepopulationand patchlevelsare
Table 1 also suggests
probableconstraints
thatcan
identical.We expecta spectrumof individualplant- be dominanton the different
ecotonehierarchies.
It
edgeinteractions
forthemosaicrepresenting
thepop- seemsclearthatat thebiomescale,climate(expressed
ulationthatdependsonthejuxtaposition,
sizesofplants, throughweather)interacting
withmacrotopography
distancebetweenneighbors,
understory
species,and constrainsthe speciesof a particularlife-form
that
plantspeciesinteractions.
No singleplant-edge
study characterizes
a biome(Neilson 1991). In contrast,
at
(i.e., -iM2 plot)cancharacterize
thepopulation
orpatch. thescaleofindividualplantedges,climateand macroThe strengths
oftheinteractions
betweenpopulations topography
are constants,
and thecontrolsare likely
are influenced
by populationsizes, spatialarrange- to be factorssuchas microclimatology,
facmicrosite
ments,distancesfromothersubpopulations,
etc.,and tors(soilmoisture,
chemistry),
plantgenetics
expressed
arewelldocumented
in theliterature.
The challenge
is through
plantchemistry,
physiology,
competition,
and
todevelopprocedures
tointerrelate
processesandcon- plant-animalinteractions.
The constraints
at interstraints
at thesedifferent
levelstoallowinterpretationsmediatescales are characterized
by a progression
of
at all levels.
overlapping
controls
thatoperateatthoseintermediate

372

JAMES R. GOSZ

Ecological Applications
Vol. 3, No. 3

fora biometransition
area.Each theinstability
Ecotonehierarchy
near suchbifurcation
points:(1) increases
has a rangeof constraints in the relaxationtime fromsmall perturbationscould
levelin theecotonehierarchy
inx symbolizes
betweentheconstraints;
andinteractions
vary be measureddirectly;and (2) increasesin the variance
constraints
Theprimary
constraints.
teractions
between
inthenumber of observed fluctuationsbeforethe discontinuity.
withan increase
withthescaleoftheecotone,
scales.
at finer
ofpossibleconstraints
The spatial patternsof ecotones provide a spatial
analog forthese temporaldynamics.A transectacross
Probableconstraints
Ecotonehierarchy
an ecotone is expectedto show increased spatial variClimate(weather)x Topography ation (e.g., increasednumbersof small patches of difBiomeecotone
x Soil
Weatherx Topography
ferentspecies) at the ecotone (see next section). TemLandscapeecotone
characteristics
(mosaicpattern)
poral studiesare expectedto showthegreatestvariation
x Biological
Soil characteristics
Patchecotone
in spatial change at the ecotone as micrositeschange
vectorsx Speciesinteractions fromone communityto another.Away fromthe ecox Microx Microtopography
tone the changes are primarilyan increase in the diclimatology
ameter of the patch ratherthan a state change. The
x
Intrainteractions
Interspecies
Populationecotone
x Physio- exceptionto thismightbe thechangeofan entirebiome
speciesinteractions
(plantpattern)
logicalcontrolsx Population
(e.g., forestto savanna) if the climate changes. This
x
geneticsx Microtopography
would be an ecotone in time (Neilson 1991).
Microclimatology
Grover and Musick (1990) describea ratchet-pulse
x Intrainteractions
Interspecies
Plantecotone
x Physio- dynamic for landscape change in a transitionalarea.
speciesinteractions
logicalcontrolsx Plantgenet- Certainmicrositesor landscape facetsare at a threshold
x Soil
ics x Microclimatology
withrespectto the conditionsallowingthe persistence
x Soil faunax Soil
chemistry
