Recovery Patterns i n Stream Communitie s Impacted by th e Mt . St . Helens Eruptio n by Peggy Wilzbac h Thomas L . Dudle y James D . Hal l Water Resources Research Institut e Oregon State Universit y Corvallis, Orego n WRRI-83 August 1983 RECOVERY PATTERNS IN STREAM COMMUNITIE S IMPACTED BY THE MT . ST . HELENS ERUPTIO N by Peggy Wilzbach, Thomas L . Dudley and James D . Hal l Fisheries and Wildlife Department, Oregon State Universit y Final Technical Completion Repor t for Project No . A-059-OR E to United States Department of the Interio r Washington, D .C . 2024 0 Project Sponsored by : Water Resources Research Institut e Oregon State University Corvallis, Oregon 9733 1 The research upon which this report is based was financed in part by the U .S . Department of the Interior, Washington, D .C . (Project No . A-059-ORE), a s authorized by the Water Research and Development Act of 1978, P .L . 95-467 . Contents of this publication do not necessarily reflect the views an d policies of the U .S . Department of the Interior, nor does mention of trad e names or commercial products constitute their endorsement or recommendatio n for use by the United States Government . WRRI-83 August 1983 ABSTRAC T Recovery patterns of benthic communities in seven small streams impacted t o varying degrees by the eruption of Mt . St . Helens were examined in winte r 1981-82, particularly in relation to habitat constraints . Overall abundance an d diversity of the communities were low, and instability of channel banks an d bottom appears to severely limit re-establishment of biota . Communities appea r to be undergoing cycles of establishment and denudation in concert with change s in landscape geomorphology . Disturbance events of varying frequencies were als o simulated in experimental stream channels to more closely evaluate the respons e of benthic communities to periodic, non-catastrophic disturbances and to compar e this with tie response observed to catastrophic perturbation associated wit h volcanic eruption . Results of simulations suggest that non-catastrophi c disturances may not play a major role in determining stream community ove r ecological time because the predictability of these events may have enable d evolution in species of adaptive responses to cope with disturbance . Catastrophes appear to have different consequences on stream communities fro m other disturbances, and should be distinguished from them . t FOREWOR D The Water Resources Research Institute, located on the Oregon State _ . University campus, serves the St:a.te of Oregon . The, Institute foters , encourages andrfa•cilitates water resources research and education inyo .l . viig al l aspects of the quality and quantity of water available for beneficial use . Th e Institute administers and coordinates statewide and regional progpmms o f multidisciplinary research in water and related land resources . The Institut e provides a necessary communications and coordination link between . the'agencie s of local, state and federal government, as well as the private sector, and-M e - broad research community , at universities in the state on matters of water related research . The Institute also coordinates the . inter-disciplinary program of graduate education in water resources at Oregon State University . It is Institute policy to wake available the results of significant water= related research conducted in Oregon's universities and colleges . The Instirtut e neither endorses nor rejects the findings of the authors of such research . I t does recommend careful consideration of the accumulated facts by those concerned with the solution of water-related problems . ACKNOWLEDGEMENT S We thank the U .S .G .S . Cascades Volcano Observatory, Vancouver subdistrict , for providing access to and field assistance in sampling sites within th e restricted area surrounding the volcano . The Weyerhaeuser Company (Tacoma , Washington) generously donated use of the Kalama Springs research facilities . P . Bisson is especially thanked for his assistance in logistical matters . K .W . Cummins made helpful comments on the manuscript . N .H . Anderson an d C .P . Hawkins are thanked for use of their data . ii TABLE OF CONTENT S I 1 INTRODUCTION N II METHODS "3 3 Mt . St . Helens Sites 3 Biological Sampling 3 Habitat Characterization 6- • 6 Kalama Springs 6 Site Description . Experimental,Manipulations 7 9 III RESULTS Recovery Patterns Of Streams Impacted By Mt . St . Helens Habitat Constraints 9 9 ' : Biological Properties 13 16 Experimental Simulations of Disturbance 16. Benthic Invertebrates Periphyton IV DISCUSSION V LITERATURE CITED VI APPENDICES 27 r 29 31 1. Channel Stability Evaluation 2. Invertebrate Taxa Collected From. Streams Impacted By Mt . St . Helens, winter 1981-82 32 33 'rJ 111 LIST OF ' FIGURE S Figure Pag e 1. Map showing location of study sites . 2. Substrate stability indexed by mean particle size an d the coefficient of variation . 14 3. M croinvertebrate density in Ape Canyon since the eruptio n May 1980) of Mt . St . Helens . 17 4. Taxonomic richness of macroinvertebrates in Ape Canyon an d an unimpacted stream since the eruption of Mt . St . Helens . 18 5. Abundance of macroinvertebrate taxa at Kalama Spring s three months after disturbance simulations . 19 Proportional biomass of invertebrate functional groups a t Kalama Springs three months after disturbance simulations . 15 6. 4 LIST OF TABLE S Table Page 1. Impact experienced and physical characteristics of strea m sites near Mt . St . Helens . 2. Habitat characterization of the study sites, winter 1981-82 . 3. Channel stability evaluation of study sites nea r Mt . St . Helens, winter 1981-82 . 