The Ecology of Interfaces: Riparian Zones Author(s): Robert J.

Naiman and Henri Decamps Reviewed work(s): Source: Annual Review of Ecology and Systematics, Vol. 28 (1997), pp. 621-658 Published by: Annual Reviews Stable URL: http://www.jstor.org/stable/2952507 . Accessed: 17/07/2012 09:46
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Annu.Rev.Ecol. Syst. 1997.28:621-58 ? Inc. reserved Copyright 1997 byAnnual Reviews All rights

THE ECOLOGY OF INTERFACES: Riparian Zones

Robert NaimanandHenriDecamps J.
Washington of Seattle, SchoolofFisheries, 357980,University Washington, Box and National la Recherche de 98195 USA, naiman@u.washington.edu Centre Continentaux, rue 29, Centre Aquatiques Scientifique, d'Ecologiedes Systemes Jeanne e-mail:decamps@cemes.cemes.fr France; Marvig, 31055 Toulouse,
KEY WORDS: corridor riparian, buffer, interfaces, ecotones,

ABSTRACT of diverse array speciesandenvironmental Riparian zonespossessan unusually geographis to flood regimes, processes. ecological The diversityrelated variable influences altitudinal climate shifts, upland and icallyuniquechannel processes, on thefluvial environment a of dynamic supports variety corridor. resulting The adapted cyclesand rates, organisms and life-history strategies, biogeochemical in to disturbance scales. Innovations regimes overbroadspatialand temporal in zone management beeneffective ameliorating have many ecological riparian zonesplayessential issuesrelated landuse andenvironmental to quality. Riparian and planning, restoration aquaticsystems, in in of rolesin water landscape and for efforts. cooperation these catalyzing institutional societal and

INTRODUCTION
First described nearly a centuryago (36), interfacesbetween environmental change disconor systemproperties patches occur where structural functional tinuouslyin space or time. which describe interfaces The termstransition zone, ecotone, and boundary, in (91, 182), are used synonymously thisreview. Interfacesbetween adjacent uniquely definedby space and ecological systemshave a set of characteristics between the adjacent ecological time scales and by the strength interactions of systems (102). Thus, interfacespossess specific physical and chemical atbiotic properties, and energyand materialflowprocesses, buttheyare tributes, 621 0066-4162/97/1120-0621$08.00

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(183, 231). The with ecologicalsystems uniquein their interactions adjacent and scales, which vary over widetemporal spatial strength these of interactions, or between adjacent resource patches ecological is controlled thecontrast by In to may of analogous a semiperunits. general, interface bethought as being an between adjacent the of and meable membrane regulating flow energy material control energy and haveresources, environmental patches(183). Interfaces between biological sensitive sitesfor interactions material flux, potentially are highbiodiversity, variables, haverelatively populations their and controlling and maintain species,and are refuge critical habitat rareand threatened for may (91, sourceareasfor pestsandpredators 102,183,231). Some interfaces or zones) alsobe sites longitudinal for migration alongwindbreaks riparian (e.g. and genetic (e.g. forest/agricultural pools or sitesforactivemicroevolution are ecosystems between terrestrial freshwater and interfaces-76).Interfaces change(155, 182). Examplesinclude particularly sensitive environmental to lakes and lake zones,floodplain wetlands, littoral riparian forests, marginal Thisarticle rewater forests, areaswith and groundwater-surface exchanges. encompass with streams rivers and becausethey views riparian zonesassociated important interfaces. for mostofthecharacteristics and Natural zonesaresomeofthemost diverse, dynamic, complex riparian habitats theterrestrial on portion theplanet(184, 185). This of biophysical of article an of characteristics riparian zones, provides overview important on describes effects adjacentenvironments, summarizes ecological physical of alterations on characteristics, discussesconsequences environmental and ecosystem form function. and

THE RIPARIANZONE
is "of to Riparius a Latin wordmeaning or belonging thebankof a river" term New Universal Unabridged Dictionary 1976). The anglicized (Webster's and lakes. to on riparian refers bioticcommunities the shoresof streams mosaicof landforms, zones are an unusually diverse communities, Riparian andenvironments within larger the and serveas a framework landscape, they and of forunderstanding organization, the diversity, dynamics communities of associated withfluvial ecosystems 84, 183,184,185). A variety natu(53, few mosaicwith a and environmental raldisturbances creates spatial temporal in parallels other systems.

Zones and Riparian Defining Delineating

The spatialextent theriparian zone maybe difficult delineate to precisely of and in of strategies becausetheheterogeneity is expressed an array life-history whilethefunctional attributes successional dependon community patterns,

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the composition theenvironmental and setting. riparian The zoneencompasses portion the of stream channel between low andhighwater the marks that and where vegeterrestrial landscape from high the water mark toward uplands the and tation maybe influenced elevated by water tablesorflooding bytheability ofthesoilstoholdwater researchers). (184, 185;exactdefinitions among differ vegetation Thewidth the of riparian zone,thelevelofcontrol thestreambed that attributes in(e.g. hasonthe stream environment, the and diversity functional of the formation biogeochemical flow, cycles)arerelated thesize ofthestream, to of within drainage regime, and position thestream the network, hydrologic the thelocal geomorphology zone maybe (53,126,182,236,238). The riparian embedded smallinthenumerous headwater streams arealmost that completely in theforest.In mid-sized beingreprestreams, riparian the zone is larger, is by by bandofvegetation whosewidth determined long-term senfted a distinct zones channel and Riparian (>50 years) dynamics theannual discharge regime. of largestreams characterized well-developed physically are but complex by channel with lateral of migration, floodplains longperiods seasonalflooding, a community, moist and oxbowlakesin old river channels, diverse vegetative is influenced the by soils (155,240). Vegetation outside zone that notdirectly matter hydrologic conditions that but contributes organic (e.g. leaves,wood, or influences physthe dissolved to or materials) thefloodplain channel, that ical regime thefloodplain channel shading, or part of by maybe considered ofriparian zones(26, 84). Theseattributes that zonesarekey suggest riparian for they may systems regulating aquatic-terrestrial linkages (182,271) andthat provide early indications environmental of change (52, 183,230). reaDefining riparian zonesis important both for ecologicalandmanagerial sons. Riparian from stream a where landuse buffer distance zones,a defined an are are activities restricted stream for protection purposes, becoming incommon tool generally creasingly management (20,80, 136,242). Definitions of extent herbaceous incorporate ecologicalcharacteristics as thespatial such of resources aquatic for to plants adapted wetted soils,production nutritional localgeomorphology, areaofsediment and (84, systems, generation 136,242). modelevation information Increasingly, geographic systems (GIS) anddigital zone els (DEM) arebeing an estimate riparian areaand of usedtoprovide initial has in the distributiondrainage networks vegetation (13,274). Where riparian to the beenremoved, methods beingdeveloped determine spatial are potential for regeneration its (136,210).

Life-History Strategies
Streams nonequilibrium are effects thebiotic on charthat systems havestrong of The and are acteristics riparian communities. active channel floodplain harsh and environments theestablishmentplants animals for of (185). Herewe focus

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on themorphological, and of physiological, reproductive strategies thevegetationeventhough equallyinformative an review couldbe madefortheanimal community 225). (see Seasonalvariations discharge wetted in areascreate and environmental conditions challenge that eventhemosttolerant species.Nearly every year, most riparian plantsare subjected floods, to erosion, abrasion, drought, freezing, and occasionally of in toxicconcentrations ammonia addition thenormal to biotic the of are challenges; life-history strategies most riparian plants suchthat extreme conditions either are endured, resisted, avoided(4, 86, 185). or In general, riparian plantcommunities composedof specializedand are disturbance-adapted species withinmatrix less-specialized less-frequena of and tlydisturbed upland forest (185). The classification plants of intofour broad categories functional of adaptations useful understanding is for processes leadingtoriparian forest succession distributional and patterns. 1. Invader-produces of and large numbers wind- water-disseminated propagulesthat colonizealluvialsubstrates. 2. Endurer-resprouts breakage burial either stem roots or after of the or from floods after or eaten. beingpartially 3. Resister-withstands flooding weeksduring growing for the season,moderate or fires, epidemics. 4. Avoider-lacksadaptations specific to disturbance types;individuals gerin minating an unfavorable habitat notsurvive. do Some widely the of distributed stratespeciesillustrate variety life-history and gies. Sitkawillow(Salix sitchensis) Scouler'swillow(Salix scouleriana) arepioneer a of plants welladapted living to under number disturbance regimes. if or Individuals thesespeciesinvadepost-fire of landscape resprout theroot remains in as endursystem intact a fire (185). Theyare wellsuited invaders, In conditions. contrast, ers,or resisters depending uponlocal environmental Sitkaspruce(Picea sitchensis), whichcan colonizewoodydebrisand mineralsubstrata thefloodplain 165) andresist and on (94, both flooding sediment where is a regular fire deposition, notpersist does occurrence. in Floodtolerance trees In includes bothmorphological physiological morand adaptations. general, and phological adaptations suchas adventitious roots,stembuttressing, root in soil subare to flexibility riparian plants a response either anoxiaorunstable strata. in Morphological adaptations anoxiabyvascular to plants periodically flooded air areasinclude spaces(aerenchyma) theroots stems allow in and that
MORPHOLOGICAL AND PHYSIOLOGICAL ADAPTATIONS

