AN ABSTRACT OF THE THESIS OF in Geoffrey Lynn Dorsey for the degree of Master of Science Fisheries and Wildlife presented on June 6, 1982 Title: An Analysis of Avian Coimnunities on Three Islands, Lower Columbia River, Oregon Abstract approved: Redacted for privacy Dj John A. Crawfo, major professor Avian communities were described for t habitats (marsh, upland, beach, and tree-shrub) on 3 islands (West Sand, Miller Sands, and Jim Crow Sands) in the Lower Columbia River. June 1978 through May 1979. Research was conducted from The variable circular-plot census method was used to determine avian commmunity characteristics (number of species, density, H', and J'). Percent cover of various habitat characteristics were measured to determine relationships with avian community characteristics. Marsh habitats differed in number of species observed and vegetative cover. Increased number of avian species was associated with increased vegetative cover for marsh habitats. Shorebird usage of marshes was related to the incidence of bare ground. Upland habitats were similar with regard to avian community characteristics; however, a marked difference in species composition was observed for barren versus vegetated upland habitats. Tree-shrub habitats were not different with respect to avian community charac- teristics although the transition habitat was markedly different with regard to structural complexity. Beach habitats were similar in avian and habitat characteristics. Comparisons of avian community charac teristics among habitats grouped by island indicated that the most successionally advanced and structurally complex habitats tended to differ from less advanced and less complex habitats. An Analysis of Avian Communities on Three islands, Lower Columbia River, Oregon by Geoffrey L. Dorsey A Thesis submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science June 6, 1982 APPROVED: Redacted for privacy essor of Wildli Ecolo- in chfilge of major Redacted for privacy Head of Department bf Fisheries and Wildlife Redacted for privacy an of Graduate Date thesis is presented Typed by LaVon Mauer for January 7, 1982 Geoffrey Lynn Dorsey ACKNOWLEDGEMENTS Many individuals have contributed to the development of this thesis and to each I am grateful. Initially, C. Wayne Eshelinan took an interest of mine and nurtured a desire to learn. and gratitude to him continue to this day. took the baton as my major professor. My admiration for Dr. John A. Crawford then I-U.s guidance, support, insight, and dogged determination have brought me to this point where further attainment lies fully with me. I truly appreciate the leadership he has provided me. Many fellow graduate students provided assistance in various aspects of this research. Russ Oates, Dan Edwards, Dirk Van Vuren, Mike Morrison, and Cecilia Noyes are just a few to whom I owe thanks. Dan helped me initiate this study; Russ provided advice and support; Dirk edited one draft and discussed various aspects of the research and results; and Cece and Mike provided invaluable assistance on statistical analysis. Larry Morrison taught me field identification of plants on my study islands. Phil Hamm assisted me in the field and lent moral suport, but more importantly has been a friend. Eric Beals often led me through the morass of coinputerdom. The Department of Fisheries and Wildlife faculty have provided advise, discussion, and an education. this thesis. period. Dr. Robert Anthony has reviewed Dr. Richard Tubb provided monetary support at a critical Thanks seems inadequate but I am truly thankful for their contributions and hope one day I can repay them in full. The U.S. Army Corps of Engineers, Portland District provided funding and provided assistance and direction throughout the research. Dr. Thotnas Morse faciitated the initiation of this research and pro- vided frequent advice and support throughout the research. What words appear on this page and thoughout this thesis are due in one sense, to only one individual--LaVon Mauer. Her assistance, innovativeness and persistence are appreciated. Finally, I owe a great deal of thanks and love to my wife, Joyce, for her patience, understanding, and tolerance of my frequent absence from home. My parents were also supportive of this endeavor and I thank them. My sincere thanks to all who have contributed in any manner to this research. TABLE OF CONTENTS Page INTRODUCTION I STUDY AREA 5 METHODS 9 RESULTS 14 Marsh 15 Upland 28 Tree-Shrub 36 Beach 43 West Sand Island 43 Miller Sands 50 Jim Crow Sands 52 CONCLUSION 52 MANAGNENENT RECOMMENDATIONS 52 LITERATURE CITED 54 APPENDICES 62 LIST OF FIGURES Page Figure 1. Habitat locations for West Sand Island, 7 Columbia River, Oregon. 2 Habitat locations for Miller Sands Island, 8 Columbia River, Oregon. 3 Habitat locations for Jim Crow Sands Island, Columbia River, Oregon. 10 LIST OF TABLES Page Table 1 Number of species, and avian density, diversity, and 16 evenness values by season for each habitat group, Lower Columbia River, 1978-79. 