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Donald B. Zobel, Departmentof Botanyand PlantPathology, State University, Corvallis, Oregon 97331- thanmost others in Oregon, 2902 [email protected] 1983).The tenitorycontrolle villages extendedsouth se' the currentlocation ofLincr the crestof theCoastRange ( EcosystemUse by IndigenousPeople in an OregonCoastal Landscape The SalmonRiver is smt The SalmonRiver has306 tat, including 223 ha salt n ( Abstract and32 ha submergedland 1974).Thereis evidencein of the Data regardingprobable uses of biological materialsby the indigenouspeople of the SalmonRiver-Cascade Head area quake-inducedsubsidence 256 were Oregon'co"rt-*ere used to estimatethe people'suse of variousecosystems near villages.Of 308 uses identified' for the tion, which coveredlndian h attri-butableto a given species;124 specieswere identifred.All local ecosystemswere important sourcesof organisms one (Minor andGrant 199 people,with no app"rent concentrationol highly used speciesin any particularecosystem. One specieswas cultivated, 1700 from the domesticated,and five acquired by trade. Fouimajor plant resources,camas, yew, hazel, and betngrass,are not known ing most of the estuarymou without use of CascadeHead landscape,but may have been uuuilubi. from elsewherein village territory.House construction est and grassland,althougl in the SalmonRiver cedarplanks, as in

304 NorthwestScience, Vol. 76, No. 4' 2002

O 2002 by the Northwcst ScientificAssocialion All rights reserved ersity,Corvallis, Oregon 97331- thanmost others in Oregon(Murray andMarrant fish (Oregon Departmentof Fish and Wildlife 1983).The tenitory controlledby the SalmonRiver 1997).In 1923-1921,before a majordecline, com- villages extendedsouth severalkilometers into mercial catchof salmonand steelheadfrom the thecurrent location of Lincoln City andinland to SalmonRiver was much lessthan that in larger thecrest (Beckham CoastalLandscape ofthe CoastRange et al. 1982). systems,lVa that in Tillamook Bay, 2Voof the The SalmonRiver is small,draining 2 l0 km2. Nehalem,and 4Vo of the Nestuccaand es- The SalmonRiver has306 ha of estuarinehabi- tuaries(Cleaver I 951). tat, including 223 ha salt marsh,5l ha tideland, Extensivered alder(Alnus rubra) and conifer and32 ha submergedland (Thompsonand Snow forestsdominate slopes above the SalmonRiver 1974).There is evidencein the estuaryof earth- valley. on River-CascadeHead area of the The conifers are primarily Sitka spruce . Of 308 usesidentified, 256 were quake-inducedsubsidence and tsunamideposi- (Piceasitcherzsls) near the shoreline,with west- ortant sourcesof organisms for the tion, which coveredIndian hearthareas aboutA.D. ern hemfock(Tsuga heterophyllct) becoming im- n. One specieswas cultivated,one 1700(Minor andGrant 1996).A sandspit block- portant a short distanceinland. Severalkilome- I beargrass,are not known from the ing mostof the estuarymouth supports dune for- ters farther inland, Douglas-fir (Pseudotsuga Houseconstruction without useof est and grassland, :stemredcedar in theSalmon Rir er althoughthere was probably menziesii)becomes abundant and often dominant. tern redcedar,may have limited the lessstable vegetation there before the introduc- and abundanceof sprucedeclines. This fbrest tionof Europeanbeachgrass (Ammophila arenaria) patternis not an artifactof logging; it existedin (Wiedemann1984). A 69 ha grasslanddominares theuncut forests near the time of Euro-American the southwesternpromontory ided nearCascade Head on of CascadeHead, settlement,according to timbercruise records for justuphill from majorvillage egonCoast, in an areawhere sites.Cascade Head Tillamook County (Anonymous 1908).Most of alsosupports smaller grasslands above istoricand modern plant and theocean the forestedarea burned in the 1840s(Munger cliffs farther north. Little butionis unusuallycomplete. rocky shore can be I 944),although areas of old-growthand isolated reachedon foot, but much d wascompared to historic is accessibleby water. old trees survived (Greene 1982). The conifer Steller'sand California (Eumetopias distributionin local and re- sea lions forestsare productive and dense, especially when jubatus,ktlophus califomianus)haul \n estimatewas made of the out on small young(Harcombe et al. 1990). rockyislands <0.25 km offshore(Lyman 1991). and animalused, based on Village sitesnearthe mouth of the SalmonRiver The rocksprovide roostingand nesting relativeto village sitesand sitesfor havebeen dated to as early as A.D. 1020 (Ross thousandsof seabirds of severalspecies (Munay in eachecosystem. Thus, the 1990).In 1854,, rhe sourhem- andMarrant 1983).The beach,rocky shore,rocky r indigenouspeople of each most Salish-languagegroup, occupiedthe area islands,estuary, dune, salt marsh, grass- :stimated.Local scarcitywas andupland (Beckhamet al. 1982).Other Salish-speaking landare within minutesby foot or canoetravel of widely usedspecies. I sug- groups lived north of the . The majorvillage sites. rmpensationsfor local scar- SalmonRiver people,referred to asthe Nechesne nt resourceorganlsms. The SalmonRiver estuaryis the smallestof (alsoan early namefor the SalmonRiver), Kowai, sevenmajor estuariesand bays used by the or SalmonRiver Indians,continued to live there Tillamooktribe; others are 1.6-12times its size well intohistoric time (Beckham1915, Beckham (Thompsonand Snow 1974).TheSalmon River et al. 1982).Their populationwas about 10 in is in and near the Salmon estuaryhas the lowestproportion of submerged 1854(Beckham et al. 1982),but had probably e northcentral andtideland of any of them, and a limited inver- beenreduced by diseasebefore 1840, ashad other y andNeskowin, just north tebratefauna. It supports4 of 16 estuarineinver- coastalgroups (Seaburgand Miller 1990).The Lncludesprimarily the Cas- tebrateslisted byThompson and Snow (1974) and SalmonRiver peoplelived primarily by fishing .esearchArea and Cascade Emmettet al. ( 199I ), comparedto 5 for theSiletz, andgathering shellfish, although hunting and use Forestof the USDA Foresr 7 for theNehalem, I I for Tillamook Bay,and l6 of marshlandsand forest were also critical to their tionalForest. I refer to this for NetartsBay. SandLake and the Nestuccaes- survival (Mumay and Marant 1983),like other Headlandscape, named for tuary supportfewer of these speciesthan the Oregoncoastal groups (Lyman 1991,Connolly d borderingthe Salmon River SalmonRiver, althoughthey are larger.The es- 1992,Erlandson et al. 1998).They apparenrlydid marshes.and grasslands in tuarinefish fauna is relativelyricher (Cornwell not hunt in the openocean. udiedscientifically anddis- etal. 2001). Of 22majorfish species(Emmetr et r plants The CascadeHead landscape was includedin and vertebratesre- al. 1991),the SalmonRiver estuarysupporrs 14, theSiletz Indian Reservation, established in 1855, :neand Blinn 1991;unpub- comparedto 15in theSiletz estuary, 18 in Netarts on file, Forestry althoughthe life of the local peopleprobably re- Sciences Bay, 19 in the Nehalem estuary,and all 22 in Forest mained unchanged.Most of the landscapewas Service,Corvallis, TillamookBay. Pre-hatcheryruns of salmon and Riverestuary excludedfrom the reducedreservation in 1875. is lessdisturbed steelheadin theSalmon River totaled4000-5000 althoughthe SalmonRiver itself remainedin the