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forthediffusion oxygen of of from aerialportions theplantto therootsand adventitious that roots grow zoneenabling abovetheanaerobic oxygen absorption theplant.Thedevelopment these by of structuresmediated increased is by levelsof ethylene, production which initiated anaerobic conthe of is by soil ditions (17). Root and stemaerenchyma common speciesof families are in Cyperaceae Juncaceae, and which normally are found poorly on drained floodplains.Adventitious occur a variety tree in of roots genera (e.g. Populus, Salix, in Alnus, Sequoia) living riparian environments where sediment and deposition wetted soilsarecommon (17,248,252). Flooding also mechanically disturbs plants eroding by substrata byabraand sion. Stemflexibility among woody genera (i.e. Populus, Salix,Lanes) imparts and endurance resistance potentially levelsofshearstress to high accompanyingseasonalfloods.Floodsoften occurduring periods whenthevegetation is without leaves,further reducing potential damage(73). Anoxicconditions challenging plants onlybecausethey are to not needoxymobilize soluble reduced ions(suchas gen,butalso becauseanoxicconditions manganese) aretoxic(171). Rhizosphere that reduces threat this oxygenation from rootto theadjacent to form very soil a smallbut by moving the oxygen effective oxidizedzone. However, riparian of species show a largevariety to in responses flooding. example, thetropical For forests Brazil, of gallery Sebastianaklotzchyana accelerates withethanol themajorend as glycolysis of product anaerobic metabolism without but detectable diffusion to oxygen therootsystem maintains aerobicroot (123), whereas Hymenaeacourbaril metabolism (witha 50% decreasein metabolism) diffusion through oxygen from aboveground the stemsystem. Chorisiaspeciosadevelops hypertrophic lenticels improve aeration therootsystem, to the of although does notreach it fullmetabolism. fourth A whichdoes not species,Schyzolobium parahyba, accelerate glycolysis to enough maintain rate energy the of production required does notgrow that bytheroots, in environment (123).
REPRODUCTIVE ADAPTATIONS Plantlife-history includea suiteof strategies co-adapted characters enhance that reproductive successin specific environments Theprimary (9). reproductive characteristicsriparian of are plants tradeoffs between sexualandasexualreproduction, size, timing dormancy, seed of of seed and timing seeddispersal, dispersal For mechanisms, longevity. example,several plants (e.g. Populus, Salix)disperse seedsinphasewith seasonal the retreat floodwaters, of ensuring moist seedbeds successful for and germination plant colonization (120,248). Manyspeciesuse transport flowing by water, a known hydrochory, dispersal seedsas wellas vegetative phenomenon as of for to fragments newsites(116,196,198). Bald cypress and (Taxodium distichum) water tupelo(Nyssaaquatica) in a SouthCarolina swampforest on water rely

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NAIMAN DECAMPS &

in levels smallincreases water seeds(244). Further, wind disperse to more than communities. Dispersal animals by adjacent plant tend introduce to seedsfrom may but and (zoochory) bywind(anemochory) be evenmoreimportant, little comparison. empirical existfor data floraand also structures riparian dispersal for The mechanism propagule et In patterns. Sweden,Johansson al mayhelp explainspeciesdistribution of floating capacity thediaspores between a relationship (116) found positive Seeds of water-dispersed vegetation. in and speciesoccurrence theriparian between timedid notdiffer seeds. Floating better thanother speciesfloated or by seedsdispersed animals wind. (e.g. asexually).Otherwise vegetatively plantsreproduce Many riparian whichare shedin theprofor tips,of cottonwood, example, healthy branch or (59, cess of branch abscission, cladoptosis 81), maydevelopadventitious (164). identical trees roots growintogenetically and

Patterns Successional and Vegetative

geomorpholamonghydrology, PHYSICAL CONTROLS Complexinteractions and dynamics, compoinfluence structure, the temperature, fire and ogy, light, (and suggests hydrology that zones (23, 155). The literature sition riparian of For factor. example, with itsinteractions local geology)is themostimportant vegetative patch topography, in riparian a having ridge-and-swale floodplains lows to alternate between thoseon topographic adapted longhydroperitypes in highswith speciesalso found mesicuplands ods andthoseon topographic (23, 168,240). of Brinson's modelis that powerandfrequency inundation (23) conceptual high-power, lowfrom proportional existin a continuum and are inversely high-frequency to floods affect wholefloodplain low-power, that the frequency channel.The former to floods that influence theareaadjacent thewetted only of for to create that features persist hundreds thousands years largegeographic floods intermediate-frequency relic levees).Medium-power, lakes, (e.g. oxbow of to that of determine structure havelifetimes tens hundreds patterns ecosystem is at of years. Treecommunity zonation influenced thisscale becausemany floods times (8,96,109). The low-power treespecieshavesimilar generation short-term such as seed germination thatoccurannually determine patterns in Colorado, and seedling survival 137,138). For example, southwestern (8, in are of (Populusangustifolia) mostabundant years seedlings cottonwood and withcool winters, springs, cool, wetautumns.Bothgood seedling wet in of with winter blocking storms yearsare associated yearsand stand-origin is Arizonamonsoon needed but late-summer theNorth Pacific, a persistent 3-4 forsurvival Good seedling (aboutevery yearsoccurmorefrequently (8). 10-15 years). stand-origin (aboutevery years years)than

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of and to of Theability soilsandsediments holdwater theexistence tributary distrivegetative in flows and groundwater areequallyimportant determining and the bution (23,50,109,131,262). Distancefrom river microtopographic in rising discharge themainchannel between lag determine times variations of the has Oncewater arrived, composition thesoil on-site. andarrival water of how determine long rateofevapotranspiration) (as andthealluvium wellas the to can water sources allowvegetation External saturated. remains thesubstrata older (14,50, 131,262). Indeed, regime of independent theflood largely persist riparmature and camaldulensis), most river cottonwood, redgum(Eucalyptus because presumably creekwater, rather nearby than use ian trees groundwater source(50,155). is groundwatera morereliable forest developsis conuponwhichtheriparian template The geomorphic (23, 180). The regime changeinducedby thedischarge undergoing stantly a representsmosaicofsites to from drainage network, headwaters theestuary, a state. Even sites or degrading, maintaining steady that maybe aggrading, is materials of movement deposited wherethedownstream state, in a steady will channels the upstream, stream from of balanced thetransport alluvium by features so and laterally down-valley thatthephysical to continue meander riparian gallery to of theriparian zone continue change(68). Levees support in result but normally forests mayfloodfrequently, thecoarsedeposits that of hydric the levelsdrop.Oxbowlakesarethemost when water drainage rapid and flooding anaerobic to speciesadapted constant supporting habitats, riparian soils. (68). fluvial geomorphology describes A richandcomprehensive literature in patterns processes and for important understanding particularly Twoaspects channel and erosion deposition, lateral and are vegetation site-specific riparian may migration be slow (cm yr-1)to fast(102 m Lateralchannel migration. (12,23,239); hydraulics and on yr-1),depending thetypeof stream channel of and the thissubstantially influences composition demography thevegetative communities 113,127). (111, Rates and activity. uponlanduse,climate, tectonic supply depends Sediment anmeters to a of erosion and deposition rangefrom fewmillimeters several in whichhas been occurring theAndesMountains activity, nually.Tectonic for forest mechanism riparian for to speciation appears be a central millennia, Amazonian basin(226,239). rich in communities thefloristically upper on vegetative community and Thephysical influences light temperaturethe of or Understory arelesswellinvestigated either than hydrology geomorphology. toward declines forest edgebutrapidly at tends be highest theriparian to light levels where trees), by theforest interior (unlessgapshavebeencreated fallen are densities diversity and seedling are often However, <2% offullsunlight. that in with suggesting other light notcorrelated variations understory intensity,