2 Discriminant function results (avian community 18 characteristics) for habitat groups for which habitat and season effects were determined significant by multivariate analysis of variance. 3 Habitat comparisons of species composition values 19 (MacArthur's Difference) for similar habitat groups, Lower Columbia River, 1978-79. 4 Seasonal comparisons of species composition values 20 (MacArthur's Difference) for combined similar habitats, Lower Columbia River, 1978-79. 5 Percent cover of litter, forbs, and grass/sedge 21 for marsh habitats, Lower Columbia River, 1978-79. 6 Results of discriininant function analysis (habitat 23 characteristics) for habitat and season effects determined significant by multivariate analysis of variance. 7 Percent cover for each habitat characteristic, West 30 Sand Island, Miller Sands, and Jim Crow Sands uplands. 8 Percent cover of tail grass/sedge ( 15 cm) for 32 MSU, WSIDU, and WSIU, Lower Columbia River, 1978-79. 9 Diversity indices for "edge" of individual habitats, Lower Columbia River, 1978-79. 34 Table 10 Pa&e Percent cover values for each habitat characteristic, 38 tree-shrub habitats, Lower Columbia River, 1978-79. 11 Percent occurence of bare ground for 4 beach 44 habitats: MSSB, MSB, JCSB, and WSIB, Lower Columbia River, 1978-79. 12 Mean values of each avian community characteristic 46 for habitats comprising each island, Lower Columbia River, 1978-79. 13 Habitat comparisons of species composition values 48 (MacArthur's Difference) for habitats grouped by island, Lower Columbia River, 1978-79. 14 Seasonal species composition values (MacArthur's Difference) for combined island habitats, Lower Columbia River, 1978-79. 49 An Analysis of Avian Communities on Three Islands, Lover Columbia River, Oregon INTRODUCT ION Previous investigations of dredged material islands were principally directed toward determining plant succession (Barnes 1971, Coastal Zone Resource Corp. 1977, Dunstan and Lewis 1974), nesting by colonial species of waterbirds (Buckley and Buckley 1975, Buckley and McCaffrey 1978, Chaney et al. 1978, DuPue 1974, Landin 1978, Morrison and Shanley 1978, Olsen 1975, Peters et al. 1978, Scharf 1978, Soots and Parnell 1975, and Thompson and Landin 1978) or animal colonization (Cammen et al. 1974). Several investigations of avian communities have been conducted in the Columbia River estuary. Woodward-Clyde Consultants (1978) investigated both floral and faunal aspects of Miller Sands between 23 April and 24 June 1975 and determined that bird species diversity was highest in the tree-shrub association and lowest in upland and sandspit habitats. Crawford and Edwards (1978) conducted research on the responses of bird, mammal, and macroinvertebrate populations to plantings on Miller Sands between June 1976 and August 1977. Avian density and diversity were greatest in the tree-shrub habitat; number of species was highest in the beach and marsh habitats (Crawford and Edwards 1978). Number and type of avian species present were occa- sionally altered by plantings, but density and diversity were usually unchanged (Crawford and Edwards 1978). Edwards (1979) determined that avian community characteristics (density, number of species, diversity 2 (H'), evenness (J') and standing crop bioinass) associated with upland and tree-shrub habitats on Miller Sands were significantly different. Avian community characteristics of upland and tree-shrub habitats on Miller Sands were significantly different from marsh and beach habitats (Edwards 1979). Seasonal differences also occurred among avian communities (Edwards 1979). Avian community characteristics for spring differed from summer (1976), fall, and winter and those for summer (1977) differed from fall and winter. Avian community charac- teristics and species composition were influenced by season, vegetative complexity, and the physical attributes associated with each individual habitat (Edwards 1979). The variation in avian species composition, density, and diversity with respect to vegetative communities observed by investigators on Miller Sands also may be a function of the soil moisture gradient. Investigations conducted on Mott Island, Columbia River, Oregon, by the Coastal Zone Resources Committee (1977) revealed that soil moisture gradient, a function of topography and tidal influence, iiad an important role in determination of vegetative communities. Plantings also influenced the vegetative communities on ?lott Island (Coastal Zone Resources Committee 1977). Factors affecting vegetative succession on dredged material islands in Florida included climate, substrate, wave action, nearness of plant sources, and human use (Schreiber and Schreiber 1978). Scharf (1978) found little correla- tion between age of islands and succession of vegetation on islands in the Great Lakes. He ascribed this lack of a relationship to variation in plant succession and parent dredged material and the influence of 3 Retardation of succession on some dredged material islands in ice.. the Great Lakes was attributed to parent material, bird usage effects, erosion, and/or inundation (Scharf 1978). Yeaton and Cody (1974) reported that