IndigenousEcosystem Use 305 reservationuntil I 894 (Beckham1975). The area example:eating meatis one useof elk; wear- TABLE 1. List of plantspecies repor! appearsto havebeen used as pastureby settlers ing elk skinsis a seconduse; fashioning a wood- use and most likelv ecosvs from the by 1851.Although splittingwedge fiom elk antleris a third use. Type Common name earlysquatters ofEuropean lineage settled the north Each speciesof use was assignedto its most Maidenhair fem bankof theSalmon River in the 1870s,most settle- likely origin.For mostspecies, the origin wasthe Lady-fem ment occurredin the 1890s,primarily along the type of local ecosystemfrom which it was easi- Deer-fern lowlandsnear the SalmonRiver and Neskowin est to collect, basedon ecosystemaccessibility Horsetail Creek. to the villagesites and species'importance in the Licorice-fem ecosystems.Other possible origins werecultiva- Sword-fern Methods tion andtrade. Some important fiber plantsfrom Bracken fieshwatermarshes were grouped with saltmarsh; Tree Red alder The primary recordabout the SalmonRiver people BigJeaf maple theygrow on the upperfringes of saltmarsh veg- came from informantsfor two anthropological Sitka spruce etation(Frenkel and Morlan 1990).Elk (Cervus studies(Boas 1 923, B amett 1931 ). Local archeo- Shore pine logical studiesare describedby Murray and elaphus),deer (Odocoileus hemionis), bear ( Ursrzs Douglas-fir Marrant(1983) and Beckham et al. (1982).Ide- americanus),mountain Iion (Puma concolor), Cascara velopeda list of plant and animal speciescol- bobcat (Lynx rufus), and raccoon(Procyon lotor) Westernyew lected,and of their probableuses, by the Salmon occur in many habitats,and thus were assigned Westernredcedar River people.I basedthe list primanly on Bamett's to a separatecategory. Similarity of distribution Westem hemlock Vine maple detailedquestionnaire (1937), and summariesof amongorigins of speciesand uses(Table 3) was Serviceberry localhistory by Beckham(1975) and Beckham testedusing Spearman's rank correlationand the Kolgomorov-Smirnovtest. Kinnikinnick et al.(1982). Secondary sources were Boas' (1923) Hairy manzanita interview,Murray and Marrant's(1983) excava- OtherOregon coastal peoples used many spe- Hazelnut tion, Seaburgand Miller's (1990)tribal summary, cies that occur at CascadeHead but were not Salal and informationabout other Tillamook villages: countedhere (Lindsay 1995),as they were not Oceanspray Newman's(1959) excavationsat NetartsBay, referredto in literatureabout the Tillamook. Many Twin-berry Sauterand Johnson's(1914) book aboutpeople usefulspecies undoubtedly went unrecorded,es- Currant in theTillamook Bay region,and Zontek's ( 1978) pecially thoselimited to medicinal,ceremonial, Rose excavationsat Oceanside.Because data about use or emergencyuse. Thimbleberry Salmonberry of plants by the Tillamook were limited, I also For six widely-usedplant species absent or rare Willow includedplants for which groups Salish-speaking in the CascadeHead landscape, I checkeddistri- Red elderberry in southwesternWashington had a distinctname butionsof herbariumspecimens (ORE, OSC, Evergreenhuckleber (Gunther anduse 1973).lnformation from Salish WILLU) andused the preliminaryversion of the Huckleberry language groups in northern and OregonPlant Atlas data base (Scott Sundberg, Red huckleberry Canadawas not used,as they lived in a region Oregon StateUniversity, personal communica- Yellow sandverbena culturally different from the Oregon coast tion) to locatesightings of species. Yarrow (Erlandsonet al. 1998). Wild onion Angelica Speciesmentioned in works from outsidethe Results Camas CascadeHead landscapewere countedonly if I At least68 plantand 56 animalspecies were used Coast strawberry couldconfirm theirpresence in thelandscape, either parsnip by the SalmonRiver people (Tables1, 2), with Cow from observationor specieslists. In somecases, Waterleaf 308 recordeduses, counting multiple usesof a the exactspecies identity is uncertain(e.g., sev- Seashorelupine single type (e.g.,use of spruceroots in several eral speciesof bird eggs),but a reasonableesti- False lily-of-the-vall kinds of baskets).Fifty-two of the mate of the number could be made. Scientific 308 useswere Water-parsley associatedwith a resource,such as wood or nomenclature(Tables 1,2) followsHitchcock and bone, Oregon oxalis without identificationof a specificorganism, and Wild tobacco Cronquist(1913) for plants,Verts and Caraway thuscould not be assignedto an origin. To gather Pacihc silverweed ( 1998)for mostmammals, Csuti et al. ( I 997) for the speciesassociated with the 256 Wapato birds and amphibians,Maser et al. (1981)for usesof an identifiablespecies, the peopleused Goldenrod mammals not included in Verts and Carraway, all the ma- jor ecosystemsin landscape(Table Stinging nettle Emmettet al. (1991)for frsh and most inverte- their 3). The most speciesand usescame from the forests,es- Edible thistle brates,and Hafele and Hinton (1996) for other Grass-like Bentgrass tuary, and ocean, and many uses from animals invertebrates.I counteduses as in the followins Brome that occupyseveral ecosystems. A few resources