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NAIMAN DECAMPS &

and (152,219,220). factors moreimportant germination establishment are in is different upland from forests, nostudies but Thetemperature regime markedly to (26). Compassorientahaveattempted linkthis to difference thevegetation regimes, significant and floristic diftiondoes influence andtemperature light and zonesonnorth- south-facing slopes(256). ferences appear between riparian in zones of humid where mostvegFireoccursrarely theriparian regions role etation cannot withstand evenmildfires (185) butplaysa significant in Colorado River, nearly 40% oftheriparian vegdrier climates In thelower (4). withhalophytic shrubs recovering faster than etation burned within years, 12 mesophytic (30). trees As corridors within watersheds, riparian zoneshavea unique on of controls themovements water, longitudinal pattern exerts that substantial (76) eight comnutrients, sediment, species(76, 155). Forman recognized and ranging from strictly linear (mostly uplands) monshapes riparian for corridors, that to highly variable valley bottoms). Cross-sectional profiles (low-elevation setan third dependuponthelocal geographic provide important dimension onesdownstream and complex ting.Thesimplest tobe upstream themore tend (76,235). on influences riparian vegetation, Despitestrong hydrologic geographic and herbivory, soils,and disease) are ecologicalinfluences (such as competition, is in reEven though competition probably significant shaping communities. duced because of frequent a does exist. hierarchy disturbance, competitive beyond their Thereis evidence that somespeciescouldexistin environments in for of except thepresence present range, specifically thedirection lessstress, ofcompetitors (129). In regions influences herbivory exerts strong with intact animal populations, activities assoonvegetative and characteristics 190). Thephysical trophic (33, the that consequences go far beyond ciated with herbivory ecosystem-level have is (121, 124,179). Thehabitat requirements individuals foodandhabitat of for and animals they burrow wallow of as further modified theactivities larger by is The result that insoilsandbuild activities. net damsonstreams, among other and is of theheterogeneityriparian habitats resource (or patches) increased, the rates elements of distribution cycling and (suchas N andP) aremodified. also Soil conditions the of pat(especially degree saturation) coincidewith terns sediment of size and microtopography, affecting distribution plant grain in Disease later this review. from river the the to uplands (250) andarediscussed as an agent biotic is studied, although pests(suchas influencing patterns little are to Choristoneura fumiferana) known spread rapidly budworm caterpillar, corridors alongriparian (76).
BIOTIC PATTERNS

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than of forests as great greater that is or In general, basalareaofriparian the within largely due valuesvarywidely regions, uplandforests (23). Although Riparian less an of to stand age,thevariation usually than order magnitude. is tropics tendtoward forests thesoutheastern in UnitedStatesand thehumid and stem and greater density basal area thanthosein morearidregions more northern latitudes (23). Theaboveground forests ranges between and300 tons 100 biomass riparian of few 1-10% ofthe total.In general, ha-l with exceptions (23). Leavesrepresent from biomass aboveground biomass, ranging belowground tends be lessthan to in haverelatively rates production high of 5% to 120% of it. Riparian forests in with forests. Muchofthe variation apparent belowground comparison upland and biomass-12 to 190 tonsha-l maybe due to sampling methods siteconditions. valuesforaboveground rangefrom production specific Published fall averages47% of the annualproduction. 6.5 to 21.4 t ha-' yr-1; litter or gradients.Rates The limited data do notreveallatitudinal successional in of no attention studies of belowground havereceived virtually production are limiting factors thatthere no strong riparian forests.The data suggest in associated with or result unusually production. low that water nutrients would to zonation often existsas a transverse gradient perpendicular the Spatial to by responses localvariations wetted channel is madecomplex vegetative that typically to patterns in topography susceptibility flood(236). Vegetative and follow in features patterns (273) anddisturbance predictable patterns physical forest succession a (185). Decamps et al (54) presented modelof riparian fortheGaronne in occur cyclicalsuccessional processes River, France, which within floodplain and are the where flood-induced erosion deposition common. on terraces without the repeated flooding, successional However, thehigher are dominate. and autogenic forces dynamics notreversible internal in to The earliest of zonesdidnotrefer studies vegetation dynamics riparian illustrated process.Many the theconcept succession of (75) butnevertheless in but areas are primary succession, an equal successional patterns riparian or of beginwithplantfragments, propagules, number successional patterns from communities flood, wind, biomass (155). Avalanche, remaining previous influences the on vegetation and physical fire, drought, disease, herbivory, other in successional leaveuniquebiotic are patterns legaciesthat displayed various them recover to possessadaptations allowing (185,201). Manyriparian plants on root fragandreproduce rootsuckering, adventitious development plant by to remaining initiate of material ments, stem and flexibility. amount biotic The and succession itsviability and on intensity, frequency, duradepend thetype, the tionofthedisturbance Disturbances prepare siteforinvasion also by (53). additional speciesfavored thenewconditions. by

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PHYSICAL FUNCTIONS OF RIPARIANZONES Mass Movements Materials of and Channel Morphology

Material supplied streams to comesfrom erosion stream of banks,a process influenced root by strength resilience and (85), as wellas from uplands the (178). Stream bankslargely devoidof riparian vegetation often are highly unstable and subject mass wasting, to which widenchannels several tensof can by to meters annually (111,239). Majorbankerosion 30 times is more prevalent on nonvegetated banksexposedtocurrents on vegetated as banks(11). Riparian vegetation modifies also sediment transport either physically by entrapping in materials, whichappearsto be mostimportant relatively lowgradient environments, by altering or channelhydraulics.In experimental channels, Kentucky bluegrass (Poa pratensis) entrains sediment thebase of at thevegetation, 30-70% retained with on dependent bladelength Accretion (3). of sediment organic and matter vegetation be substantial, can by especially floods during from (107, 146). Sediment deposition 1880to 1979on a coastal in UnitedStatesaveraged plainriver the southeastern 35-52 Mg ha-l yr-l (146). Alteration channel of is hydraulics accomplished either rootsor by by largewoodydebrisin thechannel low flowsand by stemsat highflows. at All provide physical structure slowswater, that decreasesstream power, and holdsmaterials place. Experimental in manipulations the involving removal of largewoodydebrishave resulted dramatically in increased erosionrates (16,151, 191). Erosional depositional and events shaping channel are morphology thesubject ofa largeliterature (68). In general, spatial introduced into heterogeneity thechannel either vegetation thelargewoodydebrisproduced the by or by thevegetation flows water of shapeschannel morphology redirecting and by sediment, and sorting sediments, either or materials retaining moving (110).

in Wood Streams Riparian and Zones

roles at the land-water interface Woodydebrisplays important biophysical Inthe onexposed instreams, (95, 157). alluvial and it riparian forest, substrates, in accumulates floods discrete conspicuous and during piles. Each pileusually includesat leastone largepiece of dimensionally complexwood (i.e. a key which resist member) can mostflow physically and smaller capture piecesof the of wood, making pile evenlarger.In temperate densities up to forests, 160 woodypiles perkmof stream bankmaybe found (251), butin tropical wheretermites prevalent, density woodydebrispiles is are of regions, the reduced in some substantially (190). Further, ofthis wood,especially thelarger pieces,maybe quiteold. Nansonet al (192) found wood > 17,000yearsold inthefloodplain a Tasmanian of havebeenmeasured stream; ages >300 years

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(1). of coastalrainforest Woodydebris forlargewoodin streams thePacific habitat. Depending on materials, form and pilesdissipate energy, moving trap in and water and they size, position thechannel, geometry, can resist redirect heterogeneous, power water becomespatially of to currents, causing erosive the patches theriparian in and thereby creating mosaicoferosional depositional a of channels, spatialarrangement pools is the corridor (173, 191). In steeper of debris water currents (173), so redirected independent theamount woody of have and erosional materials (157). Suchprocesses widen channel capture the in rivers (228). beenexamined by extensively geomorphologistsalluvial times also results longer in water residence (69), andthetemdebris Woody of removal wood of can Experimental porary storage materials be substantial. in matter rates thefirst toexceedbaseyear allowssediment organic and export experiments lineconditions several hundred percent (16,151). Analogous by similar insights. Leaves materials haveproduced with addition organic the of withwoodydebrismoveonly and smallpieces of wood added to a stream of traveled leavesandsmallwoodin 65% and8%, respectively, thedistance by carcassesadded streams without (69); 80% ofthesalmon largewoodydebris within are coastalrainforest streams retained woodydebris to nutrient-poor by 200 m ofthereleasesites(34). within habitat fishand macroinvertebrates the for Woodydebrisprovides within terrestrial stream channel 95, 157,167); itsroleas habitat the portion (5, is ofthe corridor only nowbeing Woody debris physically investigated. riparian and protects them (seedsandplant fragments) further retains plant propagules and (74, and,in somecases, drought herbivory 110). from erosion, abrasion, and are On exposedcobblebars,mostseedling germination survivorship associated with which a and moist, provides protective relatively woodydebris, for microenvironment. debris affords also protection small Woody nutrient-rich and of suchas mammals birds;thediversity abundance smallmammals and in greater areas withwoodydebris shrews, voles,and mice are significantly for use whileseveral birdspeciespreferentially woodydebris accumulations, and perching feeding 25 1). (61,