the principal effect of the addition of canopy cover was the addition of more bird species to the community. Edington and Edington (1972) noted that different species forage in different vertical strata, thus the more vertical stratifi- cation, the tre species opportunities (see also MacArthur 1958, Morse 1970). Avian density generally increases with succession (Shugart et al. 1975, Kendeigh 1948, Odum 1950, Johnston and Odum 1966, Haapanen 1965, Karr 1968, Karr and Roth 1971, and Shugart and James 1973). Adams (1908) and Bond (1957) noted that species richness was higher in intermediate successional stages. Meents (1979) stated that addition of a shrub layer led to greater species richness. McElveen (1977) reported that species density and avian density was greater in edge situations than for either vegetative community forming the ecotone. Stauffer and Best (1980) reported an increase in mean densities of breeding birds from herbaceous habitats (uplands-this study) to upland woodlands to floodplain woodlands. Karr (1968) observed a general increase in number of species, density, and diversity from bare ground to bottomland forest habitats on former strip mine sites. Wilison (1974) determined that increased environmental patchiness, specif i- cally the addition of trees, strongly influenced the addition of avian species to a habitat. The increase in number of species was greatest when formation of a tree layer was initiated. Des Granges (1980) stated that equitability depended on physiognomic diversity of a 4 forest stand; high equitability was associated with more complex stands. An inventory of riparian habitats and associated wildlife for the Columbia River was conducted by the Oregon Cooperative Wildlife Research Unit (OCWRu) (1976). Investigations were conducted in habi- tats similar to and in the vicinity of present study islands; however habitats on dredged material islands were not investigated. West Sand Is land was included in Segment 1; Miller Sands and Jim Crow Sands were included in Segment 2 of the study. Alder communities in Segment 1 possessed the highest number of species, avian density, and avian biomass (OCWRU 1976). Seasonal variation in maximum numbers of birds and densities were observed in Segment 2 (0CwRU 1976). Sitka spruce (Picea sitchensis) habitat contained the greatest number of species during all seasons except spring; cottonwood communities contained the highest number of species during spring. Avian density was greatest during fall and winter in Sitka spruce habitats. Cottonwood/willow communities were represented by the highest avian density during spring. Avian density was highest during summer in cottonwood coni- munities. Since habitats on these islands are potentially subject to drastic changes in structure from dredged material deposition, this study was initiated. Habitats that varied in vegetative complexity were investigated. The purpose of this research was to describe avian com- munity and habitat characteristics associated with 4 gross habitats (marsh, upland, beach, and tree-shrub) on 3 islands (Miller Sands, Jim Crow Sands, and West Sand) representing varying successional stages 5 and determine relationships between avian community and habitat characteristics among similar habitats and for island habitat complexes. STUDY AREA The study islands, West Sand, Miller Sands, and Jim Crow Sands, were chosen for investigation because they are current disposal sites for dredged material. Each island was typified by a complex of habi- tats (marsh, upland, beach, and tree-shrub). Ages of specific habi- tats varied among islands and were related to date of formation from dredged material deposition. The climate of the Lower Columbia River is typified by wet winters and dry summers (U.S. Army Corps of Engineers, Portland District, 1975). Astoria, located centrally among study sites received 143.8 cm of precipitation with average maximum and minimum temperatures of 14.7 C and 5.8 C, respectively, for the period June 1978-May 1979 (National Oceanic and Atmospheric Administration 1978, 1979). Severe cold weather from December until early February resulted in the freezing of surface layers of intertidal marshes and mudflats and a virtually complete blockage of the main river by ice floes during two periods. West Sand Island (257 ha) was located in Baker Bay, east of Cape Disappointment, Washington. Formally a natural bar, West Sand Island was shif ted to its present position by channelization activities that occurred at the mouth of the Columbia River (Chief of Engineers, U.S. Army 1937; Oregon Historial Society 1980). Dredged material deposi- tion altered and added habitats to West Sand Island (e.g. diked upland). Avian censusing stations were established in 6 habitats on West Sand Island (Fig. 1): marsh (WSIM), diked upland (WSIDtJ), upland (WSIU), beach (WSIB), transition (WSIT), and tree-shrub (WSITS). Grass/sedge, principally Carex sp. and creeping bentgrass (Agrotstis alba), was the dominant ground cover of WSIM. Red fescue (Festuca rubra) and tall fescue (F. elatior) were the