of a certaincommunity.Relativelysmall changesthat
etc.
microflora
exceed the thresholdconditions(e.g., threesuccessive
dryyears)cause certainsitesto be changedto a different
communitywhiletheothersremainunchanged.In these
constraintsoperate areas thepatternof wetvs. dry,cold vs. hot yearsmay
scales. The point is that different
acrossscales be more influentialthan the conditions of any single
levels,and studiesextrapolating
at different
constraintsas well. year.The effectiveness
need to integrateacrossthedifferent
of a patternof wet or dryyears
Anotherpoint made in Table 1 is that,in addition in causingchangemay also be relatedto thelife-history
controlsdominatingthe finerscales (e.g., strategiesof the plant species. Most plant species have
to different
plant ecotone), thereis an increase in the number of strategies(e.g., long-lived,seed banks) that allow tolprobable controls and interactionsbetween them at erance of short periods of stressreiative to the lifethese finerscales. This contributesto the increased historystrategy.Certainmicrositesor landscape facets
in mod- magnifytheinfluenceof a climaticsignalor reducethe
variationat finescales as well as the difficulty
of the life-history
strategyforsome speellingand predictingresponses.The spectrumofresults effectiveness
forinteractionsat finescales is potentiallylargerand cies, resultingin mortalityand/ornew establishment.
The temporaland spatial dynamicsof ecotone commorevariablethanthespectrumat broad scales. Much
of the fine-scalevariation is integrated(or averaged munitiesare alleged to be much more variable than
out) at broad scales, leaving only the broad-scale con- those of biome or patch core areas. This variability
straintscorrelatedwith broad-scale ecotone patterns may be seen as a curse or a benefit.For researchprogramsthatutilizevariabilityas a sensitiveindicatorof
thatchange over broad (long) temporalscales.
environmentalchange, ecotones are logical areas for
THRESHOLD BEHAVIOR
study.
Gosz (1991) suggestedthatspatial patternsin landSEVILLETA NATIONAL WILDLIFE REFUGE
scapes have nonnormal,spatiallyautocorrelated,nonCASE STUDY
stationary,discontinuous,and irregularlyspaced pain
The Sevilleta National WildlifeRefuge(NWR) is a
rameters.The dynamicsof ecotones landscapes also
are likely to be nonlinear,perhaps chaotic, and can Long-Term Ecological Research (LTER) site in New
behave in waysthatare not simpleaveragesofadjacent Mexico. This site is in a biome transitionarea where
resourcepatches(Naiman et al. 1988, Gosz and Sharpe these multi-scalestudies are being performedto de1989). Thresholddynamicsare expectedto occur near velop cross-scaleextrapolationsand explanations.Figs.
results
2-5 demonstratesome oftheintermediate-scale
boundaries.Biological systemsare metastable(O'Neill
et al. 1989), meaningthat the propertiesof the land- that are being used to relate plant to biome extraposcape remain stable only over a limitedrangeof con- lations.In a biome transitionzone, species respondin
ditions.As those conditionsreach a criticalthreshold, an individualisticmanner to the gradientsof factors
the systemmay reacha discontinuity(i.e., bifurcation) limitingtheirranges.Fig. 2 demonstratesthat domiresultingin a radicalchangein thesystemstate.O'Neill nant species of two biomes-Great Plains Short-Grass
et al. (1989) discussed potentialmethodsfordetecting Steppe and ChihuahuanDesert-have been able to in-