12 4. Biological parameters of benthic assemblages in streams near Mt . St . Helens, winter 1981-82 . 15 5. Community composition parameters of benthic invertebrate s prior to and after disturbance simulations at Kalama Springs . 23 6. Similarity of invertebrate assemblages among channels prio r to and after three months of disturbance simulations a t Kalama Springs . 24 iv 5 - 10 I . INTRODUCTIO N Life cycles of organisms are adapted to the stability of the habitat i n which they occur, and organisms may evolve a dependency on the disturbance ` regime that regulates habitat persistence . When the ratio of disturbanc e interval to generation time becomes too large, or when disturbances are extreme , organisms are not capable of an adaptive response and cannot persist in, an are a (Paine 1979) . For this reason, Harper (1977) suggests that catastrophes (large , extreme events destroying large portions of the population) should b e distinguished from disasters (more frequent, locally disruptive influences) eve n though they form a continuum, because of the different consequences these event s have on community structure . The effects on the biota and habitat of small streams near Mt . St . Helen s from the May 1980 eruption were certainly of catastrophic proportion . Impact s on stream channels associated with their eruption varied from light to hep y tephra deposits to pyroclastic flows, mudflows, flooding, and landslides . Th e effects realized by the biota included tissue abrasion, injury by impact wit h rock, smothering, burial, and incineration . Rates-and patterns of recovery of the impacted stream commumities ar e dependent on the type and intensity of the impact received, and on its spatia l scale . The areal extent of the damage affects the distance from a source o f potential colonists, and this may determine the life history and behaviora l attributes of the initial colonizing species . The type and intensity of th e impact affects the extent of mortality in the community from the disturbance , and may also affect susceptibility of the stream channels to disruption fro m further perturbation . Rates of recovery will vary inversely with disturbance intensity, an d recovery will be faster in streams with refugia than in streams in which al l internal sources of colonists were destroyed . Return to pre-eruption state s should occur most quickly in streams in which damage was slight and patchy . Th e mechanisms and course of succession in this situation may be similar to thos e occurring an a small spatial scale in 'typical' streams experiencing patch y disturbance from periodic, non-catastrophic floods . 1 Little is known, however, of the role non-catastrophic disturbances play i n determining stream community structure . For this reason, evaluation o f recovery to prior equilibria) conditions is difficult . Periodic disturbance s - may prevent the attainment of local equilibria, and the species composition of a stream community may be continually changing . The objectives of this study were to examine recovery of benthi c communities in small streams impacted to varying degrees by the eruption of Mt . St . Helens, particularly in relation to habitat constraints, and to compare thi s with the response of benthic communities to periodic, non-catastrophi c disturbances . The latter was evaluated by simulating disturbance events i n artificial outdoor channels and monitoring colonization patterns .o f invertebrates and algae . Use of the Weyerhaeuser's Kalama Springs researc h facility for this purpose enabled us to examine response of a species sourc e pool that is similar to that potentially available for recolonization o f streams impacted by the eruption of Mt . St .-Helens . II . METHODS • Mt . ' St . Helens:, Sites : Seven sites, in small streams impacted to varying degrees byavalanc .he-s acme( ' mudflows from the May 1980 and subsequent eruptions of Mt . St . He]•erns, wer e selected for biological sampling . These were chosen to coincide with sites ., established by the U .S . Geological Surve y ' s Cascades Volcano Observatory and the . U .S . Forest Service for studies of erosion and sediment routing . Site locati-€is. are shown in Figure 1 and a description of impacts experienced and physica l characteristics of the streams is listed in Table 1 . One of the sites, on th e upper Kalama River, received only minor mudflow from the eruption, and it wa s selected to serve as a control . Biological sampling and habitat swrveys- wer e conducted one to two times at each site from late October 1981 to early,Febru a .ry~ 1982 . Winter access to the sites was limited'to helicopter travel, an d unfavorable flying weather prevented more frequent sampling . Additional data from Ape Canyon are provided from short-term intensive "Pulse" studies (funded _ by the National Science Foundation and the U .S . Forest Service) during the ' summers of 1980 and 1981, and from subsequent studies of N . Anderson, and C . Hawkins (Oregon State University) . Biological Sampling : Benthic invertebrates were sampled with a modified Hess, sampler . Three t o six samples were taken at each site and the substrate type associated•with eac h sample was noted . Qualitative samples of the invertebrates attached to larg e woody debris were collected . Samples were preserved in 70% alcabQl, and sorted - in the lab . Invertebrate biomass was estimated from length-weight regression s (Rogers et al . 