Microclimate
on but controls themicroclimate streams, there of forests exert strong Riparian water microclimate Stream arefewcomprehensive studies theforest of itself. soil are with and correlated riparian temperatures, strong temperatures highly in soil,andsurface and microclimatic temperatures inrelaappear air, gradients or tive but solar speed(26). Riparian humidity notinshort-wave radiation wind in stream cimates seasons,influence and discharge forests, especially warmer difficulReduced streamflow causesphysiological through evapotranspiration. tiesfororganisms coolertemperatures (100). preferring

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Riparian ZonesAs EcologicalCorridors

Riparian zones, as networks distributed largeareas,are keylandscape over components maintaining in biological connections alongextended dynamic and environmental gradients (184,218,220). Perhaps bestevidence plants the for usingriparian zones as corridors comesfrom exoticinvasions. Exoticplants rapidly movebothup and downriparian corridors preference overland in to routes (58). However, is notclearthat it riparian zonesfunction dispersal as in corridors all cases. Certainly, adaptations many of plants allow vegetative fragments seedsto float various and for distances (196, 198,241),whilemany in feces of other riparian speciesaredispersed wind animals by or (especially the birds).Schneider Sharitz & (244), for example, found seedsof Taxodium that distichum Nyssa aquatica could floatfor42 and 85 days,respectively, and covering downstream distances up to 2 km. of for Fewdatadocument riparian zonesas corridors terrestrial animals, despite thecommon assumption models.Twoexceptions theuse oftheriparian in are the between Central forest for alongtheGaronne River, France, birds moving in field Massifand thePyrenees (55), and a three-year experiment Alberta, of movements juvenile birds enhanced Canada,demonstrating through riparian of strips before after and harvest adjacent forest (153).

ECOLOGICAL FUNCTIONS OF RIPARIANZONES SourcesofNourishment: Allochthonous Inputs Herbivory and

from is for Organic matter riparian vegetation a sourceofnourishment aquatic from organisms (112,261). In temperate zones,valuesvary about200-900 g AFDM (ash-free mass) oflitter m-2 in smallandmedium streams to dry (1st 3rdorder) about20-50 g AFDM m-2 in larger to rivers (35,39,272). As a matter the of general trend, proportion coarseparticulate organic (CPOM; > 1 in For mm size decreases river increases. example, eastern as diameter) Quebec, from declinesexponentially 307-539 g AFDM m-2 in a 1stannuallitterfall order stream stream 15-17g AFDM m-2ina sixth-order to (39). Lateral inputs influenced size from soil surface notrelated stream butare strongly the are to and of structure entrapment organic matter byriparian during spring flooding (272). in in Similar local effects arefound intermittent streams Kansas also prairie wheretotalannualinput CPOM is lowestin theheadwater reaches(90). of Prior thewetseason,storage benthic to of CPOM in thedrychannel and on thebankis 320-341 g AFDM m-2in theupstream reaches 999 g AFDM and m-2 in the4th-and 5th-order of gallery forest reaches. The storage CPOM

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is channels where retention high, increases during wetseasoninheadwater the reaches, CPOM than downstream do reaches havemore with result these the that in is reaches. although bankstorage alwayshighest downstream litter influencing entering appearsto be themainfactor Riparian structure floor.Dependor from forest the laterally streams either directly transported rangefrom 52 zone,annualinputs coverof theriparian ingon thevegetative (64) and from to 474 g AFDM 63 to 295 g AFDM m-2 in Alaskanstreams laterally rainforest stream, stream (150). In an Australian m-2in a Moroccan in to varying response litter 6.8% ofthetotalannualinput, transported forms of and bankslope and microtopography The proportion direct laterally (15). influence in-stream comlitter streams maysignificantly transported entering (45). Even quality and timing as of munity dynamics a consequence input litter input, litter though laterally transported maynotexceed 10% ofthetotal due important a sourceof nourishment, to a higher as it maybe qualitatively (15). of directly thestream into than nitrogen concentration that leavesfalling highdischarge only the material maybe rearranged during However, organic floor litter may the periodsand notduring leaf fallperiod(156), and forest of amounts floods movemeasurable to effects several of require cumulative the (172). litter laterally thechannel to substantial In addition particulate riparian zonescontribute organic matter, to to Soil ecosystems. amounts dissolved of matter organic (DOM; <0.5 ,um) river zones water unsaturated regions riparian of directly from DOM mayoriginate discharge floods indirectly thesaturated or from through-flow atmedium during in zonealso caninfluence DOM originating theriparian rates (193). Soil water water. transfer subsurface of through macropore stream communities five At thescale of theAmazonbasin,McClain& Richey(158) identified blowlitterfall, to direct of organic matter streams: transfer pathways terrestrial stormin from soil surface groundwater baseflow, (e.g. lateral movement), the from and fringing wetlands.Directlitterfall overhanging flow, seepagefrom in comwere contributions similar massand elemental canopiesand blow-in to amounting 700 g m-2 yr-1. Freshand laposition amongall landforms, in Groundwater bile organic zonesexamined. material dominated all riparian acids of and organic baseflow DOM concentrations proportions hydrophobic werestrongly by correlated withsoil type(characterized old and refractory were overland by contributions dominated saturated molecules). Stormflow a wide spectrum disof floworiginating riparian in areas,whichtransferred Fringing solvedandparticulate fresh labilematerial. and organic matter, largely in of high wetlands contributed concentrationsDOM, particularlythelowlands. In contrast theother to matter wetlands dominated of is by sites,theorganic diverse within compositionally a dissolved compounds refractory hydrophobic the and molecular somewhat preliminary, results conceptual array.Although

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NAIMAN DECAMPS &

modelofMcClain& Richey (158) mayprove useful a framework quanas for tifying organic matter concentrations compositions and between contrasting trophic pathways. for Living riparian vegetation a source nourishment many is of animals, from insects mammals, can considerably system to that alter function their by feedingactivities. Outbreaks defoliating of insects alter can riparian forest productionandthereby water alter yield, nutrient cycling, streamwater and chemistry (254,263). Through selective browsing, largeanimalssuchas moose(Alces the from trees conifers, to alces) can shift riparian plant community deciduous altering formation nutrient soil and cycling ultimately and affecting proplant ductivity moosepopulation and dynamics (206). Beaver(Castorcanadensis) of also exert substantial a on and impact thestructure function riparian systems, successional enhancing floodplain complexity multiplying and vegetative paththe for ways,someofwhich affect landscape centuries (186).

Zonesas Nutrient Filters Riparian

Karr& Schlosser's (128) demonstration theland-water that interface reduces movements streams tounderstanding roleplayed riparian to led the nutrient by zonesincontrolling in nonpoint sources pollution sediment nutrients of by and watersheds agricultural (114, 147,208). Important biogeochemical processes affect that streamside wellas aquatic as the occurwithin riparian on systems zone, whichis influenced one handby watershed and hydrology on theother handbychannel hydraulics (264,265). The subsurface transfer water materials mostly of is unidirectional toward and thechannel acrosstheterrestrial In it boundary. contrast,is bidirectional across the aquaticboundary, whereoxidizedhyporheic waterfrom streambed the mixeswith(often the reduced)interstitial water comingfrom riparian zone. Bothboundaries for appearto be majorlocations regulating diminishing and thetransfer inorganic of from and subsurface waterto nitrogen phosphorus stream water (159). in carried surPHYSICAL BUFFERS Sediments sediment-bound and pollutants face runoff deposited are in forests well as in as effectively mature riparian flow which streamside is grasses.Sediment trapping facilitated sheet by runoff, allowsdeposition sediment erosion acof of channelized particles prevents and sediments. 80-90% ofthesediments cumulated areasremove Riparian leaving in fields North agricultural Carolina (42,47). Sediment deposition be submay in stantial thelongterm, with within fewmeters a coarsesediments deposited into ofthefield-forest sediments further theforand boundary, finer deposited in est and nearthestream wherethey mix withcoarse sediments deposited overbank flows (42,146).