TABLE 1.

August1993

ECOTONE HIERARCHIES

373

mentpatterns,whichare influencedby the interaction

of environmentalconditionswith the micrositeenvironments.The netresultofthepatchpatternsforthese
species is increased patch heterogeneityand a finegrainedmosaic patternin the ecotone (Neilson 1991;
Fig. 1). The transectrepresentedin Fig. 2 is one of
manypatchecotonesin thebiome mosaic. The pattern
is expectedto be common across virtuallyall ecotones
in the region.
The resultsshown in Fig. 2 representa snapshotin
time of the changingpatterns.A comparison of site
photographsfrom1915 and 1900 demonstratesmarked
movementof L. tridentatain the ecotonal area of the
transectand a movement of the mosaic patternhypothesized in Fig. 1 (J. Betancourt, unpublished
photographs).Aircraftphotographyis a valuable technologyforevaluatingthe ecotonal dynamicsof broader-scalefeatures(Fig. 3). Studiesofa seriesofair photos
(1935-1984) have identifieddirectionalmovementof
populationsoftheselife-forms
and mosaic patterns(B.
Musick, unpublisheddata). Long-termmeasurements
of the line-intercept
transectsin the 1-km2studyareas
(Fig. 2) will quantifythe dynamicsof this broad-scale
mosaic pattern.The photos also detect broader-scale
I
-I
patternsof otherspecies (e.g., B. eripoda, B. gracilis)
I km
controlledby soil substrates.In Fig. 3, thedark-colored
mo- patchesare dominatedby B. gracilison Haplargid soils
(scale= 1:10000) showing
FIG. 3. Aerialphotograph
speciesintheSevilletabiometransition and the light-coloredpatches are B. eripoda on Calsaicsofthedominant
area. In thisimage,the darkerpatchesare dominatedby ciorthid soils. The small dots identifykangaroo rat
patches
Boutelouagracilison Haplargidsoilsand thelighter
are B. eripodaon Calciorthidsoils.The smalldot features (Dipodomys spectabilis)mounds. This rodentmay be
an importantvectorin the movementof plant species
mounds.
spectabilis)
kangaroorat(Dipodomys
identify
across the ecotone, throughdisturbanceand dispersal
activities.
vade (or be eliminated) to differentdegrees along a
The mechanisticexplanations for the dynamics of
transectin the Sevilleta LTER studyarea. The species the mosaic patternscome fromstudies of lower level
arrowsindicaterelativedirectionsand extentof move- ecotones. Finer scale studiesalong the transecton the
ment of these three species along the transect.The SevilletaNWR demonstratethedynamicsofplantedggraphs show the abundance and sizes of patches of es. Figs. 4 and 5 show patternsof movement for inthese dominants.The graphresultsare based on 1600 dividual plantsduringdry(1990) and wet (1991) summ of transectlines (1-cm resolution)in each of three, mer conditions. The figuresare computer-analyzed
1-km2study areas-Deep Well, Five Points, and La
photographsof 3 x 4 m plots taken with a boomJoya-along the 10-kmgradient.The patternsof these mountedcamera. The plots were photographedin the
threespecies supporttheconceptsoftheindividualistic springand autumn. The photographswere digitized
nature of movementand the change in patch scale as and classifiedinto plant and bare ground areas, and
thedistributionlimitis reached.In thiscase thepatches the images were overlaid to identifychanges for the
also are population mosaics since the patch was iden- interval. This technologyprovides a nondestructive
tifiedby the occurrenceof a singledominant species. analysis of the same plot throughtime. Traditional
Each species demonstratesa decreasingpatch size as methodsofvegetationanalysistendto be slow because
it approaches the limits of its distributionalong the theyare labor intensiveand involve destructivesamtransect.Bouteloua eripoda and Larrea tridentataare pling(e.g.,clipping).As a result,theyare oftenconfined
Chihuahuan species; however,B. eripodahas extended to relativelyfew and small, local area measurements,
furthernorththan L. tridentata.Bouteloua gracilis is and rely on statisticaltechniques to quantifychange
a Great Plains species and its patch size decreases to ratherthan repeat analyses of the same vegetation.
the south. The decreased patch size in these ecotonal With image-processingtechniques such as that used
areas is hypothesizedto be a response to increased forFigs. 4 and 5, numerousplots (i.e., manyhundreds)
sensitivityto micrositeconditions at the range limits can be analyzedthroughoutthetransitionarea to quanof a species. It also reflectsthe individualisticnature tifythe fine-scalechanges associated withbroad-scale
of species movement;i.e., germinationand establish- changes in patch pattern.The image-processingsoft-

JAMES R. GOSZ

374

.....