1976, K .W . Cummins, unpubl . data) . Standing crop of chlorophyll= _ a and phaeophytin was estimated from rock scrapings to provide an indexof al l biomass and photosynthetic capacity of the stream sites . Twenty rocks, eac h 8-12 cm in diameter, were collected from each site, and chlorophyll levels fro m these were measured by acetone extraction (Wetzel and Likens 4.9{7) . to S 3 4 m L a) -- DI (tl C- •,- .. - o LC) CT rl Cl C) 01 C) I- C) Cl CO 0-1 M M LC) LC) CD LC) Cr) t- ~--r ri CD l0 C) LC) LC) CD LC) I- LC) N. N N CV N r\ LC) C)1 M CO 01 d' d CO •k a, L. C O •r 4-) '0 ,N E CI) w L CD L L 4-) tT C a) ^ E d,-4 CL) L. [n C) CO 01 0 -1-) r •r 4- N 0 U Q 1-4 >1 (Cl CO -1-) U (O a) 0. -C E +-~ a) C 0 •r 4-) 0. L E w 0 N RS +3 r U) U (0 0r L >) 0_ 3 0 r 4TJ E a) L. a) > a) U) 0 N Q) • .wr 4-) N r U CA C 0'0 a) r -Cl ^ 3 44) 0 V) r CO 4r .0 E 4-) r CA 0 a) -C7 4-) N (0 r CO . C 0 E N r Cl) 0 4- L (0 }> r /0 > $ CLI a) C 4-) CO L E t a) C E a) a) T1 0 E L 4- C 0 >> C Cl) 4.) .r (0 U a) Q 16 (0 7 (0 L. CO /0 L. CO CI) CZ 0. > CD 0. 0. > a) - 5 0 J a) •I-•) 0 4- E 4-) C L O 0 U C •r E - CA 01 (Cl C C •r -0 L C/) (0 L) 4•r Cl) t0 (0 r0 r +O Habitat . Characterization : Ten line transects were established at b-m meter intervals at each site . .At each transect3 width of the wetted channel, average depth, water velocit y (measured at 0 .6 water depth), and substrate type were recorded . Substrate ' categories were broken down into the following : sand (< 1cm), fine gravel (1-2 ' cm), coarse gravel (3-4 cm), pebble (5-8 cm), rubble (9-16 cm), cobble (17-3 2 cm), small boulder (33-64 cm), large. boulder (> 64 cm) and bedrock . Pebbl e counts (measurement of the length of the longest linear dimension of 10 0 randomly selected sediment particles) were taken at each site to-Provide estimates of mean particle size . Cherrn.el stability of each site • was evaluate d with the Pfankuch rating scheme (1975, Appendix 1) . Drainage areas, eleva•tion , , ' and stream length of the sites were estimated from U .S .G .S . topographic maps . Stream gradients were measured with a clinometer . Kalama Spring s Site description : Three outdoor artificial streams, built and owned by the WeyePir use r Company, were-used to experimentally investigate the response of stream bentho s to periodic, non-catastrophic disturbance . The streams are fed by Kalama Springs, a cold-water spring draining into the Kalama•kiver, in Co►1litz Co . , Washington . The three streams are designed as a series of alternating pools ah .d riffles . The riffles are 6-10 cm deep and 1-2 rn wide, in lengths varying fro m 8-16 m . The pools are 30-50 cm deep . Approximately 10% of the flow from Kalam a Springs is diverted into a series ;of ponds above the streams . Manipulation o f wiers between the ponds allows .a constant flow to be maintained in each streams, Substrate of the channels consists of coarse gravel ., 2-4 cm in diameter, whic h overlays a fine pumice base . Water temperature is 5-7°C throughout the year . The streams are open, with no riparian canopy . 6 Experimental manipulations : Disturbance events were simulated in two of the three channels by releasin g a flood of water from the holding ponds above the channels and by agitating th e substrate with rakes . In channel 3, disturbances were simulated once every 2 weeks, and in channel 1, once a month . Channel 2 was left unperturbed . Benthi c invertebrates and algae were monitored biweekly for 3 months following tM e initial perturbations in the first three riffles below the .holding pond of eac h stream . Invertebrates were sampled with a modified Hess sampler, and 8-16 tile s were placed in each riffle to investigate algal colonization patterns . Thi s experiment was conducted from June to September , 1982 . 7 III . RESULT S Recovery Patterns In Streams Impacted By Mt . St . Helen s Habitat constraints : Although a survey of habitat parameters based on one or two sampling date s can be misleading, the data provide some indication of constraints imposed b y the physical setting that may limit the colonization and persistence of benthi c invertebrate and periphyton populations . Measurement of such properties as stream gradient and mean channel widt h and depth provided a means of scaling the study areas so that comparisons coul d be made . Values that were measured are all fairly similar among the site s surveyed (Table 2), indicating that comparisons among sites are appropriate . Water temperatures reflect the time of day and ambient temperatures at sites o n the dates at which these were measured, and values are included only to indicat e that these sites were not thermally affected at this time from volcani c activity . Discharge values that were estimated are not abnormally high, an d flows at this time of year were probably stable . Peak flows in streams at thes e elevations generally occur in late sping - early summer and late fall - earl y winter . Unit stream power, the rate of doing work of a stream per unit o f '±1_, where p = density of water, g = acceleration of gravity , width, equals w Q = discharge, s = slope gradient, and w = channel width (Leopold et al . 1964) . Channel widths of the impacted sites were somewhat greater than width s typical of streams of low (first to third) order . This sugggests that th e channel banks of the sites, except where they are controlled locally by bedrock , are being continually eroded and re-worked . Because rooted riparian vegetatio n and large organic debris are absent from most of the sites, the stabilizin g influence of these elements on channel morphology is likewise absent . Pfankuch's (1975) rating scheme was used to evaluate channel stability of the impacted streams (Appendix 1) . The value of this approach is that , particularly in sites experiencing a variety of impacts associated with th e eruption, many different attributes of the channel banks and bottom ar e considered, and are scaled relatively in importance to sediment yield . Whe n making an evaluation, one is less likely to be influenced by a single obviou s destabilizing factor . Mass wasting, for instance, appears overwhelming in mos t streams surrounding Mt . St . Helens, but the presence in some sites of large , angular rocks and woody debris provided stabilizing influences . 