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and concentrationslabilenutrients adsorbed of Finersediments carry higher of from runoff the occursas a consequence theinterpollutants; removal their dilution, adsorpand activeprocessesof deposition erosion, and infiltration, In watersheds with with soil tion/desorption reactions forest andlitter. forested riparian zones can be sourcesor sinks relatively nutrient low concentrations, conditions. example, For the on fornutrients, depending oxidation-reduction Tennessee a net was forest stream eastern in riparian zoneofa smalldeciduous concentrations riin whendissolvedoxygen sourceof inorganic phosphorus were when dissolved concentrations oxygen parian groundwater low,buta sink werehigh(174). zones Although riparian may complex. Phosphorus dynamics be particularly particulate phosphoas traps (sinks)for incoming mayacteffectively physical (60). Sigwaters availablesolublephosphorus in rus,they mayenrich runoff in zones,then of mayfirst accumulate riparian nificant amounts phosphorus form in be transported aquaticecosystems a different via shallowgroundto of matter of decomposition organic water flow, possibly a result increased as (71,268). when from morethan 50% ofsediments uplands Grassy riparian areastrap the water flowsare <5 cm deep (60,154). Grassyareas influence overland of channelized flows expanded into shallow uniformityrunoff transforming by whichare morelikely deposit the to sediment. However, performance flows, when of grassy seemsto be highly variable and of short duration vegetation occurwithin limited a sediment trapping period.For example, several floods to rainfall simulation event 5% in from 90% in a first efficiency decrease may a sixth rainfall simulation event (60). an important mechanism nutrient for BIOLOGICAL BUFFERS Plantuptake, in accumulain forests (43,72,88,208), results a short-term removal riparian in tionofnutrients nonwoody in biomass a long-term and accumulation woody sites forests especially are biomass. Riparian important forbioticaccumulathe increasing mass tions nutrients of becausetranspiration be quitehigh, may and rootsystems, becausemorphological flowof nutrient solutestoward and nutrient flood-tolerant of speciesfacilitate physiological adaptations themany In under conditions. some species,suchas water tupelo, uptake low-oxygen & conditions enhancenutrient and growth (103). Peterjohn saturated uptake N of Correll vegetation uptakes 77 and 10 kgha-l yr-lfor and (208) estimated with rates.However, potential are P,respectively, that comparable upland rates N uptake ratesmaybe muchhigher, shown Cole (37): Poplar(Populus as by with efflufertilized a nutrient-rich 213 nigra)assimilated kgN ha-' yr-lwhen nutrient entata rate 400 kgN ha-l yr-lfor of three but receiving years, sitesnot due saturation effluent assimilated only16 kgN ha-l yr-1.Further, tonitrogen

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factor tree for growth, particularly in (2), phosphorus becomethelimiting may phosphorus sink. wetlands (260), making vegetation effective an filters maybe reduced restricted by The importance plantsas nutrient of to of and accessibility water, theseasonalphenology uptake releaseofnutriby Water accessible plants is to only ents, bythesaturation mature and of forests. demand moveswater and solutes if thewater tableis highor iftranspiration concentrated surface flowand intotherootzone. During intense storms, rain to (1 macropore-dominated percolation not available plants 15). Nutrient may be declines stops precisely when high discharges occur. uptake or during winter, the In addition, to soilsstored during litter decomposition releasesnutrients forest thegrowing nutrientswoody in biomass season.Finally, ability sequester the to to (2, of maydeclineas trees mature, leading saturation 88). The contribution retention remains be elucidated. to individual riparian speciesto nutrient tree in results the of similar plant to uptake, initially Microbial uptake nutrients, by death, decomimmobilization dissolved of nutrients followed cell growth, in to and nutrient release.Nitrogen, contrast other nutrients, position, eventual in riparian forests. Denitrifihasanalternate pathway major of importance most on of a cation (43, 88, 114,125,212,214) depends thepresence nitrate,suitable moisture, and carbon substrate, theabsenceof oxygen.Soil temperature, and rate.Soil pH affects whether or N20 thetype carbon of influence reaction the with decomIn microsites associated N2is produced. riparian zones,anaerobic inotherwise well-drained posing organic matter fragments denitrification allow rates soils. Within riparian a zone,denitrification of30-40 kg ha-l yr-' have where been recorded; fastest the boundary ratesoccurat theriparian-stream soil nitrate-enriched enters water organic surface (40). Denitrification groundwater-borne is less well established.Carof nitrate anaerbonavailability limits microbial subsurface activity, preventing usually is concentrated in obic conditions from developing (99). Sincedenitrification theupper12-15 cm of thesoil (whichis onlyoccasionally partof theshalthe low aquifer), from shallow groundwater require may nitrate disappearance nitrate to role riparian vegetation playa primary (144). Groundwater-derived of in soil plantuptake thenitrate maybe eventually denitrified surface after from litter and release,followed by groundwater, decomposition, ammonium with denitrification interacts microbial nitrification (88,92,93). Therefore, via and carbon availability litterfall root vegetation nitrogen uptake organic and rivarieswithin between and nitrate.Such an interaction decay to remove undertheinfluence subsurface of water, plantcover,and soil parianforests characteristics (101). on flow effects nuBUFFER VARIABILITY Subsurface water paths havestrong In trient locations the of CoastalPlainoftheChesapeake characteristics.several

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boundary nitrate concentrations7 of Bay watershed, average annual terrestrial groundto 14 mgN03-N L-l decreaseto 1 mgN03-N L-1 orless in shallow site a water near streams (145). However, thesamearea,a single with nitrate in of concentration 25 mg N03-N L-1 at depthhad a concentration 18 mg of Lowrance al (148), who et N03-N L-l in shallowgroundwater thestream. at 31 estimated annualdenitrification to average kg N ha-l yr-1in thetop rates 50 cm of soil,measured rates 1.4 denitrification between kgN ha-l yr-1in a received fertilizer) 295 kgN no to riparian zoneadjacent an old field to (which and subsidies. Suchresults ha-l yr-lunder conditions high of nitrogen carbon insurface soilsas wellas thehigh variillustrate potential denitrification the for to in is measurements. ofthisvariability driven Most ability be expected field and soil as by fine-scale differences between rooted nonrooted layers well as on flow between anoxicand oxic conditions, itdepends subsurface paths. and On thewhole,enrichment riparian zones maylead to significantly higher of litter and soil inorganic-nitrogen contents, potentials concentrations, nitrogen saturation. for nitrogen net and all nitrogen mineralization nitrification;suggest enriched of and usually maintain Nevertheless, rates denitrification storage high nitrate (92). riparian zonesas sinks upland-derived for nutrithe of coveralso influences efficiency riparian zonesin filtering Plant with poplaris moreeffective entsand pesticides.A riparian zone vegetated are forwinter retention one vegetated grass(97). Some trees with nitrate than better others filtering in removes x effectively than nitrate: Populus canadensis accumulation nitrogen root of in nitrate from saturated with subsequent soils a seemtofavor colonization by biomass (203). Rootsofalder, willow, poplar and for and and, proteolytic ammonifying microorganisms particularly alderroots, plantcovermayaffect to inhibit nitrifying microorganisms (221). Changing zone in New Zealand 12 yrafter retirewater quality:In a set-aside riparian returned native to tussock (Poa cita), ment from grazing, dominant vegetation and to for nutrients dissolved leading a zonelikely be a sink sediment-bound to but dissolved (41). nitrogen a sourcefor phosphorus of redoxconditions theavailability disand Soil characteristics influence that to solvednitrogen plant roots.Pinayet al (215) calculated sandy riparian but flux a forest soilsretained 32% ofthetotalorganic nitrogen during flood, on soils. McDowell et al (162) compared 70% was retained loamyriparian rain different in tropical riparian nitrogen dynamics twogeomorphologically water of subsurface forest sites.Atonesite, deeplayer coarsesandconducted a to theadjacent oxic stream belowmostplant roots, through and anoxiczones from at whereas theother site,a dense upslopeto downslope, respectively, movedthrough a and water rapidly claylayer impeded infiltration, subsurface intense bioticactivity shallowand variably zone. Although oxidizedrooting soil controlled of at sites, differences strongly hydrologic export nitrogen both