dominant species forming WSIDIJ ground cover. Red fescue, seashore lupine (Lupinus littoralis), coast strawberry (Fragaria chiloensis), and black knotweed (Polygonum paronychia) characterized WSIU. of WSIB. Bare ground was the dominant feature Carex sp. was the dominant plant species in WSIT. The overstory of WSITS was composed of red alder (AJ.nus rubra) and willow (Salix sp.). Willow, twinberry (Lonicera involucrata), and salmon- berry (Rubus spectabilis) were the dominant understory shrubs. Litter and grass/sedge, principally Carex sp., comprised the ground cover. Miller Sands (228 ha), located between river miles 22 and 25, was formed in 1932 by deposition of dredged material (Edwards 1979). Creation of a spit began in 1948 and was completed in 1976. Presently, deposition is limited to the spit area on Miller Sands. Censusing on Miller Sands was conducted in 6 habitats (Fig. 2): marsh (MSN), spit upland (MSSU), upland (MSTJ), spit beach (MSSB), beach (MSB), and tree-shrub (MSTS). Carex sp. and bare ground typified MSM during the summer. MSSU was characterized by bare ground. Moss, f orbs, litter, and grass/sedge constituted the primary ground cover of MSU. MSSB and MSB were devoid of vegetation. Red alder, cottonwood (Populus trichocarpa), and willow were the main constituents of NSTS overstory. Salmonberry and elderberry (Sambucus raceimosa) were the Figure 1. Habitat locations for West Sand Island, Lower Columbia River, Oregon. 7 I caie - 1 Figure 2. . lEO Habitat locations for Miller Sands Island, Lower Columbia River, Oregon. most abundant understory shrubs. Ground cover of MSTS was typified by litter, forbs, and grass/sedge. Jim Crow Sands (86 ha), also formed from dredged material, was located between river miles 26 and 28. Deposition has occurred on Jim Crow Sands since at least 1957 (U.S. Army Corp. of Engineers Pipeline Disposal History, Portland District 1980). sused on Jim Crow Sands (Pig. 3): beach (JCSB). Three habitats iere cen- marsh (JCSM), upland (JCSU), and Deposition on the eastern portion of JCSU had occurred just prior to study initiation, hence, regarding succession, the eastern portion of JCSU was chronologically younger than the western segment. Bare ground was the prominent feature of the west and east ends of JCSU. Grass and forb plantings during September 1978 on the recent deposition site markedly reduced the amount of bare ground. Bare ground typified JCSB. Bare ground and grass/sedge were the domi- nant features of JCSM. METHODS The variable circular-plot method (Reynolds at al. 1980) was used for avian sampling. This method was selected because variations in detectability among species of birds and among seasons and habitats for a species is accounted for when calculating species densities, A total of 15 habitats on 3 islands were sampled monthly from June 1978 through May 1979. Two habitats were sampled each day and 10 sta- tions were censused in each habitat. at each station. Data were recorded for 10 nun. Birds, detected visually and/or aurally within the boundaries of a habitat and under 40 m in altitude, were recorded by j ficale - 1 cm 103 m New upland Marsh Old Uplflfl(i Figure 3. Habitat locations for Jim Crow Sands Island, Lower Columbia River, Oregon. 11 species and distance from observer for each individual or group of individuals. Density estimates were based on an effective detection distance determined for each species (Reynolds et al. 1980). recorded in concentric 10 in Birds were bands from the observer to 150 in, in 25 bands from 151 in to 200 in, and in 100 in in bands for greater distances. Numbers of individuals for each band were recorded for each habitat. Similar habitats (e.g. MSM, JCSM, WSIM) were combined to increase sample size for determination of the effective detection distance. Seasonally, the effective detection distance and subsequent species densities were calculated by the method of Reynolds et al. (1980). The percentage of paired singing males was not determined, thus no correction factor for avian density during the breeding season was utilized. Species density estimates for each habitat were determined from the formula: D = (10,000 m2ha)N = birds/ha; where x the xir- number of stations sampled (10); r the radius of the circle deter- mined by the inflection point; and N is the number of birds detected within the radius of the circle. yielded total Summation of species densities avian density for each habitat. Densities for groups of birds (e.g. lands, hirundinids) were apportioned according to the density determined for individuals composing the group or by proportion of individuals observed during incidental counts. Diversity indices (Shannon and Weaver 1949) were calculated from the formula H' = in the ith species. p. logp; where p = the proportion of individuals Where density estimates for groups (e.g. lands, hirundinids) occurred for a habitat, p. was calculated only from those 12 densities associated with identified species. Evenness (J') was determined by dividing H' by the logo of the number of species observed in each habitat. Species composition