9 (CI IU r- cci a) 4-•) (Un r C C o - L L 0_ go C-, v .) LC) 01 g-- i r+ d• CD CD N cal N CO CO 3 O r1 O CD O M O CD Lo d• O O tO .-I M CO CO a> Lo Lo CD 01 O C M d M 01 CV rI N o t-) CO .-- 1 ba) c E O N CC) N- M M N 0 M M ' -4 n O I\ M O r♦ tO r .0 M CO M N 71 a) 01 LC) C b a) '-I CD 0 C S] CO O N 01 r- I 112 01 N Co 01 rr 1 r♦ O 01 rr L a) 4-) ra 4-1 4 L O LC) b L CT O d- n t0 a) 3 CT r1 N CD L[) r1 N M rl LC) l0 O J C /C CU M n 0 C . IO 0 N U *-I N C ra +-) a) +-) R) L ra O CD Lf) LC) d L O a d ' -I '0 -4 CT N --I O L a) 0 a) O - d' '-I w O .4-, N M a) O CO O 0 .0 . N O E M U N a) 0 0 C O •r •rNL a) I tUO L Lo 0_ C 0 U a) UU( O a) R3 F- C O T C) w LC) 4-' r a' a b R ( a.1 O^ E (.3 o O C5 N E L 1 cry M N CD C) co Lo • r- I r-1 n CO a) r- O1 -CI -C) O U hw tO .-- I tO 00 O1 d• d CO r--1 LO N O • O N • a) r n M _ w d' r-I E 4-)C a) L CO rt O CO O CC) n O tO E tO CO CD rt +.) N ri LC) a) 0. N CD O L . co O C ra C a) U .- as C 4--I a) a) -a b U 3 is R C a) 3 rt3 a) 10 a) O 4 C) "O U r• 0,4 O 4-) O 4-) 0 •r 4-a Iq L r- 1 ro r U 'I NU N n i u) v) C) E d a L) C77 L -- Q E 'U L N •I-+ ia ) a) L a) 3 O r aC +) +•) C ra L rC1 C +) •r N E _o O 0 ON x N a) v U •r w }1 __-. L E (U 0_ U 0 co I C a) ra N a) • r N •k Ix Channel stability of the impacted sites was generally evaluated as poo r (Table 3), and ranking of scores reflect reasonably well .the type and magnitud e of impact experienced . Carbonate Springs received a very poor rating . Strea m power (indicating its potential to move sediments) of the site is high (Table 2) ; it is a very active, braided channel cutting a floodplain in the avalanche zon e of Mt . St . Helens . Lower Maratta Creek received the worst 'score . It, too, i s located in the avalanche zone and received blast deposits . Upper Maratta , 0 .5 km upstream from the lower site, received a better stability rating . I t received blast deposits but lies just outside of the landsl=ide area . Differences in rating of the two sites largely reflect difference•,$ in channe l capacity, especially indexed by width to depth ratios, and in-cutting of th e lower banks . The bank rock content of bath sites i& fairly high, but mu .chr mor e mud is present in the lower site . The Pfankuch scheme does, however, possess limitations, particularly i n relation to biological activity . It was developed for use in the Rock y Mountains, and was designed to predict horn likely a channel is to yield'sedimen t at high flows . The scheme is of ecological importance inasmuch as unstabl e sediments are of ecological importance . Considerable biological activity ca n occur, however, in unstable streams . The major limitation of the Pfankuch scheme for evaluation of channe l stability in streams of the Pacific Northwest, and perhaps the entire country , is the inadequate emphasis it places on the role of wood debris . The pretenc e of wood debris on slopes is rated inn the scheme as a destabilizing facto r because it increases sediment yields when it is moved . The role large woody debris plays as a major .structural feature within stream channels (Ke1 . le an d Swanson 1979, Swanson and Lienkaemper 1978) is not considered . Wood debri s creates a stepped gradient and although it increases stream power locally ove r channel drops, it decreases stream power throughout a reach ; this lend s stability to the channel . The 'control site' surveyed on the upper Kalama Rive r received a stability rating of good . Even though the site has a high st•rea . m power (Table 2), the presence of considerable in-channel debris and of roote d 'riparian vegetation provides .a geomorphic control that appears to override it s capacity to move sediments, and the site stability should probably be rated_, .a s excellent . '11 Table 3 . Channel stability evaluation of sites near Mt . St . Helens, winter 1981-82 . Item Rated Ape Canyon Upper Muddy Upper Maratta Lower Maratta Carbonate Spgs . Swift Kalam a UPPER BANK S Slope 6 6 8 8 6 6 2 Mass Wasting 12 9 12 12 12 12 6 Debris Ja m Potential 2 2 6 8 2 8 6 Vegetatio n cover 12 9 12 12 12 6 3 Channe l capacity • 2 3 2 4 4 4 2 Bank Roc k content 8 4 4 4 8 8 2 'Obstruction 4 4 8 8 2 6 4 Cutting 12 8 12 16 16 4 4 Deposition 12 6 11 12 16 12 4 Ro c.k _ Angularity 2 2 1 1 2 1 1 Brightness 2 2 1 1 2 1 1 Consolidation 6 4 6 6 6 4 4 Bottom Siz e Distrib . 16 16 16 16 16 16 8 Scouring , Deposition 24 24 24 24 24 20 12 Moss and Algae 4 3 4 4 3 4 2 (Mood Debris) + - + + - ++ ++ 124 102 127 136 131 112 61 LOWER BANK S BOTTO M i TOTAL SCORE (Poor) (Fair) (Poor) (Poor) (Poor) (Fair) (Good, 4 The presence in the channel bottom of large angular rocks, accordin g . to the Pfankuch evaluation, is considered to be beneficial as these are not easil y moved . Small, rounded rocks are destabilizing . More critical, , perhaps, from a biological perspective, is the presence in the substrate of a divers e assortment of sediment sizes . Because they can interlock, sediments ar e probably less likely to shift if they are of different sizes . Substrat e heterogeneity also prbvides habitat complexity, which is positively correlate d with biological diversity (Gorman and Karr 1978) . As another approach t o channel stability