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have consequences waat modified interactions. Such interactions important tershed landscape the retention ammonium of or and scaleswhenever transient nitrogen cycling, thereby altering nitrate riparian in sediments influences biotic thetiming form dissolved and of nitrogen export (266). inorganic scaleswith respect natural to dishowplant speciesrichness varies regional at are effective evaluatin turbances (218). Profiles alongriver courses especially for biodiversity patterns ingtheintermediate disturbance hypothesis explaining in sections river of (199). Plantspeciesrichness notalwayshighest median is courseswhereflooddisturbances at intermediate levelsof intensity and are duration (8,57,199,259). in In northern of results species Sweden, post-glacial history thelandscape into durwhere rivers deposited diversity peaking begintodowncut sediments rivers Spain,semi-arid inga higher coastal stage (9200yrBP; 198). In southern as showirregular of richness alongtheir courses, a consequence patterns species of irregular and tributaries may water availability (259). Main channels their differ in theAdourbasinofFrance, where exoticspeciesaremorenumeras than ous in themainchannel display different longitudinal patterns native and Sweden,themainchannel species (57,216). In theVindelbasin,northern in speciesrichalso has a higher speciesrichness thetributaries; addition, than but ness is greatest mid-altitude themainchannel leastin thetributaries at in (58,217,257,258,259),or (195). The distributions native exoticflora of and in markedly longitudinal long-lived ruderal species(195, 196,199,200) differ profiles richness. of It remains be demonstrated biodiverriparian forests maintain to whether from other catchments continuous movement species of sity through upstream envior from extension speciesdistributions of downstream favorable during has dispersal water a role in by ronmental periods (195). Clearly, propagule such the mechanisms as anemoflora structuring riparian (116), butdispersion and deserve consideration (169), as do thedifferent stages chory ornithochory for establishment and of a plant'slifecycle,whichare important successful growth (56). river banksin North from Pacific Black cottonwood (Populustrichocarpa) as to climatic selection at regional well as local Americarespond pressures inhigh in and variations survival, growth, photosynthesis scales,with significant is to the crown traits respond whether habitat light regimes; additionally, and leaf mesicorxeric discontinuities among riparian populations (65,66,67). Genetic of P trichocarpa coincidewithupstream-downstream along thesame river occur moisture levels(66,67). Such discontinuities changesin atmospheric
VEGETATIVE PATTERNS DIVERSITY Riparian forestsprovide insightsinto OF

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even though black cottonwood dispersesexual and asexualpropagules can overwideareas. zone Floodscreate heterogeneity the within riparian andthereby niches facilitate coexistence the of create distinct regeneration that despite similar congeneric species.Forexample, speciesofSalix co-occur, six adultecology, pealongtheSorachiRiver, Hokkaido, Japan (194). Dispersal spring floods. riodsofthese speciesoverlap water as levelsdecrease following The dominant species,S. sacchalinensis, establishes a widerangeof soil on coexistwith textures.The subordinate speciesS. roridaand S. subfragilis S. sacchalinensis thefinest coarsest where domthe on and soils,respectively, The of other inant speciesdoes notgrowas efficiently. occurrence thethree stages flooding to rarer subordinate speciesalso is related thefitness early to of and soil characteristics. for Periodic flooddisturbances various of intensities also critical mainare forest the tree podocarp taining four dominant speciesofthelowland floodplain in New Zealand (63)-Dacrycarpus dacrydioides, cupressinum, Dacrydium Prumnopitys ferruginea (conifers), Weinmannia and racemosa(angiosperm). trees. Thetwoformer upper-canopy andthetwolatter sub-canopy are are trees, on to Intense floods denude largeareas,allowing dacrydioides establish silt D. with on microsites covered substrates D. cupressinum establish elevated and to suitable sites organic debris. Less intense floods exposedebris, providing more forD. cupressinum occasionally P ferruginea. for do Minorfloods not and the of disturb canopy provide but opportunities establishmentP ferruginea for in under canopy cover andW racemosa smallcanopy gapscausedbynonflood of conifers. mortality themature At local scales,floods of plants through affect speciesdiversity herbaceous and of and physical heterogeneities created theerosion deposition litter silt. by Innorthern of 150 Sweden, litter leaf accumulations approximately g m-2result ina maximum speciesattheupper number diversity adult and of vascular plant in limit riparian elevational of zones(197). Likewise, southern France, organic in rates matter accumulations 150 to 300 g m-2 result highgermination of of and decreased whereas accumulations 600 to 1200 g m-2 mortality rates, to decrease 150 rates. corresponds maximal germination Around g m-2oflitter of and silt density speciesrichness (137). Similarly, accumulations about500 of to and (138). g m-2correspond maximal density speciesrichness seedlings
NATURALDISTURBANCES INVASION BY EXOTIC SPECIES The richest communities havethegreatest also of as sites. This specific proportion exotics, alongtherivers well as within in corridors thattherichest communities riparian maybe themost suggests

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heterogeneity created by invasible because of thesubstantial environmental plantcommunities theAdour of moderate floods. Comparing riparian the Oregon, Planty-Tabacchi al (217) et River, France, with Mackenzie the River, species, discovered about1400(24% exotic)and850 (30% exotic)constituent alongtheAdour (46%) than along respectively. Morewoody speciesareexotic theMackenzie communities theMackenzie of (17%). In contrast, herbaceous these are moreinvasible (32%) thanthoseof theAdour(21%). Although in differences management Oregon in than France, parallel more intensive forest theecologicalmechanisms similarities remain obscure. explaining these of Several interactive to establishmentexotic species processes appear control in biological invasions have inriparian zones(56). Although common nature, beenaccelerated (142). Life-history characteristics through human activities and of invaders control various the stabilization, exstagesof establishment, the the that shorter life pansion.For example, Pysek& Prach(222) reported thehigher rate invasion the of four speciesalientocentral Europe, span, among and Reynoutriajaponica, Impatiens glandulifera, Heracleum mantegazzianum, R. sacchalinensis. such along Landscapecharacteristics as connectivity rivers patch structure (223) also exert (116) andhistorical development landscape of control. Naturalenvironmental features may also slow therateof invasion. For in exotic from Sweden, plants example, thesubarctic environment northern of other in where area the Australia, continents rare riparian are zones. In northern 1.2 the every yr, rate expansion of occupied theexotic by Mimosanigra doubles wet is related theamount rainfall theprevious season (143). At the to of in due regional scale, thearealdoubling timeis 6.7 yr,probably to thespatial savanna.Seed predation combined isolation wetland of habitats eucalyptus by becausetheseedsof with folivores also likely slowtherateofexpansion is to thisspeciesaredispersed flotation. by there exotic can native plant species diversity, is no Although invasions reduce clearevidence does. However, it the annual glandulifera, tallest plant Impatiens in Europe,is expected reducespeciesdiversity to out-compete native to and Tamarix habitats (10,216,224). Similarly, light-demanding speciesinriparian of habitats exoticwoodyplantsin aridand semi-arid riparian spp.,invasive muchof thenative to western UnitedStates,are expected replaceor inhibit do these cleardatasupporting refuting expectations not or flora (25). However, appeartoexist.
REFUGES FOR REGIONAL DIVERSITY

as forest haveacted safe patches Riparian humid sitesforregional flora tropical during periods.Some present-day dry zonesappear haveexperienced Pleistocene but is to droughts, there noindication is of ofmassextinctions, there indication rapidspeciesre-expansion whereas

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for mayhavebeenrefuges patches forest Holocene.Riparian the during early (166). Thefloristic America in diversity Central of maintenance mesicplant the continuous to are forest similar thosecharacterizing of attributes theriparian thus and is stemdensity higher biomasslower, in forests thearea,whiletheir trees riparian Moreover, speciesrichness. to their increasing potential maintain are trees removed becausemoremature younger are matrix often in a savanna removal (132). Frequent forests to as by windand fire compared continuous the from community exclusion the reduces rateofcompetitive trees ofmature species. of numbers coexisting for the whileenhancing potential greater > with 10-cm (trees 52 America average species/ha in forests Central Riparian for lowerthan to comparable or slightly a height), richness at diameter breast occurin diverse forests extremely in forests thearea (166). However, upland suchas in Amazonia(225 treespeciesha-l) and in rain other tropical forests forests role the speciesha-) where refuge ofriparian Asia (283 tree southeast (62). mayhavebeenmorelimited (45). vegetation of a from knowledge riparian may characteristics be predicted facimportant most trees of or Thepresence absence riparian couldbethesingle (255) communities sinceriparian these that activity affects by tor altered human on its through effects primarily diversity macroinvertebrate affects vegetation vegeof the (21, habitat 82, 140,204,227). In contrast, effect riparian benthic is for as tation a sourceof nourishment invertebratesnotso well established. consumer a thus is Muchofthewoodydebris unpalatable, preventingdiverse (202). Also, changesin species composition, from developing community For excomposition. followchangesin riparian often thandiversity, rather a may of community chironomids replace shredder a ample, gatherer-collector in litter riparian decomposing in is whenthere a reduction slowly community (89). streams smallwoodland
community COMMUNITIES Stream macroinvertebrate MACROINVERTEBRATE

Habitat

changesin and and in Withvariations floodduration frequency, concomitant of is the succession, environmenta complex shiftand tabledepth plant water scales (155). spatio-temporal on and destroyed different created inghabitats of vegetation with remarkable a variety woody zonesarecovered Mostriparian and nesting to offering for as shrubs serving refuges smallmammals trees from as of develops a result enherbivory sitesforbirds.Also, sustained perching stability debris provides woody and and hanced productivity foodquality, fallen act forests communities. Riparian as for terrestrial wellas aquaticinvertebrate and for in areasand,in somecases,as corridors migration as refuges adjacent (24). dispersal