among similar habitats and between seasons for individual and combined similar habitats was examined with MacArthur's (1965) difference measure: ik e (lit-H) p.,+p; where lo difference. ih of the ith species of Hh and HK (where H, for the first census and and ik = equal the proportion equals the diversity index for the second). Differences ranged from 1.0 to 2.0; a value of 2.0 denotes 2 communities with no species in common. Coinniunities are more similar as the difference value approaches 1.0. Vegetation sampling was conducted seasonally. Habitats were characterized by 80 0.25u2 quadrats (1 m2 for beach habitats and uplands devoid of vegetation); 2 quadrats were randomly distributed along lines in each cardinal direction at each station. Habitat characteristics evaluated were percent cover of moss/lichen, rock/litter, bare ground, grass/sedge, and f orbs. Additionally, per- cent overhead cover, canopy height, canopy base height, and circumference of trees were determined for MSTS, WSIT, and WSITS. Canopy and canopy base height, indices of foliage height diversity, were determined for these habitats by determining the number of individuals (p.) in each 1 ni increment above 2 in. The Shannon-Weaver (1949) for- mula for diversity was again used for calculation. Percent overhead cover was determined by visually estimating the area of a leveled 13 Brunton Pocket Transit mirror obstructed by overhead cover (leaves, branches, and/or stems of trees). This characteristic was nasured seasonally at 8 randomly selected sites for each avian census station; 2 sites were randomly located along a line extending in each cardinal direction from the census station; with 4 measurements taken at each site. Habitat edge effect was quantified based on the formula deve- loped by Patton (1975): DI TP where DI = diversity index, TP 2I total perimeter (meters), and A equals area (meters squared). Multivariate analysis of variance (SPSS-MANOVA) (Cohen and Burns 1976) was used to test for differences, with respect to the avian community and vegetative characteristics among: habitats; 2) habitats grouped by island; and 1) combined similar 3) seasons. Correlation analysis was utilized to eliminate those variables which exhibited a high intercorrelation. Interaction (season by habitat), habitat, and season were the main effects tested by the multivariate procedure. A significant interaction effect normally precludes further analysis of main effects (Kim and Kohout l975)e of Statistics OSU, pers. Ramsay (Dept comm. 1980) indicated that if a biological basis for the interaction effect was determined, then further analysis of the main effects (habitat and season) was not necessarily precluded. Significant differences (P 0.05) detected by the SPSS-MANOVA procedure (Wilk's lambda value) were further analyzed with the Hotelling T-squared statistic (Morrison 1967) to determine which habitats or seasons were significantly different from one another. Univariate analysis of variance, a subunit of the SPSS-MANOVA 14 statistical package, was used to gain an indication of the specific parameter(s) which led to significant differences among habitat cornparisons. Inadequate sample size obviated separation of seasons for avian community data. Discriminant function analysis (Kiecka 1975) was used for separation of habitats and seasons by avian and vegetative characteristics when MMOVA indicated a significant difference. Discrim.inant function analysis provided an indication as to which characteristic(s) was most important for group (habitat or season) separation. Canonical corre- lation Cr) was utilized as a measure of the degree of group separation for each discriniinant function. Canonical correlations that exceeded 0.70 were considered strong indicators of habitat differences. 0.15 was used for determination of signif 1- An alpha level of P cance among multiple comparisons unless otherwise specified. For multiple comparison tests alpha (0.15) was divided by the total number of possible comparisons. This procedure established the actual alpha level for entry into the appropriate F-table and thus determination of significance for comparisons. Hence, fifteen possible comparisons existed among the 6 habitats present on Miller Sands; therefore, x 0.15/15 0.01, the alpha level with associated degrees of freedom for critical F-values. RESULTS AND DISCUSSION Results are separated on the basis of major habitat groups (marsh, upland, beach, and tree-shrub) and islands (West Sand, Miller Sands, Jim Crow Sands). Habitats were grouped on the basis of similarities 15 in vegetation, substrate, tidal influence, and general structure. Combination of similar habitats based on these features was conducted to determine if subtle differences influenced avian community characteristics and composition. Grouping habitats by island allowed com- parison of various habitats. MARSH Among marsh habitats, JCSM:WSIM was the only comparison for which significance was determined (Table 1) and number of species was the only characteristic that differed (P 0.05). The observed difference in