evaluation, we plotted the coefficient of variation agains t mean sediment size (Figure 2) . Large sediment sizes with a small coefficient o f variation may provide substrate stability, but generally large particle siz e with a large coefficient of variation is indicative of good substrate stability , and small particle sizes with a small coefficient of variation indicate les s stability . Evaluation of the stability of the study sites ; with the exceptio n of two outliers, corresponds fairly well with the Pfankuch rating . Uppe r Maratta, because of its large sediment size, and Lower Maratta, because of th e large CV, were included among the more stable sites (Figure 2) . The strea m power of these sites may be too low to affect sediment distribution . Biological Properties : Overall abundance and diversity of winter assemblages of benthi c invertebrates in Kalama River relative to the other sites (Table 4) indicat e that it serves as a reasonable control site . Composition of the invertebrate fauna present at each site suggests habita t : conditions and the nature of the food resource base . In Carbonate Springs, onl y burrowing and/or case-building midges were present (Appendix 2) . This indicate s that the most likely food resource is fine particulate organic matter (FPOM) , which is probably trapped in sediment interstices or in the periphyton film . The absence of taxa that require stable substrate for attachment, such a s filter-feeding collectors or large mayfly or caddisfly scrapers, reflect s substrate instability . That habitat instability may override food resource - limitations is suggested by the presence in Carbonate Springs of a fairly hig h standing crop of chlorophyll-a (Table 4) . The only site in which scrapers wer e collected is Kalama River, which exhibited slightly lower chlorophyll levels . Shredders were also abundant and diverse at Kalama River . This reflect s availability of allochthonous coarse particulate organic matter (CPOM) from th e 13 6 Site 7. 6. 3. 4. 2. I. 5. Maotoinvertebrates Density Richness Ka lams 26 Swift 29 U. Maratta 7 L. Maratta U. Muddy 478 Ape 14 Carbonate Sag, . 9' 14 3 2 1 7 4 7 10 20 Channe l Stability Good Fair Poor Poor Fai r Poor Poor hood ++ ++ + + + 30 40 Mean Sediment Particle Size (CM) : Fig . 2 . Substrate stability indexed by mean particle size and the coefficien t of variation (CV) . CV = 100 s x X Dashed line represents a proposed boundary separating stabl e and unstable habitats . 14 VT 1• a a) 03 +) X b • a) .O R3 C 1 +-) O ,, N CM 0 . CO O 4- L a -0 4- O M -Cl Ir-1COU O I COO L ~ g Q) N a) ri V) C_0 • L Y (O E C Cl) L U Q) (O d a) 0 E C 7 a •I•-) C E r- •r >) 0 ( O 3 4-) L a •.) w C 4- 0 -. a ^a) +) +) Q) V) 3 -Cl 4- IL C 4-) Cl) a) • r Q) +) s 3 C ) r- E CO +•) V) ri W O E = L •r V) -0 C 11:1 w 4- +-) . N U ▪ a) V) a) V) C .) V) `] C a) a) ••r L C CO C +) O. (O .C .--1 0 L N•r w C 4-) Cl) L U V) L (O n:) CO a! N E U E ~ C 4- 0 .8- (O a) 0 •r E r- d V) -0 O b < E N C ]G `•~ (0 C 0 CL) rts C X V) N a) C t CU U }) • r (CI Ce L -0 U a)• r +) E L O Q) > C ~-•1 • i d t` Lt) N M . M r-a r'I N -I N- ) d CO .-1 M 01 • . • d r4 C O X ( 3 I- N N (O ^ Ec.L 0 E r CO LC) a) . +) b\ L O) E a1 u 4-) L N • N • r-1 C) -1 N n LO M N • 0) _I .1 a) > C P-4 La) CO CO • C 4-) E I- U rO H 4-) n ~ a •r N C O ^ c.L a) CO L L N a) a) 0) a) U•r >+) C . C (O r O r I .C IO I ~' 4) 1 0C . - C) E 0 a) r • C (O C .Cl a a) • r V) (O .C +) r4- Q •r Q Cq L L N r(3 a) a V) 0 .C V) 4•) d L .- U • r CU C (3 W to 4) U Cl) C a) O s E4- 4-3 U C (O O (O C CO ( O L O a) 7 (O0 .a) Q O 4•) V) 4.) LC) L (O a) (0 .-- I +) LC) (6 O L O CU +) 0 LZ d r-1 CO N- d r+ 0) CO N LO N 1 • a) > C t-r .3 -C 0 L\ 0 0) r- .._. E L L) O d r-1 n r U CS .-rtf 0- 0 L) D)o)E (U .--, •r C C (O +-) (1 ) C)• .- •r N U O) O-0 r- Cl) C r C Q N r-• r q (O E V) ~- L •r 4-) (O W O 0CO N V)= UV) n N CO LC) M N .i LO L Cl) 4•) 0 J 3 V) o N M LO . .--I (O 4-) •1•) CO L b 4-) -1• ) (O g X L Cl) O. '.O N Q) C 0 a C U a) CL < 0 O E L. Cl) Q d 15 L Q V) Cl) 4-) (a C 0 -CI L rO L) (3 0 L R3 3 4r - E (O ]C undisturbed riparian zone . A nemourid stonefly was the only shredder found a t any of the impacted sites . Its food resource probably consists of CPOM burie d during the eruption and periodically re-excavated as the sediments move about . The dominant functional group of invertebrates at impacted sites is coile4tors . Their food source is also derived from buried organic matter as well as fro m sloughed periphyton . Predators were found only at sites that supported 'a fairl y high richness of other invertebrate taxa . Because predators are typicall y long-lived, witp life cycles of one or more years, habitat disruption may als o prevent their re-establishment in ,impacted stream reaches . The life cycles o f the dominant taxa (midges) found in the impacted streams are relatively shorter , with two or more generations per year . Apart from burrowing, another strategy enabling persistence in unstabl e sediments is exhibited by Baetis mayflies, which actively' swim about and utiliz e FPOM . This was the only species collected at Upper Maratta . Baetis spp . als o occurred in Ape Canyon and Upper Muddy . Taxonomic richness and abundance of the invertebrate fauna appear to b e fairly closely linked with the evaluation of channel stability of the stud y sites (Figure 2) . An exception to this relationship is exhibited by Upper ' Muddy, which has poor stability and a high invertebrate standing crop . No explanation is readily apparent for this anomaly . Perhaps an unidentifie d refugia exists nearby that provides a constant source of colonists . Changes in density (Figure .3) and in