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The occurrence anyspeciesinriparian of areasis probably toseveral due interrelated reasons. Juvenile woodturtles (Clemmys insculpta) prefer remain to nearstream channels wherethey can movecomparatively distances short to find appropriate thermal moisture and conditions atthesametime less and are exposed predation to (22). Drinking water obviously important is an reason for mammals visit to riparian zones. However, smallmammals respond may more in to differences tree communities. Higher capture rates somesmallmamfor malshavebeenreported streamside in habitat dominated redalder(Alnus by rubra)adjacent uplands to dominated Douglasfir by (Pseudotsuga menziesii) (160), butno difference detected was whereuplandand streamside habitats in weresimilar vegetative structure composition and (161). The white-tailed deer (Odocoileusvirginianus) uses riparian zones almosttwiceas muchas nonriparian areas,supposedly an antipredation as strategy (139). in Like mammals, moreindividuals speciesofbirds found riparian and are morethan60 habitats thanin adjacentones. In thelowerMississippi River, and and speciesofmammals, about190 speciesofreptiles amphibians, about 100 speciesofbirds seasonally are associated with riparian habitats (133). For Coloradooccurin riparian example,82% of thebreeding birdsof northern vegetation (135). Bird assemblages riparian of zones and adjacentuplandsare interdepenof and dent;thenumber shared speciesvariesseasonally longitudinally along river courses with apparently greater interdependence atintermediate elevations (135, 141). However, individuals speciesmaynotbe morenumerous bird and dewhere riparian the habitat either or is similar upland to habitat notclearly lineated Riverin (176). About90% ofthepresent faunaalongthePlatte bird northern Coloradohas arrived sincethedevelopment a gallery of forest within thelast90 years bird increase 30% (134). In boreal forests, densities reportedly to 70% inprotected riparian forest strips yearafter the clear-cutting; then they decline the while adjacent the clear-cut during following years regenerates (48). Birdcommunities sensitive thequality riparian to are of vegetation 141). (44, Destruction riparian the of vegetation causeslocal extinction also reduces and River of to sites(135). AlongtheColorado ability somepopulations recolonize in theGrandCanyon, black-chinned hummingbirds (Archilochus alexandri) nestonlyin exotictamarisk-dominated than0.5 ha habitats are greater that in area (27). In disturbed 2 areas,woodystrips m widepermit onlyportions to of birdpopulations occur; widths to >25 m on each bankare necessary from m 25 maintain sensitive species(44). In a survey 117corridors of ranging to 800 m wide,Kelleret al (130) concluded theprobability-of-occurrence that increased mostrapidly between m and 100 m. An important 25 conclusion ofrecent work riparian on must habitat birds that for is conservation be based on specific birdspeciesand account differences behavior example, in for (for

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between generalist-opportunisticriparian-obligate and species). A drainage basinperspective absolutely is necessary (135). The inhabitants riparian of zonescan modify habitat structure function. and Through ponding water storing and sediments, beaver create wetlands alter and thevegetative composition in-channel riparian of and communities temperin ate North America, strongly influencing riparian landscapes(186,188,189). Through selective cutting trees, of theychangethecomposition riparian of communities. examplein Minnesota, For under beaverinfluence, trembling in aspen(Populus tremuloides) decreased abundance, whereas alder(Alnus rugosa) andblackspruce (Picea glauca) increased (122). Besidesbeaver, species suchas moose,elk (Cervuscanadensis), brown and bear(Ursusarctos)create and maintain of banks. In southern tight networks trails alongriver Africa, hippopotamus (Hippopotamus amphibius) modify riparian habitats, gathering in poolsduring daywhere the they up sediments deepenaquatichabistir and the tats;during night they createpathsbetween pools and terrestrial grazing between areas,thus maintaining connectivity patches (190). Warthog (Phacochoerus transform ofhectares ploughed tens to fields digging aethiopicus) by in soils 10-15 cm deep to feedon underground storage plant organs riparian the forests wetlands and of rhizoma(229); consequently replacement perennial tousgrasses annual is by grasses forbs favored. and Feedingactivities havelong-term consequences thestructure funcfor and tionof riparian the forests.Zoochory, particularly ornithochory, facilitates selective expansion certain of speciesthrough feeding propagule and transport. in Moose browsing reportedly affects decomposition indirectly through changes thequality litterfall North in of American riparian systems (163,205). Similar effects occurinriparian in corridors Africa through browsing elephant bulk by (Loxodonta africana),bulkgrazing hippopotamus, selective by and browsand ingbykudu(Tragelaphus strepsicerous), giraffe (Giraffa camelopardalis), bushbuck (Tragelaphus scriptus) (190).

ENVIRONMENTALALTERATIONS Human Alterations

include neolithic deforestaalterations human-induced AlongEuropean rivers, and tionandland-clearing Gallo-Roman medieval during periods (207). Civil in in and works thenineteenth engineering century hydroelectric developments events have thetwentieth thesealterations accelerated century (211). Similar in timescale sinceEuropean settlement occurred North on America a reduced (246). in which necessary for are Flow variability fluctuations channel and width, the of maintaining biodiversity riparian systems (78), havebeendramatically

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in through impoundment, manriver water decreased many parts theworld of in changes riparian vegeagement, lowering water and of tables. Substantial flow, riparian as vegetation is tation mayoccurwithout changing meanannual and flows (6). In many especially sensitive changes minimum maximum to in in plant community comcases,hydrologic alterations result shifts riparian in communities (18,31,54,77, 104,196, position wellas senescence woody as of 198,207,232). in UnitedStatesby The rapid invasionof floodplains the southwestern water courses (269), inducing other disTamarix ramossissima desiccated has of species(e.g. Salixgoodturbances as fire such (32) anddisplacement native floodplain, dingii Populusfremontii; 269). AlongtheColoradoRiver and 253, for maybe at theoriecophysiological tolerances competition moisture and in structure a gallery from forest riparian to ginof shifts riparian community of Salix) andthedisappearance thickets (with Tamarix dominating replacing or Populus(32). historiin floodfrequency duration and AlongtheRiverMurray Australia, from native gum(Eucalyptus red camaldulensis) establishing callyprevented of river on grassplains. By reducing and regulation depth duration flooding, extinction has favored redguminvasion is expected cause a complete a to that of (19). Alterations water oftheonce-extensive plainsin thenearfuture grass declinesand losses of cottonwood regimes also havecontributed dramatic to as of forests throughout western North America a consequence drought-induced from of tables(233,234). and mortality abrupt flow reductions lowering water A possible is a of diminishes mechanism this for decline that reduction flooding therateofrecruitmentnewstands of (170). In fact, maintenance phreatophytic the woodlands heavriparian depends of traits between river flow life-history ofcottonwood and ilyon therelationship within between and as vary years (108). Forexample, growth conditions widely at thesame site,an important of to factor survival theability seedlings for is overa rangeofmoisture et establish conditions (247). Tyree al (267) demonto for strated regimes several Populusspecies species-specific responses water differ xylem in cavitation. that water and potentials consequent in of to Even though alterations water regimes mayresult declines cottonenvironmental wood forests, leasttemporary at increases occurin someother comMissouri situations River, poplar-willow (249). Alongthemeandering erodeouter banksand deposit sediments munities dependon flow peaksthat on inner banks(118, 119,120). A reduction peak flowsafter of completion of oftheGarrison and the Dam in 1953prevented meandering thus formation in desuitable areas forpoplar-willow establishment, resulting a continuous in creasein pioneerstagesand an increase olderstages(119). In contrast, River communities dramatically have expanded alongthePlatte poplar-willow

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after completion damson itstwomain of tributaries, North South the and Platte rivers, reduced flow exposed and large areassuitable seedling for establishment andsurvival (120). Invading riparian trees mayhavean impact channel on morphology, particularly whenthey replaceformerly grassy areas(79,237). Active investigations are neededon mechanisms linking water, landforms, speciesin various and landscape settings order predict specific in to the effects manipulating of flow on floodplain forests (53). In northern Sweden,species-richness thepercentage and vegetative cover are bothlowerper site in a regulated compared a nonregulated as to river (196,198). The proportion annualplus biennial of species-richness higher is and perennial species-richness loweralongregulated is rivers. Waterlevel regime and mean annualdischarge certainly amongthemostimportant are variables maintaining for speciesrichness plant and cover. The speciescomposition theriparian of is plant community also important to consider predicting in responses alterations 196,198,207). Several to (18, scenarios be expected a plant may for as alterations: community a result flow of reproduction on-site by regeneration, colonization other of parts thefloodof plain,or replacement a new typeof plantcommunity by (207). Alterations ofriparian communities plant affect and obviously aquaticmacroinvertebrates fishes a consequence modifying as of and habitat trophic pathways in-stream of (49,238) as well as thespeciesdiversity amphibians, and birds, mammals (44,130,181).