number of species for JCSM:WSIM comparison was consistent for all seasons except winter when similar counts of species occurred (Table 1). Discrimination among marsh habitats also was accomplished on the basis of number of species; density was of secondary importance (Table 2). Based on this separation, JCSM was least similar to WSIM, and MSM was intermediate to those habitats. The first discrimiaant function was not a strong separator of marsh habitats (rc = 0.55). composition of all marsh habitats was similar (Table 3). Species Seasonally, the contrast in species composition was greatest for the suminer:winter comparison (Table 4). Analysis of marsh habitat characteristics indicated a significant season by habitat interaction effect (P < 0.05). This effect was attributed to the paucity of forbs in JCSM which obscured seasonal changes in f orb cover as observed for MSM and WSIM (Table 5). Comparisons among marsh habitats and habitat characteristics asso- ciated with each indicated that all comparisons were significant Thbk 1. Nwtr of bpcie. a1 twlan dentty, dirfty aiii eveoiiw va1ic,i ti ucu br edi AvLn Cotiiuntty Nwgeb of 11thitt grwp ctes FLIII Ithter SprirE 12.3 10.7 10.7 12.0 11.4 NfH 12.3 17.3 11.0 14.3 WSIM 16.0 17.0 11.3 16.3 c,4 .iL 1trutby Spr1r I4asi Siaeer hilt WInt.r 3.4 18.7 12.5 Lit 1.50 [.52 9.9 1.4 11.1 1.2 1.72 1.88 5.5 6.6 6.2 10.4 1.2 135 l'aU WInter 8.9 18.8 13.7 5.9 15.2 *u grwp, 0r (1uthta K1ur 19il$-1J. rjcL0rI1t1C (b1nW1i) urIwr JCSH lkl Suwor btan iieer hiLt Witer 0.94 1.42 0.69 0.t3 0.05 039 0.59 1.69 1.66 1.79 0.00 0.00 0.79 0.58 u.00 1.87 1.00 1.82 1.01 0.03 0.00 0.41 0.0/ 0.59 Spd r1rE I*iiwi Up1as 6.0 8.3k 8.5 8.2 2.3 3.6 6.6k 5.2 1.40 1.82 0.88 138 1.31 0.58 0.85 o.57 0.tE5 43.66 WSIW 10.0 11.3 1.3 8.0 9.2 43.0 4.0 1.9 4.2 5.8 138 1.49 0.81 1.29 1.24 0.64) 0.61 0.41 0.liS 0.51 J4J 10.7 6.3 3.3 [4.3 83 4.8 4.8 0.8 1.9 4.6 1.40 0.73 43.34 2.01 1.12 0.59 0.41 0.22 0.19 0.50 MS9l 9.1 7.3 7.0 9.0 8.3 17.4 6,9 Li 18.1 10.9 1.21 1.11 1.58 1,38 1.25 0.9 0.56 0.14 0.64 0.64 WSIU 16.3 1.7 5.0 12.3 10.3 8.1 1.4 0.5 4.7 3.6 1.56 1.61) 0.448 1.66 1.43 0.50 0.19 u.62 43.00 0.06 12.3 9.7 8.7 11.7 10.6 19.8 12.9 1.8 19.6 13.5 1.23 1.41 1.36 1.4/ 130 0.49 0.0/ 0.63 0.61 0.64) )H 10.3 9.0 11.0 15.0 11.3 6.1 2.3 2.6 17.0 1.0 4.70 1.37 1.78 1.53 1.62 0.79 0.04 0.10 0.5/ 0.60 JtB 14.3 43.3 8.3 14.0 11.5 15.5 7.5 /.4 135.6 41.5 [.44 1.60 1.02 1.52 1.40 0.60 u.02 0.48 0.u4 0.38 4I8 40.7 13.0 10.7 10.3 41.2 12.2 9.0 4.6 7.6 44.4 1.4o 1.49 1.53 1.59 1.52 0.02 43.56 u.o 0.69 0.64 beade flhl1a 1. (conttiud) Avtai ccu.auilty cractertatic IR1tya Nnxibar at Hahttat Spthi haii Suiaiar Fall Wlaer Sr10 t*an Sua&at hiLl Whiter 5irui a.ia 9.1 10.1 12.5 1.68 1.75 1.90 2.03 1.84 0.10 0.70 0.18 0.79 0.)h$ 4.8 10.0 14.0 10.4 2.28 1.42 1.67 2.24 1.90 0.88 0.72 0.81 0.81 0.80 6.5 9.9 14.3 12.3 2.32 1.52 1.94 1.93 1.93 0.80 0.68 0.744 0.72 0.15 t$aU &.aler t1t Wthtec 13.0 10.9 6.0 24.1 8.0 16.3 11.6 12.9 12.3 15.0 14.0 18.6 Fall Wtnter 7.0 12.0 11.1 tvr 14.7 7.3 %flS 48.3 40.3 &aaer grwp (bt.1/I&a) pedeia rtng ?rae-uub g data re trawfonaid (aquare coot) print to uaiywl.w. wraity data were not Included in analyaia at upland habluata becanaie at ttai htJl correlation (r C8k aerttca1 lbnca j68iete tUcant dii nteacca (ii 0.440) a.iLUa e'&nncea. 0.05); cwaccted ltncs indicate no ditietence. unlinrlaie uI t'Lences uuors lasbitata. deticai ltraia tot habitat groop represent oulihurtate dl erencea; veLticat 1laea tar ascii asian ccusuutty tharaotetiUc repnn.ent was algnittcan4' dttfetent F SprlIUJ was a1gn1f1cant 0.025) iron fall aiai winter with roapect to najiber at species and dauaily. diffcrent F 4 0.025) fran winter with respect Lu tuiter of spectai density, and evenness. further tests weca coaui,ctu.t it tailtiwariate analysts tcvealad no slit1icant dittarences. I- Table Dtscriiainant functinu results (avian coaaunity characteristics) for habitat groups for which habitat 2. and season effects were deteri1ned signiticant by multtvariat analysis o variance. Effect tested and habitat group Kotated atandardted discriminant function coefficients (each parameter) biscrituinant function Percent of variance correlation Species 1)enaity Diversity 8veuness 1* 78.4 21.6 0.55 0.32 -1.39 -0.67 1.05 0.43 0.54 1.34 --- 72.3 26.1 0.48 0.31 -0.45 -0.32 1.24 0.35 --- 0.12 1.08 2* 56.0 39.3 0.62 0.56 1.40 0.52 -0.54 0.38 0.39 1.23 --- 0.69 0.48 -0.51 -0.12 1.33 0.06 --- 0.00 2* 65.0 21.5 1* 92.0 0.69 0.27 -0.13 1.08 0.16 1.06 -0.27 0.73 0.35 0.21 1.26 -0.27 -0.48 ----- Canonical HABITAT Harsh 2 Upland j* 2 Miller Sands 1* West Saud Island 1.00 SEASON Upland 2 Tree-Shrub 8.0 1* 88.0 2 10.9 *Significant discriminatory information existed prior to formation 1 discritainant function (P -0.11 1.04 0.04 0.05). 19 Table 3. Habitat comparisons of species composition valuesa (MacArthur's Difference) for similar habitat groups, Lower Columbia River, 19 78-79. Habitat Group Marsh Habitats MSM JCSM JCSN 1.22 WSIM 1.33 1.24 Upland MSSU NSU JCSU MSU 1.45 JCSU 1.46 1.39 WSIDU 1.75 1.44 1.60 WSIU 1.62 1.34 1.34 Beach MSSB MSB JCSB MSB 1.20 JCSB 1.19 1.23 WSIB 1.41 1.40 MSTS WSITS Tree-Shrub WSITS 1.15 WSIT 1.22 aValoes range from 1.0 to 2.0. WSIDU 1.37 1.44 1.13 Differences in species composition among comparisons are greater as 2.0 is approached. 