taxonomic'rich'ness (Figure 4) o f invertebrates in Ape Canyon since the 1980 eruption suggest that re establishment of fauna is a temporary phenomenon, and that a succession o f stable and non-stable cycles may occur in any given reach until the overal l landscape instability is lessened . Complete recovery of stream communite s affected by the eruption may be on a time scale of up to 1 0 2 years- or longer . Experimental Simulations of Disturbanc e Benthic Invertebrates : Two general trends were observed in the abundance along channels of tax a that appeared consistently in samples . One trend is that the abundance of som e taxa was much greater at the termination of the experiment in the unperturbe d channel (Channel II) than in the two disturbed channels (Figure 5a) . This tren d was exhibited by a number of taxa, occurring in different functional groups . 16 !I rI APE CANYON E r t 1 I 0 1 ! r 0 c 10 0 r I I I t I l 1 I 1 1 I l 1 1 ~ 11 I i 10 1 I I I I II I r I 1 0 1 • 1 0 5 10 1 1 15 20 Martine Since Eruption Fig . 3 . Macroinvertebrate density in Ape Canyon since the eruption (Ma y 1980) of Mt . St . Helens . Except for sample at 19 months, data ar e from Anderson and Hawkins (unpubl .) . 17 i% L • MI rp N ftj IC) - -CI Q) N w 0) I I O I • d •■- E E ni L C L) +3 QC N C 01 L0 2 )-2 Cal ItJ O. q S% 0 `■ / / rt Z R7 +Rf3 • r a c a) a) • . C CDE-C Crt1 NL +a) L) 40 RSU) q yC o. !Z (U +) fn c Cr C.)0X •. ip W N C • N }t I- n:s LC • Q) a - > a) r • •r L S U) > • C + .t ■ iOd M P0439(103 axD~ a S N utoao r~W o1p •oil g r L. a Cl) C +-) •rO C/) 3L. L• U +) N rG)0 E 4- 'C U 4.~ O O ▪ a) NN •rcO x - b b n E c ~ c o U L C ~-L O ON 4-)a) O O L U r t -csa +.1 E c a) OU E X c o f~f •r L N 4- I- Jr c3+ t 18 Abundance of Taxa (Control > Disturbed ) 6- asb Yoraper/a Plecoptera 600- °' b Ostracoda-C o Gr 2 200- He/icosama Gastropoda Gr b 3- Simuliidae-C o , Ecclisomyia Trichoptera Gr 15- 5- c . A//oiler/a Plecopter a Co 20 V a, b Perlodidae Plecopter a Pr b TurbeHaria-Pr 100- Channel ]l Fig . 5a . Abundance of macroinvertebrate taxa at Kalama Springs three month s after disturbance simulations, (groups for which control > disturbed) . Channels are ordered along an axis of increasin g disturbance frequency . Co = collector, Sh = shredder, Gr = graze r (scraper), Pr = predator . 19 Abundance of Taxa (Control > Mud > Most Disturbed ) Poro/eptop'i/ebia I5° Micrasema Ephemeroptera Co Trichopter a Sh 50 - 0 150- Harpactacoid a Co 300- a,b Hydracarino Acari- P r I00 - Sae Hs Ephemeroptera Co 2- 5i -Channel II Dicranota Tipulida e Pr 4 I II I Fig . 5b . Abundance of macroinvertebrate taxa at Kalama Springs three month s after disturbance simulations, (groups for which control > mod > disturbed) . Channels are ordered along an axis of increasin g disturbance frequency . Co = collector, Sh = shredder, Gr = graze r (scraper), Pr = predator . 20 a) L L ( O to U a) L r a) r a) C 4) C O 4- M17 U Rs s U N s o +) . L) C 0 "a a) 0 II . 010 N Q7 V C •r r a) LO U d L. C 6 rn i-a ms Cs ( Z W r i w ) ssowOig r0E OU co 4 ) r s v) b U •r •r L s d ) +) 3 a its L •rC ms o N L X ro 4- a)m aIZ) }) N L it) aU C i~ o •r r0 L. L 0)4 a) L U N 0 La) a) ~ •I•) ^ } L (J ) C b a) C a) L > 0 -,a •r I 4-3 rq L I O L r0 N 0 o (g L ~ u ~ 0 A w • /Jagwnu) ~(4isuaa a) CC v t U rtt E0rt3cD E v) a) L o a)U :co o0a )L U a) 0) a) 0 L a) V) L C4.3 a) 0 V) 0 J:) •r L. _a 11 <-0 Ov) U u) 21 Three of these taxa, the snail Helicosoma, Ostracods ; and the planari a Orotophela, were very abundant in the unperturbed channel, attaining densities . of 1800-6000 individuals m- 2 . In some instances, the abundances of taxa wer e also greater in the moderately perturbed channel (I) than in the frequently . perturbed ore (III) (Figure 5b) . Included here are the baetid .mayflies an d mites that are among the initial colonists- of heavily impacted streams- near~ .Mt . _ St . Helens . A second trend was exhibited by a few taxa in which abuncia'rce was greate r at the termination of the experiment in one or both of the disturbed channel s than in the unperturbed channel (Figure 5c) . This pattern may represent a •. release in numbers, either from the elimination of competitors o r , predators, o r an increase in resource availability that occurs in the presence o f non-catastrophic disturbance events . This trend occurred among the orthocla d midges, which largely dominated the community of the disturbed channels , reaching densities of up to 15,000 individuals per m 2 , but it is statisticall y significant only for biomass (p < .05, ANUVA) and not for density . A n unidentified orthoclad species occurred in the most frequently perturbed channe l that was fewer in number but ' much larger in size than the speci-es' inhabitin g other channels, and this accounts for the greater biomass observed in thi s channel . The clam Pisidiurnnever occurred in any sample taken from th e unperturbed channel . Total density, biomass, and diversity, of the invertebrate assemblages were - similar enough among channels prior to disturbance simulations to, make . u s confident that the differences we observed following .distu .rbance were attributable to the disturbance regime (Table 5) . Both density and 'diversity o f the assemblages after the initiation of the disturbance events were higher i n the channel never perturbed, and were somewhat greater in the channel mor e frequently perturbed, than in the more moderately perturbed . The diversity in the moderately perturbed channel (I) declined after one month of disturbanc e simulations, suggesting that one might observe in this channel a hyperboli c curve of species diversity over time, , similar to that observed in othe r successional communities (Connell and'Slatyer 1977) . A decline in di .rsity,•- • over time was not observed in channel III, probably because - distulbence s occurred too frequently to allow the colonization of competitiNily superio r taxa . Biomass following disturbance simulations was -remerkably - 4imilar amon g 22 .4 Table 5 . Community composition parameters of benthic invertebrates prior t o and after disturbance simulations at Kalama Springs . Values represent , means of six samples within each channel . Channe l II I II I Increasing Disturbance + Before Disturbance Density/0 .1 m 2 3043 346 4 378 .0 386 .9 453 .6 1 1317 .6 1451 .0 1832 .8 ' 2989 1343 178 7 Biomass/0 .1 m 2 M .U .I .