306 Zobel l meatis oneuse of elk: wear- TABLE 1' List of plant speciesreportedly used by Tillamook peopleand presenrin the cascadeHead landscape,with typesor conduse; fashioning a wood- useand most likely ecosystemor otherorigin from which it was obtained. m elk antleris a third use. Type Common name Species Usesr Originr usewas assignedto its most Fern Maidenhairfem Adiantum pedatum BF A ostspecies, the origin wasthe Lady-fem -.lb At hy r i um Ji I ix m i n a F A stemfrom which it waseasi- Deer-f'ern Blechnumspicant M S d on ecosystemaccessibility Horsetail Equisetum spp. FI A nd species'importance in the Licorice-fem Poly p od i um g ly c1- r rhi 3 IM A Sword-fern possibleorigins were culliva- Poly-stichummLtnitum BFHM S Bracken reimportant fiber plants from Pteridium aquilinum F (t Tree Red alder weregrouped with saltmarsh; AInus rubra BIW A Big-leaf maple rperfringes of salt marsh veg- Acer macrophl,llum BCI A Sitka spruce d Morlan 1990).Elk (Cervus Picea sitchensis BISW S Shorepine (Ursas Pinus contorta FIR D coileushemionis), bear Douglas-fir Ps e udots u g a menzie,s i i BW s ntain lion (Puma concolor), Cascara Rhamnuspurshiana M A r,and raccoon (Proc y-onLotor) Western yew Taxusbrevi;folia IRS sr itats,and thus were assigned Westernredcedar Thuja pLicata BHIS S rry.Similarity ol' distribution Westernhemlock Tsuga heterophtlla BIW S Shrub Vine maple reciesand uses (Table 3) was Acer circinatum BIS A Serviceberry Amelanchier nan'srank conelation and the alnifolia F \t Kinnikinnick lov test. Arctostaphykt.s uva-ursi FRS D Hairy manzanita A. columbiana 'astalpeoples used many spe- F Dr Hazelnut Con-luscornuttt BI lascadeHead but were not Salal Gaultheria shallon FM S say 1995),as they were not Oceanspray Holodiscusdiscolor IM D Lreabout the Tillamook. Many Twin-berry Lonit'era involucrata FM D ubtedlywent unrecorded, es- Currant Ribes spp. (3) F S,A ed to medicinal,ceremonial, Rose Rosanutkana F A Thimbleberry Rttbuspan,ijlorus BF S Salmonberry R. spectabilis F A edplant species absent or rare Willow Salir hookeriana CIS D J landscape.I checked distri- Red elderbeny Samhucusracemosa FM A tm specimens(ORE, OSC, Evergreenhucklebery Uaccinium0vatum F D hepreliminary version of the Hucklebeny V. ovalfolium F S database (Scott Sundberg, Red hucklebeny V.panifulium FR A :rsity.personal communica- Yellow sandverbena Abronia latifolia F D Yarrow ingsof species. Achillea millefolium MR (r Wild onion Allium sp. F \T Angelica Angelicahendersonii FS M ^i Camas Camassiaquamash F \t 56 animalspecies were used Coast strawberry Fragaria chiloensi.s F D :r people(Tables l, 2), with Cow parsnip Heracleum lanatum FM U Waterleaf countingmultiple usesof a Hy d rop h -t,I I um t enu ip es F A Seashorelupine Lupinus reof spruceroots in several littoralis F D Falselily-of-the-valley Maianthemum dilitatum F fty-twoof the 308 useswere 5 Water-parsley Oenanthesarmentosa F M ource,such as wood or bone, Oregonoxalis Oralis oregana FM S n of a specific organism,and Wild tobacco N icotiana q uadrivaLvis RS C ;ignedto an origin.To gather Pacificsilverweed Potentilla paciJica F M ed with the 256 usesof an Wapato Sagittaria latifolia F T the peopleused all the ma- Goldenrod SoLidagospathulata F DT eir landscape(Table 3). The Stinoino ncrrlo Urtica dioica FS A Edible thistle F )scame from the forests.es- Cirsium edule (t Grasslike Id many usesfrom animals Bentgrass Agro.itls sp. F D Brome )cosystems.A few resources Bromus carinatus F U Continued, next page

IndigenousEcosystem Use 30i TABLE 1. Continued TABLE 2. Continued.

Type Common name Species Uses' Origin'] Type Common na

Seagull(>l r Sloughsedge Care.robnupta F M Woodpecker Red f-escue Festucarubrct F D Unspecified Baltic rush Jtttcus balticus H M (>l spp.)eg1 Salt rush J. lesueurii H M Mammal, marine Califomia se Coastlinebluegrass Poa conJinis F D Harbor seal Small-fruitedbulrush Scirpusmicrccarpus BCF M Porpoise Sea Seasidearrow-grass Triglochin maritimunt F M otter Steller'ssea Tall trisetum Trisetum canescens F G Gray whale Beargrass Xeroph!llltm tenar BC sr "Reeds" Mammal, terrestrial Black bear BCH M Beaver "Kelp" Algae I o Chipmunk "Seaweeds" FI F Deer "Moss" Moss H S Dog Elk rTypesofusage:B=containers,suchasbaskets,boxes,orlinersfbrthem;C=clothing;F=food;H=housingorbedding, Mink including mats;I = implements,including canoe, bow, wipeq needle,rope, fish club...;M = medicinaluse; R = recreationaluse, Mole including ornaments.games, or smoking: S = ceremonialuse by the shamanor in dancesor other ceremonies;W = firewood. Mountain lio Other usesof plantsnot identifiedto species:wood, 28; grass,3; fiber, 2; rotten wood, I . Rabbits :Origin: T = probablyby trade;C = cultivated;other lettersindicate the ecosystemfrom which the specieswas most likely to be Raccoon collected A = alderforest,D = dunegrasslandorduneforest, E = estuary,G = headlandgrassland, M = marsh,both saltmarsh River otter and borderingfreshwater marsh, O = oceanand its shores,S = spruce-hemlockforest. "Sheep" (mo rindicatesthat the speciesis not listedfor the landscape,but may havebeen collected elsewhere within territorycontrolled by the Douglas'squ group. Long-tailed r Wildcat (bob rTlpes TABLE 2. List of animal speciesnoted in lists of usesby Tillamook people,with typesof use and most likely ecosystemsor of usage: B = containerssucl other origins from which they were obtained. implements, including canoe,bow, u ments, games, or smoking; S = ceren 2Uses Type Common name Scientificname Usesll Originl of animals not identified as to r 3Origin: C = domesticated animal, T = Mollusc Chiton F o to be collected-A = alder forest, D = Clam TresusnuttaLLi BFI$ o. bordering freshwater marsh, O = ocea a Cockle Clinocurdiumnutallii FO indicates that the speciesis not listet Dentalium Dentaliumpretiosum S$T group. Mussel Ml,tilus sp. FIO Octopus Octopus dofleini FO Olivellashell Oliviella biplicata SOmust have come from trade.l Red abalone Haliotis rufescens S T tivatedtobacco, as did otherwel River clam F E apparently their only attempta Snail F o ticated dog bonesoccurin Saln crustacean Barnacle F o food seems their most likely F E Crab Cancer nagi,ster Manant 1983). Crawfish Pacifastacus sp. I Fish Chinook salmon Onc o r hync h u s ts haw -,- ts c ha FI E The proportionsof species Chum salmon Oncorhynchusketa FI E among ecosystemswere signi Coho salmon Oncorhynchuskistttch FI E (Spearman'srank correlationc Herring or smelt F E p = 0.015);i.e., there were mor Lamprey Enlosphenustrident0tLts F E were more species.The "Mudcat" distribu I E origins did not differ from th Starryflounder Platichtlrys stellatus F E statistic= 0.447);i.e., there wi Amphibian Rough-skinnewt Taricha granulosa 5 A concentrationofhighly useds1 Bird Duck (5 spp.) FI E systems. Eagle HaIiaeeIus Ieucoc ep haLus S S Goose Branta canadensis FI E There were many documel Hummingbird SArnanrmalsand sometrees, inch Continued,next page lion, whale, westernredcedar (l