......l*..

~~~~~~~~~*m

EcologicalApplications
Vol. 3, No. 3

sandwichedbetweenperiodsof winterwet,
drought,
but onlywhenthe drywinterwas followedby high
of conditions
summerrainfall.Unique combinations
to increaseitspatchsize
also mayallowL. tridentata
and extensionto thenorth.A trendof drysummers
combinedwithwetspringscould cause B. gracilisto
and a movementof the
changeits mosaic structure
the
to thesouth.Theseresultsidentify
mosaicpattern
(i.e.,
or periodicities
oftemporalpatterns
importance
and preof temperature
temporalscales)in extremes
linkwith
cipitationin thisregionand theimportant
Thelong-term
studiespossiblewith
ecotonedynamics.
of thecondithisLTER sitewillallow identification
speciesin thistrancontrolling
tionsand constraints
sitionzone.

change
imageof vegetation
FIG. 4. Computer-generated
fromspring1990 to autumn1990 on a 3 x 4 m plot.The
and evaluation
Cross-scaleextrapolation
areasthatdidnotchangeduring
vegetated
darkgreyidentifies
baregroundthatbecame
whileblackidentifies
theinterval,
scales and
of studiesat different
The combination
bare
byautumn.Lightgreyidentifies
coveredbyvegetation
above,allowstheextrapolation
The summerof overtime,illustrated
soil thatdid notchangeduringtheinterval.
oftheconbetweenscalesneededforan understanding
precipitation.
1990had below-normal

trolson thedynamicsofthemosaicof thetransition

nestedsampling
themulti-level,
wareusedin thesestudiesis Khoros-a robustsystem zone.Fig.6 represents
ecotonestothedynamics
pro- designthatrelatesplant-edge
theclassification
thatmakesitpossibletoperform
is the
new ofthebiomeecotone.Remotelysensedimagery
toolsto construct
thenecessary
cessbysupplying
to evaluatebiomedynamicsof the
and graph- likelytechnology
visualization
and by supplying
algorithms
thatevaluateimtools(Rasureet al. 1990).The sys- entirearea,andthenewtechnologies
ical user-interface
ecotone,andpatch
production agesattheplantecotone,population
temhas been developedinto a turn-key
acrossthosescales.
andclassification ecotoneallowthebasicintegration
toallowrapiddigitization
technology
ofthemanyphotosbeingcollectedon theSevilleta. These studiescan be coupledwithtraditionalplotstudiesto reducetheothofB. eripoda clippingand line-intercept
TheplotsinFigs.4 and 5 havemixtures
to a minimumwithand B. gracilis.Fig. 4 showsthattherewas virtually erwiseintensefieldmethodology
andplots
The studytransects
no changein plantcoverofeitherspeciesforthedry outa lossofinformation.
summerof 1990. The plantcoverwas 67% in both are expectedto be diagnosticof dynamicsthatoccur
ecotonesin thearea.Futurestudieson
Juneand Octoberand thesmallamountofarea that on themultiple
changedfrombare groundto vegetation(13%) was
causingconversion
balancedby mortality/senescence
ofplantcoverto bareground.Duringthewetsummer
of 1991 (Fig. 5), B. eripodawas verydynamicand
fortheplantcoverincrease
responsible
almostentirely
from49 to 80%. This was caused by expansionof
areas. B.
individualplantsinto formerbare-ground
eripoda(Chihuahuanspecies)is expectedto respond
(i.e., shallowroot
to summerprecipitation
primarily
whileB. gracilis(GreatPlainsSteppe)maygain
system)
that
a competitive
edgefromthedeepersoil moisture
B. ericonditions.
wetwinter/spring
occursfollowing
poda is able to expandrapidlyduringyearswithwet
themechanistic
tillering,
vegetative
through
summers
in thetrancauseofpatchareachangesdemonstrated
sect studies.The highseed productionduringthese
of additional
yearsalso mayallowtheestablishment
ofnewpatchesinthemosaic,and
plants,theinitiation
FIG. 5. Computer-generated
change
imageof vegetation
that fromspring 1991 to autumn 1991 on a 3 x 4 m plot. The
ofitsrange.Neilson(1986) reported
an extension
theperiod1915to 1968hadsignificant darkgreyidentifiesvegetatedareas thatdid not changeduring
only7 yrduring
interval,while black identifiesbare ground that became
of B. eripodaon the
and establishment
seed production
covered by vegetationby autumn. Light greyidentifiesbare
New soil thatdid not change duringthe interval.The summer of
Desert)insouthern
theJornadasite(Chihuahuan
duringperiodsofwinter 1991 had above-normal precipitation.
Mexico.Theseyearsoccurred

Z~~~~~~VS

August1993

ECOTONE HIERARCHIES

375

Sevilleta LTER
imagery

0
0

~~~~~~Topography
Digital
m~~~~~~eleaVstion

Biome Ecotone

~~~~\
m~~~~~~~mp

F2

~~~~~~~~~~~Soil

>

maps

Weather

~~~~~~~~stations

l
l
+

Io 8 0
0

..

~~~~~~~~~~~~~~Vegetation

~~~~~~~~~~~~~~~~transects

Patch Ecotone
Population Ecotone

SPO

Ai
;rcraft

Soil
classification

Photot
plots

~~~~~~~~~~~~~~competition

studies

Plant Ecotone

~~~~~~~~~~~~~~~~~Plant

cheiistr,
Moisture

Plant

~~~~~~~~~~~~~~~genetics

Temperature
Soil
fauna
flora

FIG. 6. Multi-level,nestedsamplingdesignthatrelatesplant-edgeecotonesto thedynamicsofthebiome ecotone. Different

satelliteimagery(SPOT, TM, AVHRR)
technologiesallow integrationover the scales representedby theseecotones.Different
provides a rangeof scales and reflectancemeasurements.

additional ecotones will be used to validate the extrapolationsof currentresults.