* 2441 After Disturbance (1 mo . ) Density/0 .1 m 2 Biomass/0 .1 m 2 M .D .I .* 287 .8 333 .0 296 . 8 1447 .1 604 .3 240 .8 1 After Disturbance (3 mos . ) Density/0 .1 m 2 2800 Biomass/0 .1 m 2 M .D .I .* 1758 195 7 246 .4 238 .2 244 . 0 1483 .2 357 .2 240 .8 1 *M .D .I . = McIntosh Diversity Index = N- s E n i 2 i= 1 channels (Table 5) . This suggests that compensatory changes occur in strea m community structure that act to maintain a constancy, in at least some aspects o f community function . The structural resemblance of pairs of channels with each otaier wa s estimated by use of the Curtis percentage similarity measure, whi€h ranges from ' 0 to 100 . A value of 100 indicates complete similarity . Al .] channels wer e structurally similar prior to disturbance (Table 6) . After three months, channe l II, the unperturbed channel, was very different from both channels I and III . Similarity here was altered both because of the dominance of orthoclad midges i n the disturbed channels and because of the loss in these .channels of some sensitive taxa, such as the predatory perlodid stoneflies and the grazin g Invertebrate assemblages were partitioned into functional groups to enable an assessment and comparison among .channels of the trophic as well as taxonomi c structure . Functional group designations were assigned according to Merritt an d Cummins (1979) . The proportional biomass of shredders three months afte r 23 { Table 6 . Similarity of invertebrate assemblages among channels prior t o and three months after disturbance simulations at i(alama . Springs . PS* Before Disturbance After Lei s_turbanc e Channel s 1,2 .769 .28 8 2,3 .806 .42 7 1,3 .941 . . .835 *PS is the Curtis Percentage Similarity Measure . PS 200 E MIN ( N IH, N IK ) s (NHI + N KI ) -kJ disturbance was small in all channels (Figure 6) . This was expected, as th e resource base in the channels is largely autotrdphic . The collector component , largely comprised of midges, dominated the disturbed channels ., and a iargei, relative increase in the predator functional group was observed in .the . unperturbed channel . The proportional biomass of grazers (scrapers) in the unperturbed channel was also somewhat larger than in the disturbed channels . . The functional group structure of the Invertebrates colonizing heavily impacted , unstable streams near Mt . St . Helens is similar to that observed in th e experimentally disturbed streams . Scrapers and predators vau1r in low abundance in these streams, because the large and stable substrate require1-b y scrapers for attachement is absent, and the abundance and predictability of foo d resources required by both groups is low . Periphyton : The early colonizing species in all channels was predowimantly Diatoma hiemale var . mesodon, which forms a gelatinous matrix after one month . Successive algal taxa included the filamentous xanthophyte Tribonema and th e chrysophyte Hydrurus foetidus . Diatoma was most abundant in the mos t frequently perturbed channel . Chlorophyll-a levels of the periphyto n assemblages were highly variable within and among channels,_,ranging from 1 .3 t o over 400 Mg m- 2 . The standing crop of chlorophyll-a was generally greater, bu t not significantly so, in the unperturbed than in the perturbed channels . 24 N rO a N 0 MI -0 a) 0 L CT)H L r }1 r N C a) • r O C •r C 4-) (O N O N u) E 0 CO L. ~ .C a) EU > •r C N •r 1/) C 0 a) . 4- V a) O C a) rts 3 N .0 t!1 L O E •F) O N •r • r _o 'Q L a) r L > rO a) a ) C 4.) 04- •4 (O a) 4.) . LN L a.. 0 O. 0 C:Lo 25 IV . DISCUSSIO N A lower density and diversity of invertebrate . fa na, both taxonomically and 16 trophically, were observed in stream channels in wkiich disturbance events wer e simulated than in•an unperturbed channel . There is some indication, howe%sr• , that some community properties, notably biomass, remain constant among differen t . disturbance regimes . These findings suggest that non-catastrophic disturbances may not play a major role in determining stream community structure over ecological time . Non - r - ` catastrophic disturbances in many streams are fairly predictable events, an d species may have evolved behavioral and other adaptations to deal with them . Adaptations relevant to competitive situations, be it in usurping resources o r in withstanding attacks from natural enemies that might be necessary in th e absence of disturbance, may not have evolved . We are reluctant to generalize from our findings, however, to dismiss th e potentially large role that non-catastrophic disturbance may play in streams , because . a number of factors not considered in our experiment will also affec t the role disturbance plays . The spatial distribution of disturbance is one o f these . Effects of non-catastropic disturbance in most streams are patchy i n nature as some microhabitats are more immune to disruption than others, . A give n .disturbance regime produces a distribution of patch