20 Table 4. Seasonal comparisons of species composition values8 (MacArthur's Difference) for combined similar habitats, Lower Columbia River, 1978-79. Season Habitat group Summer Fall Winter Marshes Fall 1.45 Winter 1.71 1.27 Spring 1.36 1.24 1.26 Uplands Fall 1.44 Winter 1.69 1.31 Spring 1.23 1.34 1.42 Beaches Fall 1.42 Winter 1.74 1.27 Spring 1.32 1.32 1.32 Tree-Shrub Fall 1.24 Winter 1.33 1.14 Spring 1.13 1.32 aValues range from 1,0 to 2.0. 1.33 Differences in species composition among comparisons are greater as Z.0 is approached. 3.9 10.5 32.8 6.6 17.3 49.5 8.7 46.5 56.4 2.9 3.2 8.1 JCSM HSM WSU4 Spr1ng Winter4 50.5 73.5 2.4 2,3 1.1) 0.2 0.0 0.0 1.7 0.2 7.1 8./ 36.8 linea indicate atguiticant difference. were tranaforwed (wreath /11) for analywia, 21.11 19.4 O.l 0.1 0.0 5.5( 0.1 SueaerC 0.1 Mean gpringt Winter iail Suaar' Mean Forb ercent cover of forha cad grawe/aedge doring apring ditfered frow winter. cover of litter loting apring differed true tall. 4Vercent cover of litter, lorba, and graaa/wedge during winter d1fferd from tall. CVerceot cover of litter, forb, and graae/wedge during anawer differed true tail, winter and apriug. b8k liabttwt Fall Litter 24.1 8.2 1.4 Fail t*ercent coverU of litter, torba, and graau/aed6e for i&irai habttata, Lower Coluwbia Kivr, 19111-79. Suiuwer Table 5. 4.5 1.2 J.2 Winter 2.o 9.9 2.9 priug 11.4 Heau 22 0.03). (P Percent cover of litter and grass/sedge were the prin- cipal habitat characteristics that separated marsh habitats (Table 6). A continuum from low (JCSM) to high (WSIM) percent cover of litter and grass/sedge was developed on this basis. A high degree of separation (rc = 0.79) existed among marsh habitats based on the first disciminat ion. Identical placement of marsh habitats occurred for ordinations developed on the basis of avian community and habitat characteristics Jefferson (1977) stated that plant distribution in salt marshes was related to elevation. She developed a classification scheme wherein salt marshes were assigned to six categories: low silt marsh, 3) sedge marsh, immature high marsh, and 1) low sand marsh, 4) bulrush and sedge marsh, 6) mature high marsh. 2) 5) Although JCSM and MSM are freshwater marshes, characteristics of these marshes are very similar to the low silt (2) and bulrush and sedge (4) marsh categories, respectively. West Sand Island marsh, a saltwater marsh, appeared most similar to a mature high marsh (6), which Jefferson (1977) considered a climax situation. Thus, a successional gradient from JCSM to WSIM apparently existed and coincided with the ordination of marsh habitats on the basis of number of avian species and the habitat parameters, litter and grass/sedge. The increased elevation and concomittant increased occurrence and coverage of vegetation associated with WSIM compared to JCSM appeared influential with regard to number of species of birds. WSIM had a greater amount of edge than JCSN (DI respectively). Additionally, 3.93 and 2.58, Habitats abutting WSIM were open water, upland, and Table 6. kesulta of d1ecrIutnLInt fujicttod analysis (habItat characteitsttce) for hb1tat anU season etfecte teretuen at ttitaut by tivartate analysis of variance. hoIsted taiidardize4 diacrisinaut iuncctou coefttclenta (acU parseeter) lifted tested Discriiainant function Percent of variance coirelation 94.0 0.79 - 0.94 2* 6.0 0.3! -- 1* 60.0 0.95 2* 29.0 1* Canouicsl t4oss/ Hchcu hock/ titter bare ground Iass/ 1ercent overhead IOLbS sedge cover 0.21 0.75 - 0.10 --- 1.25 -0.18 -- 1.15 0.111 0.37 0.24 (1.53 0.91 0.20 0.30 1.2? 0.24 0.31 --- 79.7 0.92 0.26 0.19 -0.43 -0.17 1.04 2* 20.3 0.77 0.64 1.43 --- 0.50 1.99 -0.04 1* 91.6 0.98 1.05 0.12 0.14 0.14 2* 6.6 0.79 0.08 1.06 -- 0.19 0.09 --- 1* 87.8 0.77 0.34 0.55 -0.10 0.10 0.75 2* 12.2 0.41 1.01 1.20 2.15 -0.36 1.91 -- 1* 71.6 0.95 0.08 0.12 2.12 0.17 0.15 2* 16.0 0.82 0.66 4.01 2.66 1.09 3.65 --- tlabt tat Marsh Upland Tree-Shrub Miller Sands Jiia Crow Sands West Saud Island 1 I Table 6. (contInued) kotated ataudardized ala iuiinent cocUicleitta (each peraateter) lunction ierceat Effect teated Diecrtaiiiant function Percent ot variance Canonical correlation Hoab/ Rockj Craaa/ overheen Iorba sedge cover Uare lichen litter ground Season 1* 94.8 0.85 -- 1.19 -- 0.02 -0.43 -- 2* 5.2 0.35 -- -0.34 -- -0.07 -1.19 -- Upland 1* 99.6 0.74 1.80 2.63 3.40 0.02 1.60 -- Tree-Shrub j* 90.4 0.82 0.69 1.50 -- -0.49 -0.90 2* 7.8 0.39 0.17 0.51 -- 1.31 0.85 Marali *Stgntficeot (P 0.59 0.05) dlacr1utnatory Intor.ivation existed pilot to ioraat1on of diacrlwlnant function. r',) tree-shrub, whereas JCSN was abutted by open water and beach habitats. Species that typically foraged in upland sites on West Sand frequented WSIM during summer [e.g. marsh hawk (Circus cyaneus), long-billed marsh wren (Cistothorus palustris), starling (Sturnis vulgaris), common yellowthroat (Geothlypis trichas), house finch (Carpodacus mexicanus), and song