308 Zobel TABLE 2. Continued.

Type Usesr Origin: Common name Scientiticname Usesir Originr

Seagull(>l spp.)eggs F F M o Woodpecker S$ F D Unspecitiedsea bird F H o M (>1 spp.)eggs and young H M Mammrl, marine Califomia sealion ZaLop hu.s caliJo rnianus BCF o F D Harbor seal Phoca vitulina BCF o BCF M Porpoise Phocoenaphocoena F o F M Seaotter Enhl,dra lutris CH o Steller'ssea lion jubatus F tr Eumetopius BCF o Gray whale BC sr Eschrichtiusrobustus BFI o Mammal,terrestrial Black bear Ursus americ'anus BCH M CFS w Beaver Castor canadensis I o FHIR Chiprnunk Tamiastownsendii CF FI E S Deer Odoc'oileushemionu.s BCFHIS w H S Dog Canis.familiaris FR Etk Cen,us elaphus = = BCFHIM F food; H housingor bedding, Mink Muslela vison RS M medicinaluse; R = recreationaluse, Mole Scapanus sp. B (t or otherceremonies; W = firewooo. Mountain lion Puma concolor L w Rabbits S1y'r,i/agasbachma ni, Lep us amen ca n us FCIR ich thespecies was mostlikely s to be Raccoon Proc y-onktor rassland,M = marsh. Lt- both salt marsh River otter Lutra canadensis "Sheep" CS E (mountaingoar) Oreumnosamericanus H T Lerewithin tenitory controlled by the Douglas'squirrel Tamia s c i uru s do u g I usi i CF S Long-tailedweasel Mustela frenata B S Wildcat (bobcat) L1'n.xruJu.s L w rTypesofusage:B=containerssuchasbags,orlinersforcontainers;C=clothing;F=food;H=housingorbedding;l= f useand most likely ecosystemsor implements,including canoe, bow, wiper, needle,rope, fish club...; M = medicinal-use;R = recreationalusJ, including orna- ments,games, = or smoking;S ceremonialuse by the shamanor in dancesor other ceremonies;$ = currency. Usesr'r 2uses Originl ofanimals not identifieclas to species:bird bone,I, R; bone,IS; skin, BHI; antler,cl; feathers,IS. 3Origin:C = domesticated = animal, T probably by tradei other le_ttersindicate the ecosystemfrom which the specieswas most likely F o tobecollected-A=alderforest,D=dunegrasslandorduneforest,E=estuary,6=grassland,M=marsir,bothsaltmarshand BFI$ o. borderingfieshwater = marsh,O oceanand its.shores,S = spruce-hemlockfbrest, W = rioved widely through severalecosystems. r aindicates that the speciesis o not listed fbr the landscape,but may have been collected elscwherewiihin te,iitory controlled by the S$ group. FI O F o S o musthave come from trade.The Tillamook cul- TABLE 3. Percentagesol speciesand recordedor probable S T tivatedtobacco, as did otherwestem Oreson tribes. usesof biological materialsfbr the indigenous peoples F E apparentlytheir only attemptat farminj. Domes- in the CascadeHead landscape,by their most likely F o ticated ecosystemof origin or other origin. dogbones occur in SalmonRiver middens; Percentagesare for I 24 F o speciesand 256 usesat_ food seemstheir most likely use (Murrav and tributableto an origin. See F E text for basisof as_ signmentto ecosystems. IA Manant1983). FIE Theproportions of speciesand uses(Table 3) Ecosystem Percentage Percentage FIE amongecosystems were significantlycorrelated or other origin of species of uses FIE (Spearman'srank correlation coefficient r = 0.81, FE p = 0.015);i.e., there were more uses where there Ocean ll l1 FE weremore species. The distributionof usesamong Estuary t4 ll IE Dune,/beach originsdid not differfrom rhatof species(K-5 10 10 FE Salt marsh l0 statistic=0.44-71 i.e.. there was not a significant 7 SA Alder forest l5 17 concentrationofhighly usedspecies in a few eco- FIE Conifer forest 20 a1 systems. SS Headlandgrassland 8 6 FIE Therewere many documenteduses for large Roam throughseveral 6 14 SA mammalsand some trees, including elk, deer,sea Cultivated 2 I Continued,next page lion,whale, western redcedar (Thuja plicata), and, Acquired by trade 5 4