New capabilitiesin supercomputingand networking
also will make cross-scale extrapolationfeasible. For
example, currentplans are to obtain color aerial photographyat a scale of 1:7920 (1 cm = 79.2 m). Each
photographwould cover 3.28 km2with the resolution of the color film(Film Code 2448) allowing the
distinguishing
of an object on thegroundthatis 18 cm
in diameter(R. Parmenter,personal communication).
The photoswill have 70% overlap along the flightpath
to allow stereo imagery.It is estimatedthat 1000 of
these photos would be needed to cover the 1300-km2
area of the refugeand borderlineareas. These photos
could then be used to create digitized orthographic
photos that would be incorporatedinto the Sevilleta
GeographicInformationSystem(GIS). Decision-based

rulescan be used to subsampletheentirearea to create

scales. For example, subsampling
images at different
oftenresultsin the loss of informationon spatial patternin the originaldata. It is similarto averagingout
variation. A rule that subsamples but maintains the
fractaldimension of the fine-scalepatternproduces a
broad-scale pattern without losing structuralinformation(B. Milne,personal communication).Scale-dependentanalysesoftheorthographicdata setthenprovide standards for analyses that compare different
technologies (satellite vs. aircraftvs. ground-based
transects).The analysesalso allow correctionsforproblems withresultsthatare dependenton the technique
used (i.e., differenttechniques utilize differentasThe multiple
sumptionsthatinfluenceinterpretations).
image-GIS analdata sets allow calibrated/referenced
yses that provide needed informationon vegetation-

JAMES R. GOSZ

376

Ecological Applications

Vol. 3, No. 3

soil relationships,plant distributionpatterns,and the are changed.Finally,suchdata bases act as thestandard

to allow comparisonsof othertechniques(e.g., satellite
extentand patternof animal disturbancesin the ecoimagery,line-intercepttransects,etc.). Such a multitone hierarchyof this biome transitionarea.
In view of the enormous data storage,image-pro- scale, multi-technology
approachwillbe a necessityfor
cessingsoftwarerequirements,and computingrequire- the challengewe have set for modelling ecotone dynamics.
ments to process data fromthe entirearray of 1000
images,itwillbe necessaryto develop interactivecomputingwithsupercomputerfacilities.Each color photo
ACKNOWLEDGMENTS
requires 232 megabytes(MB) of storage(77.4 MB in
Many individuals in the Biology Departmentof the Unieach ofred,green,and blue bands) and the 1000 photos versityof New Mexico as well as otherinvestigatorson the
Sevilleta LTER have contributedto the ideas being pursued
necessaryforthe entirearea require 232 320 MB (G.
Shore,personal communication).We intendto utilize in this biome transitionarea. The interactivegroupresearch
that is
LTER programhas contributedboth
thenationalnetworkto initiateprocessingrequestsfrom the allowed by the
supportand discussion that makes this paper possible.
theSevilletacomputers,have processingperformedon The U.S. Fish and Wildlife Service manages the Sevilleta
a supercomputer (e.g., San Diego [California]
National WildlifeRefugeand theirsupportand encourageSuperComputingCenter) with data in storage there, mentalso makes all ofthispossible.JamesBrunt,GregShore,
and outputtheresultsback to the Sevilletacomputers. and Troy Maddux performedimage and data analyses. Financial supportwas provided by the National Science FounThis approach will dramaticallyincreaseour abilityto dation (BSR-88 11906). This is contributionnumber35 to the
performthe scale-dependentanalyses and cross-scale Sevilleta LTER program.
extrapolationscalled for in our hypothesesof biome
LITERATURE CITED
transitiondynamics.
CONCLUSIONS

This discussion of ecotone hierarchiesemphasizes

the scale-dependentnature of patternsand processes
associated withecotones. An edge at one scale may be
indistinguishable
at a different
scale. However,theprocesses thatoccur at a certainscale can have significant
influenceson both broaderand finerscales. Measuring
and understandingthese cross-scale influencesrepresents a significantchallenge for ecotone studies. The
basic strategymust be studiesperformedat a number
of scales, and thispaper suggestsone hierarchicalclassificationto identifyappropriatescales (i.e., plantedge
to biome edge). Different
technologiesand approaches
must be used at these different
scales, requiringadditionalstudieson integration
and calibrationofthetechniques. A gradientanalysisin a biome transitionzone
is one example where the different
technologiesare
performedon the same transectsto allow such crosscalibration.This analysisalso allows the identification
of the fine-scalepropertiesthat determinechanges in
propertiesin the nextbroaderscale. The highvariance
associated withfine-scalemeasurementsrequireslarge
sample sizes; however,some techniques,such as repeat
photographyon areas stratifiedby factorsthat act as
constraints(e.g., topography,soil characteristics),can
detectchange witha minimal sample size.
The availabilityof supercomputingtechnologiesallows additional capabilitiesforcross-scaleintegration.
Complete area photographyat fineresolutionis possible, and the storage and computingcapabilities at
these supercomputingcentersallows analyses of these
huge data sets. The same digitized data base can be
analyzedat fineto broad scales by a varietyofdecisionbased rules that retain the propertiesof the different
scales (e.g., fractals).These analyses also are able to
identifythe emergentpropertiesthat occur as scales

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