and Management Restoration

are Hydrological characteristics of primary in importance managing riparian zones. Riparian buffer zonesretain surface runoff as of pollutants a result their water storage and capacity infiltration Riparian (41). buffer zonesintercept the dominant hydrological that on pathways are dependent soil typeand permeability, adjacent land use, slope,potential run-off generation areas,and land installations. example, For nitrate removal that drainage requires thebulkof thewater moveseither acrossthesurface as shallowgroundwater or through active zones(145), andsediment soil removal that biologically requires surface runoff notoverwhelm buffer are does the system. Hydrological pathways likely to changewidely space andtime.Forest in variations or growth weather may affect degreeof saturation theriparian the of zone and theproximity the of water tableto thesoil surface the of (70,98), thereby complicating intricacy routes thewayswater-borne and nitrate and groundwater encounters roots soil microbes. rates zonesdifferentially to Increasing loading mayaffect riparian according thetype pollution 145). Fornitrate, of (70, rates N-removal of higher generally

646

NAIMAN& DECAMPS

of and loading ratesas a consequence denitrification vegeoccurwith higher maybe limited inefficient by buffering capacity tative uptake.Nevertheless, carconditions, organic or limited duration anaerobic of nitrate uptake rates, nutrients metals, and respiration (98). Forother bonavailability microbial for are and ratesof similar denitrification lacking, higher to biological processes of capacity the loading mayresult excessreleasewhentheimmobilization in to In riparian buffer exhausted. suchcases it is necessary manageriparian is these so sediment removal infiltration as to prevent and systems facilitate to and of Also, flooding erosion from becoming sourcesof pollutants. systems to limitation buffering capacities winter be a general may riparian soilsduring for nutrients metals and (98). use zones is to control An increasingly important managerial of riparian over when diffuse Riparian zonesare moreeffective thelongterm pollution. at practices the has through good agricultural upstream pollution beenlimited of zonesonwater effects riparian quality catchment (7,51). Theintegrated level smaller streams havinga greater will also differ according stream to order, than diffuse onestobuffer pollution (145). Thecontrol against potential larger in is strategy management ofwater catchments an effective quality headwater remain for is few oncea river contaminated, inexpensive possibilities because, improvement (98). buffer the One modelsuggests multi-species that strips provide best riparian of (106, 145,245). Thismodel against agricultural impact protection streams the uses three interactive up-slopeorderfrom zones thatare in consecutive and treesup to forest about 10 m wide,2. shrubs stream:1. a permanent 4 m wide (and managedso thatbiomassproduction maximized), 3. and is the zone influences stream up herbaceous vegetation to 7 m wide. The first channel environment temperature, habitat diversity, morphology, light, (e.g. controls foodwebs,andspeciesrichness). The secondzone,which pollutants and transin subsurface andsurface flow is biological chemical runoff, where and sediments in infiltration, deposited formations, storage woodyvegetation, and to are maximized.The first zones contribute nitrogen, two phosphorus, of zone provides sediment removal.The third spreading overland pollution this of Clearly basicmodel thus deposition coarsesediments. flow, facilitating to and orders provide must adapted various be to catchment conditions stream effective management. imitate is to systems sustainability likely occurwhenmanaged Long-term buffer strip riparian natural ones. For example,zone 1 of themultispecies better zone 2 is harvested if infrequently; functions (locatednearthestream) if better accumulated sediment and zone 3, nearthecropland, functions also is removed herbaceous and is periodically (145). The vegetation reestablished from efficient removal ranging offers divergent examples literature, however,

INTERFACES RIPARIAN

647

sinks (41). (7) nutrient loading toexhaustible of ofnitrogen 20 years high after within catchment the and Whatever example, the land improved use practices flowsby of zones forinterception groundwater of themaintenance riparian capacity) differ absorption in (which stages succession of vegetation various in (98, strips streams 270). and vitality buffer of for arekeyfactors thelong-term of and creative management restorationriparian obtained from Other benefits wildlife, of habitat terrestrial aquatic for and zonesinclude provision diversified matter to and of animal dispersion, input organic and corridors plant perhaps for value in and of streams. Enhancement thevisualquality increase recreational Management beenusedposihas benefits. ofthelandscape also important are in throughout animals upland streams of to communities aquatic tively influence are and forests recWalesandScotland floodplain (204), andmanaged natural restoration projects (209). ommended British in river floodplain and

Tools theFuture for

related to functions ecological are expected fulfill Riparian systems increasingly microclimate, cycles, tobiodiversity, information biogeochemical flow, habitat, also andresistance resilience disturbance to (187). Theywillbe expected and of recreation, to fulfill moresocial functions, including provision resources, all will system perform these no riparian Clearly, single culture, aesthetics. and at to but functions, eachwillbe likely perform leastone. at courses, An extensive linear to approach management, thescale ofriver Significant alongstreams. riparian systems and is neededto delineate classify riparian systems usingremotely beenmade in mapping progress already has be that et senseddata (175). Lowrance al (145) recommended linearforests with at of characterized a resolution 10 m to 20 m. In combination hydrogethe for data,suchprecisemapshelpin assessing potential riparian omorphic Managementto and pollutants. surface- subsurface-borne systems intercept on initiated investigations, as thoseconducted thepoplarsof thePlatte such to entire river courses understand demonstrate needfor the considering River, the (120),torestore ecosysof systems of various aspects thedynamics riparian international greentemintegrity rivers floodplains of and (46), andtomanage ways(28). are to Bothlandscape detailed perspectives required judge whether site and perspecsizes. The landscape trees willsurvive reachtheexpected to planted tivemaybe attained based on mapping pastand present knowledge through of characterizing ecologyof approthe extents floodplain plant communities, at and landscape sitescales(105). priorities both and priate species, determining based on water Detailedsiteperspective be attained through knowledge may ratesas survival growth and suitable conditions, long-term and soil regimes, metabolism (117). wellas on theeffects variable water levelson tree of

648

NAIMAN & DECAMPS

research, demonstration, and An intensive site-specific approach integrates will application riparian of zone buffers (145). Such an integration aid in approach the on discovering effects vegetation the of typeand management pollution, response riparian the of long-term control nutrient sediment of and in zones to acutestresses suchas largestorms extremes temperature and or of stress leading saturato growing seasonrainfall, consequences chronic the controlling groundwater tionofriparian zonesbynutrients, theprocesses and issuesarerequisite develfor microbial dynamics 145). Insights these (87, into (106). Intensive opingmodelsof riskassessment (149) and decision-making studies the to funcsite-specific also shouldimprove ability modelimportant on tional issuessuchas theinfluence soilwetness keynutrient of transformation should improve ability evalthe to processes (83). Finally, site-specific studies a of habitats uatetheperformances restored throughbetter knowledge riparian ofhydrologic, and geomorphic, biologicconditions. Researchat thecatchment hillslope scale, or at leastat therepresentative on to of systems hydrologic inputs scale,is essential assesstheeffect riparian in most operate effectivelyheadfrom uplands (145). As many buffer processes water involving many smallcatchments basins, downstream cumulative effects is scale must given be proper emphasis (29). Thecatchmentalsotheappropriate zones(243) andtoassessthe to improve conditions within riparian hydrologic of are of wider ones potential narrow riparian zonesthat remnants previously in mostrural urban (54,210). and temperate landscapes zone retoolsforthefuture Finally, considering riparian zones as management the of schemes cope with to related surprises quires adoption flexible, adaptive in to discontinuities synergisms and systems (177). Discontinuities riparian nutrient reachdisruptive thresholds sudthat mayoccurwhere accumulations also from sinkto a source. Synergism mayoccur a denlychangethesystem from (such as nutrient loadingand interactions between chronic two stresses (suchas a large storm). globalwarming) between chronic anacutestress or a and future issues In addition such"anticipatable unforseeable entirely to surprises," (187). require increased efforts research management in and
ACKNOWLEDGMENTS

from AWMellonFounthe of was by Preparation thisarticle supported grants Station theUS ForestService. of dationand thePacific Northwest Research and for T E We thank Nilsson, Coe, G Pinay, Chauvet, E Tabacchi comments C and and suggestions greatly that the improved content, TrevaCoe and James Helfield assistance for with bibliographic the searches.
homepage at VisittheAnnualReviews http://www.annurev.org.

RIPARIANINTERFACES
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