IndigenousEcosystem Use 309 red alder.Uses were bestdocumented fbr antler. Somecommonly used animals also are appar- 1982);thus, it seemsunlikelY t shell,and bone implements. Multiple uses of wood, ently absentfiom thelandscape. The soft-shelled caused the local scarcityof ! many recordedor inferred to be westernredcedar, cl,am(Mya arenaria),probably introduced, is the Effects of managementwith are indicatedby Barnett's(1937) infbmant and only clam reportedfrom the SalmonRiver (Th- Tillamook to make berries.gi tbr otherTillamook areas(Newman 1959, Sauter ompsonand Snow 1974),although shell from gaper perhapscamas more available(S andJohnson 1974, Beckham et al. 1982). clam(Tresus sp.) is presentin local shellmiddens l974,Boyd 1999),would have (Beckhametal. of re Some important plant resourcesfor coastal 1982).Of seveninvertebrates found though the comParisons seem tribeswere rare or absentfrom theCascade Head in middensfarther north in Tillamook territory availability in this studY (Barner of resources,al landscape.Western redcedar was not recordedin | 982),only threegrow or were found in data fot usage middens at the (Beckham distributionand abund the Tillamook County poftion of the landscape Salmon River et al. ganism 1982).Mussels (Mytilus after the time of Euro in 1908(Anonymous 1908). In the 1934timber sp.)were the mostcom- changes monly used invertebrates(Murray cruiseof the CascadeHead Experimental Forest, and Marrant Except for materialsthat su 1983),as elsewhereon the Oregoncoast (Lind- this speciesmade up only 0.26Voof timber vol- about usagedePend on the sPat say 1995).More fish thannamed in Table2 were ume (USDA Forest Service 1938) and was re- Euro-American explorersand probablyused: Oregon coastal peoples appear to corded no closer than 2.6 km to the estuary.It perience and memory of a fev haveeaten most that were available and collected grew 9 km south,also, probably within territory fulness of many speciesmay them year-round(Erlandson et al. 1998),and24 of the villages.Yew (Iaxrls brevfolia), preferred corded for the Tillamook. specieshave been identified from middens at species for bows and arrowsby the SalmonRiver people grasslandPlant Preser Seaside(Connolly 1992),on the northernfiinge from Table I wert (Barnett1937), is absentfrom the study area. but absent of Tillamook occupancyduring the historic pe- the indigenous Although yew grows in many CoastRange for- usedby Peopk riod. Washington(Norton . est types, including two with Sitka spruce western tional six were famine food (Hemstromand Logan 1986),the nearestrecord coastal tribes (Turner andDa' is from 92 km NNE. Hazel (Cory,luscornuta), a Discussion analysis of launal remainsa majormaterial for basketsand rope, is widespread Datafrom studiesofethnography, archeology, and just north of Tillamook temtor in inland CoastRange forests, but almostabsent distributionof organismsshare some problems 24 frsh taxa and 30 bird taxi where sprucedominates (Hemstrom and Logan of interpretation.Sampling is sparsecompared many more than listed in Tab 1986);it hasbeen collected on thecoast at Sand to the time and territory it is usuallyused to rep- Obtaining appropriatedistl Lake. 25 km N of the SalmonRiver and nearthe resent.Absence of an item from the recordcan- type of study is also difficu coast36 km S. Camas(Camassia spp.), widely not assureits absencefrom indigenoususe or the maps (Little 1971 , Csutiet a reporledas a staplefood plant, hasnot beencol- landscape.ln this study,most data about plant eral aspectsof organismdisl lected in the CascadeHead landscape,but her- usageare recent, from interviewsof tribal mem- dance and even var barium specimensshow that it grew at Devil's berswho surviveduntil at least1890 (Boas 1923). Presence shown on maPS.Data from Lake, at least 4 km S and probably within the Many data about animal use, in contrast,came survey and specimenc territoryof theSalmon River villages.Otherwise, from excavationof earliermiddens deposits, AD Plots the maps' failure to disPlaYt camasis rareon the centraland north-central coast. 900-1675(Newman 1959, Ross 1990, Erlandson bution required for a studYc but doesoccur in meadowsnear and eastof the and Moss 1995).Data for organismdistribution mous 1908,Hemstrom and L crestof theCoast Range. The currentherbaceous datefrom 1908to (usually)after 1970. er al. 1991). vegetationin the SalmonRiver valley,where one Two major changesin vegetationoccurred near Table 3 probably undere might expectcamas to havegrown, was probably the time of contact with Europeans.About AD tanceof someof thelesser-u riparianforest before it was convertedto pasture I 700,subsidence during an earthquakeand burial example,grasslands are repl (USDAForest Service 1999). BigJeaf maple (Acer by a tsunamideposit modified estuarinehabitats and 2 only bY food a macrophyllum),an importantsource of fiber and (Minor andGrant 1996);marsh vegetation in the Plants managedbY Tillamook wood, is uncommonin the landscape,in forests estuaryprobably recovered within severaldecades. Peotr ing of elk and deer(Sauter with spruce(Hemstrom and Logan 1986),and in In the 1840s,a large,intense fire killed most for- which are assignedto no spe this regionin general(Hines l97l ). Maple grows est in the landscape(Munger 1944).lnformants could be capturedin a variet abundantlyin the CoastRange, probably within for ethnographicstudies grew up amidst young ecosystems(Barnett 1937). teffitory controlledby the SalmonRiver people. forestthat developedafter fire. Their statements peoplehad easY access to a li Beargrass(Xeroph.vllum tenar) is absentfrom the to anthropologistsmay havediffered from those elk and deer graze,which hi landscape,but is reportedfrom foresttypes that appropriatefbr precedinggenerations. Old-growth of its perimeter. Elk bone grow on the easternfringe of the watershed.The forestthat survivedthe fire is of similar comDo- middens at the mouthof th nearestspecific record is from 92 km NNE. sitionto forestthat developed after the fire (Creene al. 1982,MuraY andMana