ages, and the effect on th e community is properly the summation of patch . effects . And, critically,_ the . exact effect of periodicity . of disturbance events is unresolved . Disturbance s icalo f at a rate of once per month, relative to the annual hydrographs ty p streams in the Pacific Northwest, a-re probably too frequent to allow more slowl y dispersing colonists to enter a community . Life history attributes and composition of the invertebrate fauna that wer e collected in streams severely impacted by the eruption of Mt . St . Helens ar e similar to those of the invertebrate fauna in the most frequently disturbe d channel at Kalama Springs . In both cases, communities are dominated by chironomid midges . Abundances, however, are greatly reduced in the stream s surrounding Mt . St . Helens, and evidence that the communities are undergoin g cycles of establishment and denudation in concert with changes in landscap e geomorphology indicates that catastrophes have different consequences on strea m communities from other disturbances, and should be distinguished from them . 21 Instability of channel banks and bottoms in streams severely impacted by the eruption appears to pose a major impediment to recovery of benthi c communities . Until these elements stabilize, communities will consist only of ' short-lived fugitive-type species that will be swept downstream with eac h successive' high flow event . Community recovery can perhaps be enhanced b y management practices that promote channel stability, such as . planting o f riparian vegetation or introduction of stable substrates into a stream . 4, 28 V . LITERATURE CITE D Connell, J . H . and R . O . Slatler . 1977 . Mechanisms of sucessian im natura l communities and their role in community stability and orgeization . Amer . Natur . 111 :1119-1144 . Gorman, O . T . and J . R . Karr . 1978 . Habitat structure and stream fi=s h communities . Ecology 59 :507-515 . Harper, J . L . 1977 . Population biology of plants . Academic Press, Lonalon. . Keller, E . A . and F . J . Swanson . 1979 . Effects of large organic material o n channel form and fluvial processes . Earth Surf . Processes 4 :3'61-3€30 . . Leopold, L . B ., M . G . Wolman, and J . P . Miller . 1964 . Fluviad processes in _ geomorphology . W . H . Freemanand Company, San Francisco . 522 p . Merritt, R . W . . and K . W . Cummins . 1978 . An introduction to the aq,l tie insect s of North America . Kendall/Hunt, Iowa . 441 p . Paine, R . T . 1979 . Disaster, catastrophe, and local persistence of the-se a palm Postelsia palmaeformis . Science 205 :685-687 . Pfankuch, D . J . 1975 . Stream reach inventory and channel stability dvairatiom . U .S .D .A . Forest Service, Northern Reg ion, 26 p . Rogers, L . E ., W . T . Hinds, and R . L . Buschbom . 1976 . A general weight vs . length relationship for insects . Ann . Entomol . Soc . Att . 69 :387-389 . . Swanson, F . J . and G . W . Lienkaemper . 1978 . Physical consequences . of larva organic debris in Pacific Northwest streams . U . S . Forest Serv . Gen . Tech . kept . PNW-69, Pac . N . W . .For . Range Exp . Sta . 12 p . Wetzel, R . G . and G . E . Likens . 1979 . Limnological analyses . W . a, Saunder s Company, Pa . 357 p . 29 w VI . APPENDICE S 1. Channel Stability Evaluatio n 2 . Invertebrate Taxa Collected From Stream Impacted B y Mt . St . Helens, winter 1981-82 . 31 ,, J . N m N m ( m 1 a +. O v C +. 4 C C 0 7 0 7 m 01 at O .4 . a --+ a O 3 7 0 V 4 m E L. a L. 0 (0 A a d '-. O ++ C C ' 0 w. 0C m u E a T O >. L m .' C > m 7 > O ++ CN. L 0 ^ . V O C r 0 .. a 0 0. 0 ¢ O O 4 Oi L T m •.-~ L. . Oti4r A 0 .- T .ay O a 17G uu 0 0 0 C' C H -a ! 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C J '•. V U .-. d 0 m .. '.. m '.+ 0 04 'C 0 00 'C a 0 . 0C > .-. Y 0C . O0 0 0 a0 0E •. u 1 C a • -. L O-O .. 0 E m V1 C v •. a U r O0)_ O 4 C 7 0 0 0 70 4 .. uW m . 0 O >, m c J U 32 4 0 c 7 ..0 . 0 . 0 01 C 3U. 1N "7 c ..C C a E VO •C L' C .00 0 m a C C .4. 7 •u 0 0 V m u0 1l >. • .0 C C 0 0 t -u. .0 u 0 -+ u V 0 co 0 0 0 •'. Cl . 0 '.. 0 .J A t ~d .T . 0. 0 m L C m a -. a 4 uO 4 u0 ni 7 t 00 u .4 7S 40 3 L 0 m U "C C .0 ~ u -C C ... -• .~ 0 t m 0 0 m C y7 O a C 4 E 0i a 0 a m 4 4 011 . . 4 0 C .-a 0 0 x O _a 0V 0 . - A. 0'0 ua w 0 C 0 C .. A u 0 T - to r O N N 00 0 V EC .0 O a 0 0 0 -4 C 4 O m O a 00 U C. al M. .. O. 4 d 7 CI O u IA t ..1 o U u C 1a O 7O 0u t i 6 0 . m .C 0 u o 00 0 0 G -. 0 CD > r r U ' t' . APPENDIX 2 . Invertebrate taxa collected in stream sites impacted by Mt . St . Helens, winter 1981-82 . S Lower Maratt a Hemipter a Trichocorixa spp . Trichopter a Pseudostenophylax sp . Di pter a Simulium sp . Empidida e Orthocladiinae spp . Diamesinae spp . Other : Naididae (Uligochaeta ) Ape Canyo n Ephemeropter a Baetis bicaudatu s Baetis tricaudatu s Plecopter a Shipsa sp . Dipter a Diamesinae spp . Upper Mudd y Ephemeropter a Baetis tricaudatu s Plecopter a Malenka sp . Megarcys sp . Trichopter a Rhyacophila sp . Dipter a Limnophora sp . Orthocladiinae spp . Kalama Rive r Ephemeropter a Ephemerella coloradensi s Drunella sp . Epeorus longimanu s Plecopter a Malenka sp . _ Megarcys sp . Perlodida e Trichopter a Lepidostoma quercin a Hydatophylax sp . Pseudostenophylas sp . ,Micrasema sp . Neotheremma sp . Rhyacophila sp . ;'Diptera Dicranota sp . undetermined Tipulida e Twinnia sp . Orthocladiinae spp . Empididae sp . Other : Naididae (Oligochaeta ) Upper Maratt a Ephemeropter a Baetis tricaudatu s $~ Swift Cree k Dipter a Orthocladiinae spp . Diamesinae spp . undetermined Tipulida e Other : Acari (exuvia ) centipede (terrestrial ) Carbonate Spring s Dipter a Orthocladiinae spp . Diamesinae spp . f +~. $ .' 33