310 Zobel ly usedanimals also are appar- 1982);thus, it seemsunlikely rhar rhe 1840sfire Not all resourcesfor which useis claimedbv lhelandscape. The soft-shelled causedthe local scarcity of western redcedar. or altributedto theSalmon River people grow in la),probably introduced, is the Effects of managementwith fire, used by the the vicinity of known villages. Four important d from the SalmonRiver (Th- Tillamook to make berries,game animals,and materialsprobably came fiom trade:the denta- 1974),although shell from gaper perhapscamas more available(Sauter and Johnson lium (Dentalium pretiosum) and red abalone spresent in localshell middens 1974,Boyd 1999),would havedisappeared. Al- (Haliotis rufescens)shells, widely usedfor cur- l2).Ofseven invertebrates found though the comparisonsof resourceusage and rency and decorationthroughout the nor-thwest r northin Tillamooktenitory availabilityin this study seemreasonable, many coast;mountain goat(Oreamnos ame ricanzs) hair ty threegrow or were found in datafor usageof resources,and all data for or- for blankets; and wapato tubers (Saggitaria elmonRiver (Beckhamet al. ganism distributionand abundance,may include latifolia) for food. Wapatomay havebeen traded tytilussp.) were the most com- changesafter the time of Europeancontact. from eastofthe CoastRange (Beckham et al. 1982), .ebrates(Munay and Marrant Exceptfor materialsthat survivedburial, data becauseaccess through the mountainswas rela- re on the Oregoncoast (Lind- aboutusage depend on thesparse records of early tively easy.Woven blanketsin coastalOreson sh thannamed in Table 2 were Euro-Americanexplorers and residents,and ex- probablycame from importedmountain goat wool. lgon coastalpeoples appear to perienceand memory of a few informants.Use- ratherthan mountain sheep; in contrast, atwere available and collected fulnessof many speciesmay not havebeen re- farther north used hair of domesticateddoss Erlandsonet al. 1998), and24 cordedfor the Tillamook. For example, l7 (Howayl9l8). In addirion,it is unclearwherh-er t identifiedfrom middens at grasslandplant speciespresent at CascadeHead beargrassleaves, used to decoratewoven items. 1992).on thenorlhern l'ringe but absentfrom Table I were used or possibly andcamas, as food, wereavailable to the Salmon tpancyduring the historic pe- usedby the indigenouspeoples of the prairiesof River people.Where camaswas absent,people westernWashington (Norton 1979),and an addi- tradedfor it (Gunther 1973). tional six were famine foods of northwestern Someresources absent from the landscapemav coastaltribes (Turner and Davis 1993). Detailed havebeen available in the inlandterrirory con- analysisof faunal remainsat Seaside,Oregon, f ethnography,archeology, and trolledby the village,which extendedto thecrest Justnorth of Tillamookterritory, identified at least anismsshare some problems of the CoastRange (Beckham et al. 1982),in- 24 frsh taxa and 30 bird taxa (Connolly 1992), iampling is sparsecompared cludingbeargrass (Hines | 971),camas, yew, and manymore than listedin Table2. itory it is usuallyused to rep- hazel.These resources are easily transported. There an item from the record can- Obtainingappropriate distribution data for this is evidencethat residenceat the Salmon River type Lcefrom indigenoususe or the of study is also diffrcult. Publishedrange estuarywas seasonal,during tall and winter (Little study,most data about plant maps 1971,Csuti et al. 1997)show gen- (Murray and Marranr 1983), presumably with om interviewsof tribal mem- eralaspects of organismdistribution, but abun- residencefarther inland in otherseasons. Bie-leaf rntilat least 1890 (Boas 1923). danceand evenpresence vary within the ranges mapleis presentat CascadeHead in small-num- shown Lnimaluse, in contrast,came on maps.Data from detailedvegetation bers,but abundantinland within territorvcontrolled earliermiddens deposits, AD surveyplots and specimencollections illustrate by thevillage themaps' failure to displaythe detailsof distri- n 1959,Ross 1990, Erlandson The most important scarcityin the Cascade butionrequired for a study of this type (Anony- tatafor organismdistribution Headlandscape was western redcedar. Availabil- mous1908, Hemstrom and Logan 1986, usually)after 1970. Emmett ity of largewestern redcedar trees revolutionized etal.1991). ;esin vegetationoccuffed near the cultureof coastalpeoples in British Colum- Table probably . with Europeans.About AD 3 underestimatesthe impor- bia as cedarmigrated into their teritory during ring an earthquakeand burial tanceof someof the lesser-usedecosystems. For post-glacialwarming (Hebda and Mathewes 1984). it modifiedesruarine habitats example,grasslands are representedin Tables1 Local spatialvariation in its importanceon the 996);marsh vegetation in the and2 only by food plants and moles,but were Oregoncoast seems likely to have affectedcul- overedwithin severaldecades. managedby Tillamook peopleto improvehunt- ture,also. According to Bamett's(1937) infor- l, intensefire killed most for- ing of elk and deer (Sauterand Johnson1974), mant, a Salmon River native, houseshad mat- : (Munger1944). Informants whichare assigned to no specificecosystem and lineddit walls,and walls and roof werenot made udiesgrew up amidst young couldbe capturedin a varietyofways in several of planksor bark. Wood (type unspecified)was :d after fire. Their statements ecosystems(Barnett 1937).The Salmon River usedfor beams,posts, and the sweathouseroof. people nay havediffered from those hadeasy access to a largegrassland where Houseconstruction without cedarplanks differs :dinggenerations. Old-growth elk and deergraze, which hascliffs along much from thestandard Tillamook and coastal practice. perimeter. thefire is of similar compo- of its Elk boneswere common in Beckham( I 975)cites Bamert ( I 937)bur also stares rvelopedafter the fire (Greene middensat the mouth of the river (Beckhamet that SalmonRiver peoplebuilt plank houses. al.1982, Murray and Manant l983). Newman (1959)explicitly dismissesBarnett's

IndigenousEcosystem Use 3l I informant'sstatement, citing the plankedhouses land grassland,and forest, appearto have less Erlandson,J. M., and M. L. Moss. subsistencechange at the Oc at NetartsBay andBoas ( I 923) potentialfor wealth than situationsusually de- he hadexcavated Tillamook County,Oregon. Nc ashis reasons.On the otherhand, one of Newman's scribedfor Tillamook and other coastalOregon ResearchNotes 29:221 -228. housesapparently had not been of planks, and tribes. Previously unrecognizeddifferences in Erlandson,J. M., M. A. Tveskov,a Boas (1923)noted that poor peoplesometimes resourceavailability may havecaused unrecog- The developmentof maritime lived in grasshouses both in winter and summer. nized variationin size,wealth, and tradeof dif- em Northwest Coast of Nor It seemslikely that building houseswithout ce- ferentvillages. thropology35:6-22. Frenkel, R. E., and J. C. Morlan. I dar was necessary,due to its scarcity near the In conclusion,all local ecosystemswere im- Salmon River salt marshes:r Salmon River villages. Cedar was used for ca- portant sourcesof organismsfor SalmonRiver UnpublishedFinal Report to noes(Bamett 1937), the usualregional custom. people.Even so,the CascadeHead regioncould tectionAgency, Region 10, S Canoeswere an imporlantitem of trade(Seaburg not supply all important speciesused by the Greene,S. E. 1982.Neskowin Crest and Miller 1990),whereas moving houseplanks Tillamooktribe. Trade supplied some needs. Col- Supplement13.1n J. F. Fra Dyrness,and C. Maser(edit many meterslong (Newman 1959)for kilometer lectingthroughout their extensiveterritory away distancesmay not havebeen feasible. search Natural Areas in Ore from coastalvillages probably suppliedothers. guide fbr scientistsand educi The scarcityof severalplants considered stan- In the caseof cedarfor houses,however, the people vice, Pacific NorthwestFores dardresources fbr Oregoncoastal peoples, espe- apparentlyhad to do without. Station,Portland, Oregon. cially cedarand camas,may have limited the Greene,S. E., and T. Blinn. 1991.F populationand wealth of theSalmon Riverpeople. the CascadeHead Experim, Acknowledgements ResearchArea, OregonCoar In they lived near a small, primarily addition, USDA ForestService Genera This researchwas supportedby a residencyat marshyestuary (with limited invertebratediver- GTR-268. Pacific Northwes sity) of a small river (with relatively small fish the Sitka Centerfor Art and Ecology,Otis, Or- perimentStation, Portland, ( runs).Access to extensiverocky coastline required egon,provided by the NeskowinCoast Founda- Gunther, E. 19'73.Ethnobotany of !\ oceantravel, not feasibleduring stormyweather. tion, andby sabbaticalsupport from OregonState Knowledge and Use of Indil There are no estimatesof populationbefore the University.I appreciatethe commentson a draft Americans.Revised edition. Washingtt introductionof diseasethat accompaniedcontact of this paperby Ro6erlaHall, SarahGreene, Ken- ton Press,Seattle, Hafele, R., and S. Hinton. 1996.Gu Erlandson,and Mark Merlin. with Europeantraders. These villages, although neth Ames, Jon Aquatic Invertebrates.Aqual locatedoptimally amidst river, estuary, sandy and Thanksto RichardHalse and ScottSundberg for 1. OregonTrout, Portland, C rocky shore,ocean (Erlandson et al. 1998),up- help determiningplant distributions. Hall, R. L. 2001. Nah-so-mahvillage, Unpublished Report to Sea State University,and the Cr file at Department of Anth Cleaver,F. C. (editor). 1951.Fisheries statistics ofOregon. Literature Cited University,Corvallis, Orego Contribution16, Oregon Fish Commission, Portland, Harcombe, P. A., M. E. Harmon, z Anonymous. 1908.Timber cruise,Tillamook County,Vol- Oregon. Changes in biomass and pro ume 2. Unpublishedreport, on file at the Pioneer's Connolly,T. J. 1992.Human responsesto changein coastal a coastal Picea sitchensis-7, Museum,Tillamook. Oregon. geomorphology and fauna on the southernNorthwest approaching maturity. Canr Barner,D. C. 1982. Shell and archaeology:an analysisof Coast:Archaeological investigations at Seaside,Or- Research 20 : | 602- | 6 10. shellfishprocurement and utilization on the central egon. University of OregonAnthropological Papers Hebda, R. J., and R. W Mathewes. Oregoncoast. M. S. Thesis,Oregon State University, 45, Eugene,Oregon. of cedar and nativeIndian cul Corvallis,Oregon. Cornwell, T. J., D. L. Bottom, and K. K. Jones.2001. Rear- VII. Or- can Pacific coast. Science2i Barnett.H. G. 1937.Culture elementdistributions. ing ofjuvenile salmonin recoveringwetlands ofthe Records,Univcrsity oI Hemstrom, M. A., and S. E. Logan. egon coast.Anthropological SalmonRiver Estuary.Oregon Department of Fishand I : l5-5-204 and managementguide, Si Califbrnia Berkeley Wildlife, Infbrmation Reports2001 -05, Portland,Or- Beckham.S. D. 1975.Cascade Head and the SalmonRiver USDA Forest Service,Pacifi egon. estuary:A history of Indian and white settlement Ecol 220- 1986a,Portland, C Csuti,B. A. J. Kimerling, T. A. O'Neil, M. M. Shaughnessy, Unpublishedreport to USDA ForestService. Siuslaw Hines,W. W. 1971.Plant communit E. P. Gaines.and M. M. P. Huso. 1997.Atlas of Or- National Forest,Corvallis, Oregon. ests of north coastal Oregol egonWildlife. Distribution,Habitat, and Natural His- Beckham.S. D., K. A. Toepel,and R. Minor. 1982.Cultura- State University, Corvallis, ( tory. OregonState University Press,Corvallis, Oregon. resourceoverview of the Siuslaw National Forest, Hitchcock.C. L.. and A. Cronquist Emmett,R. L., S. L. Stone,S. A. Hinton, and M. E. Monaco. westemOregon. Heritage Research Associates Report cific Northwest. University r 1991.Distribution and Abundanceof Fishesand In- 7. Lake Oswego,Oregon. attle, Washington. vertebratesin WestCoast Estuaries, Volume II. Spe- Boas, F. 1923.Notes on the Tillamook. Universityof Cali- Howay, F.W. 1918.The dog's hairbla cies Lif'e History Summaries.Estuarine Living Ma- fornia Publicationsin American Archaeology and Eth- The Washington Historical ( rine Resources Report 8. National Oceanic and nology20:3-16. Lindsay, L. W., Jr. 1995. Native use l -30 1n R. Boyd (editor). AtmosphericAdministration, National OceanService, Boyd, R. I 999. Introduction.Pages egon coast. Pagesl9l-210 In Land in the Pacific Nofthwest, Strategic EnvironmentalAssessments Division, Indians,Fire, and the of the Coquille Estuary: Nat OregonState University Press, Corvallis, Oregon. Rockville, Maryland.

312 Zobel d forest, Erlandson. appearto have less J. M., and M. L. Moss. 1995. Chronolosv and the Oregon Coast. Words and pictures Unlimited. h than situations subsisrenr.echange al rhe usually de- Oceansideslre 1::_ii_+7 r. Corvallis.Oreson. )ok andother Tillamook County,Oregon. Northwest Anthropolosical coastalOregon Little, E. L., Jr. 1971.Atlas of UnitedStates Trees. Vol. ResearchNotes 29:22 I -22g. l. unrecognizeddifferences Conifersand ImportantHardwoods. in Erlandson, US Departmenr J. M., M. A. Tveskov.and R. S. Byram. 199g, y may havecaused unrecog- of AgricultureMiscellaneous publicurion I l,{6.U. S. Thedevelopment o[ maritimcadaptation, on thesourh_ printing iize,wealth, and tradeof Government Office, Washington.D. C. dif- ern Northwest Coast of North America. Arctic Lyman. prehistory An_ R. L. l99l . of the Oregon-Coasr:The Ef- thropology35:6-22. tects of ExcavationStrategies and AssemblageSize Frenkel,R. E., and J. C. Morlan. 1990. Restoration on ll localecosystems were im- of the Arr.haeologicalInquiry. Acadrmic pre,:. SanDr_ Salmon River salt marshes: organismsfor SalmonRiver retrospectand prospect. ego,California. UnpublishedFinal Reportto US Environmentalpro- Maser.C., e Cascade B. R. Mate,J. F. Franklin,and C. T. Dyrness.l9g I . Headregion could tectionAgency, Region 10, Seattle, Washinston. Natural coast mammals. USDA )ortantspecies used by the Greene.S. E. 1982.Neskowin Cre:r ResearchNaru-ral Area ForestService General Technical Report pNW_ 133. de supplied Supplement 13. In J. F. Franklin, someneeds. Col- F. C. Hall. C. T. PacitlcNorthwest Forest and RangeExperimenr Sra_ theirextensive territory away Dyrness,and C. Maser (editors).1972. Federal Re_ tion, Portland,Oresorr. searchNatural Areas Minor, es probablysupplied others. in Oregon and Washington:a R., and W. C. Granl. 1996.Earthquake_induced sub- guidelbr scienrisr\ orhouses, andeduralor\. USDA ForesrSer- sidenceand bunal of lateHolocene irchaeological sirer. however,the people vice,Pacific NorthwestForest and Range Experiment northemOregon coasr. Arnericun entiquirv 6l:772_ o without. Station,Portland, Oregon. 781. Greene, S. E., andT. Blinn. 1991.Research publications of Munger,T. T. I 944. Out of the ashesof Nestucca.Americal Ints rhe Ca:cadeHead ExperimenralForest antl Scenic Foresrs50:342-345, 367_ 368. Research Munay, Area, OregonCoast Range, 1934 to I990. T. A., andJ. Marranr.19g3. preliminary sampling of supportedby a residency at USDA ForestService General Technical Reoort pNW_ theThree Rox site.35_ LNC_33. on th" trr"gon cu"ast. ,{7-55 r Art and Ecology,Otis, Or- GTR-268.Pat.ific Norrhwe5r Forest an.l RanqeEx_ Pages 1nD. E. Dumond redjtor),Coniributions portlund. ro he perimentStation. Orepon. the Archaeologyof Oregonlggl_19g2. Associa_ NeskowinCoast Founda- tion Gunther.E. 197.1.Ethnobotany ol WestemWashington: The of OregonArchaeologists Occasional papers 2, )alsupport from OregonState Portland, Knowledgeand Useol lndigenousplanr, by Native Oregon. iatethe Newman,T. commentson a draft Americans.Revised edition. University of Washing_ M. 1959.Tillamook prehistoryand its relation :naHall. Sarah Greene, Ken- ton Press.Seattle, Washin|]ton. to the NorthwestCoast culture area. ph. D. Disserta- tion. University landson,and Mark Merlin. Hafele.R.. and S. Hinton.IqgO. Cuide ro pacificNorrhr.resr _. of Oregon,Eugene, Oregon. Norton, H. H. 1979.The {alse AquaticInvertebrates. Aquatic Biology Series:Book associationbitween anihroposenic andScott Sundberg for prairies l. OregonTrout, portland,Oregon. and imponantlbud planrsin ucstemiash_ antdistributions. ington. NorthwestAnthropological Hall.R. L. 2001.Nri-:o-mah village.r iewedLhrouqh il> fauna. ResearchNotes I 3:I 75-200. Unpubli:hedRepon to SeaCranr prug.,r=m.Oregon OregonDepartment of Fishand Wildlife. StateUniversity, and the Coquille Indian Tribe. on 1997.Salmon River Basin fish managementplan. 951.Fisheries statistics of Oregon. file at Departmentof Anthropology, Oregon State Salem,Oreeon. Ros:.R. E. I990. )regonFish Commission, ponland, University,Corvallis, Oregon. Prehisrorlofthe Oregoncoast. pcges 554_ 559/a W. Sutrlesreditorr. Handhook Harcombe.P A.. M. E. Harmonland S. E. Greenc.1990. oiNonliAmeri_ can IndiansVol. 7: NorthwestCoast. ran responsesto changein coastal Changesin biomassand productionover 53 yearsin SmithsonianIn_ stitution,Washington, D. C. ndfauna on the southem Northwest a coastalPiceu sitthensis-Tsuqa heterophll/a forest Sauter,J., and B. Johnson.1974. Tillamook gicalinvestigations at Seaside,Or- approachingmaturity. CanadianJournal of Forest lndians of the OregonCoast. portland, cf OregonAnthropological papcrs Research20:1 602- 1 610. Binfbr

IndigenousEcosystem Use 313 Verts,8.J.,and L.N. Carraway.1998. Land Mammalsof Or- Zontek.T. I 978.Test excavations and cultural resource evalu- DavidM. Braun, Bi Runcheng, egon.University of CaliforniaPress, Berkeley. Cali- ation of the Oceansidesite (35-TI-47).Unpublished CraneResearch FacilitY, 1262 F fbmia. manuscript,Department ofAnthropology, OregonState Wiedemann.A. M. 1984.The ecology of University.Corvallis, Oregon. coastalsand dunes: a community profile. USDI Fish and Wildlife ServicePublication FWS/OBS- 84/021. Washington,D. C. Folivoryof VineMa1 Forest Received5 December2001 Accepted publication I6 May 2002 for Abstract

Folivory of vine maplewas docun consumption by lepidopteranlarva the oeriod fiom bud break to leaf r"u.ing to adult. Weekly folivory 1 seasonalherbivory was 9.970 oflea representedby eight taxa in the G served, suggestingthat the folivon wasa signilicantcomponent ofsta vine maple is a regionallYwidesPr communities and leaf-basedfood '

lntroduction Herbivory in forestedecosY consumption of foliage,Pl woody tissueby animals.TY herbivorylevels of7.l% for ll.l7o for shade-tolerants tropics havebeen reported, t (Coley andBarone 1996).It folivore outbreaksof the' w orm \Cho ri st on e u ra occi( (Brookes, 1987)or the wer (Lambd ina fi sc e llaria lugul (Harris et al. 1982)in ten can produce defoliation aPtr bivory also variesspatially, to 54Eo along an elevation (Lowman 1995).Studies at measuring holes Producr arthropods,and therefored( Old- growth conifer fores west west of the CascadeR have defoliator outbreaks(Pr Exceptions are outbreaksc the western hemlock looPet ally producespatches ofint( growth stands dominated (Ts u g a het e rop hy I la) (Han

rAuthor to whom correspondenc mail: [email protected]

314 Zobel