sparrow (Melospiza melodla)]. The high elevation of WSIM and the immediate proximity of upland habitat facilitated use of WSIM by upland-associated species. Few species associated with upland habi- tats were detected In JCSM during summer. Number of species differed between JCSM and WSIM during fall (Table 1). This difference was apparently related to edge effect and availability of foraging sites. Some species that typically foraged in upland sites [e.g. rough-legged hawk (Buteo lagopus), marsh hawk, and starlingJ were observed foraging in WSIM. Thus the higher eleva- tion of WSIM and its juxtaposition to WSILJ influenced the number of species present. Additionally, species that typified an open water habitat tended to occur more often in WSIM than JCSM. Representative open water species present in WSIM included common loons (Gavia immer), greater scaup (Aythya mania), bufflehead (Bucephala albeola), and Bonaparte's gull (Larus philadelphia). Other species that foraged in shallow water or uiudflats [e.g. great blue heron (Andea herodias), black-bellied plovers (Piuvialis squatarola), and dowitchers (Limnodromus sp.) also were observed In WSIM but not JCSM during fall. The juxtaposition of WSIM to Baker Bay, a large, shallow bay with extensive amounts of intertidal concentrations of cormorants, mudflat at low tide which attracted waterfowl, gulls, shorebirds, and other 26 aquatic associated species helped account for the increased number of species observed in WSIM. Hence at low tides, species typically pre- sent in open water habitat often frequented WSIM whereas during low tide species that foraged In shallow water or mudflats were present. No concentrations of birds as large and diverse were observed near Jim Crow Sands. Severe winter weather probably contributed to the consistently lower number of species observed in winter for marsh habitats (Table 1). More species were recorded in WSIM than JCSM during spring. The elevated nature of WSIM was possibly an important element for foraging by water pipits (Anthus spinoletta), starlings, marsh hawks, and possibly some shorebirds. More species of shorebirds were present in WSIN than JCSM during spring (Appendix 2 and 3). The extensive amounts of mudflats in Baker Bay adjoining WSIM attracted large concentrations of shorebirds during spring. During high tide phases WSIM represented suitable habitat for shorebirds where foraging sites were exposed for a longer duration. Jim Crow Sands marsh was exposed for less time than WSIM and did not abut an area that offered as extensive array of foraging substrate as Baker Bay did for WSIM. The lack of seasonal differences in avian community characteristics was ascribed to the numerical replacement of neotropic migrants by nearctic migrants. Thus the exodus of hirundinids and other passerines was counterbalanced by the arrival of shorebirds and waterfowl. The considerable change in species composition observed from summer to winter emphasized this turnover (Table 4). Avian use, particularly shorebird, of the 3 marsh habitats exhibited a relationship to habitat characteristics. Western sandpipers 27 (Calidris mauri) preferred marshes with extensive amounts of mudflat (JCSM) during migration; peak migration periods for this species occurred in late summer and spring. Concentrations of dunlins (C. alpina) were greatest in JCSM during fall and spring migration and lowest in WSIM; MSM was represented by intermediate numbers of dunlins. Dunlins selected marshes with the greatest incidence of bare ground (mudflat). Thus vegetation probably influenced spatial distribution of dunlins and western sartdpipers among marsh habitats. Recher (1966) reported that the size and den sity of migrant shorebird populations was limited by available space, a function of tidal inundation, rather than by available food. Substrate composition, an important determinant of abundance and availability of food organisms for shorebirds, also influenced distri- bution and density of shorebirds (Recher 1966). Page et al. (1979) reported that dunlins and western sandpipers preferred foraging sites consisting of moderately sorted fine sand with little or no organic debris. Jim Crow Sands marsh was probably a more optimum foraging site for dunlins and western sandpipers, because its low elevation precluded dense stands of vegetation yet more niudflats were available at low tide for foraging shorebirds than WSIM provided. During winter, dunlins were most numerous in WSIM possibly because marsh sites at Jim Crow Sands and Miller Sands were covered by ice twice during winter. Reduced availability of estuarine tidal flats caused by freshets resulted in reductions in number of dunlins and least sandpipers at Bolinas Lagoon, California (Page ec al. 1979). During spring, the differences in density of dunlins among marsh habitats was not as apparent and probably resulted from the phenology of marsh vegetation; e.g. decreased incidence of litter, increased amount of bare ground, and short growth forms of most vegetation which allowed easier access and foraging for shorebirds. UPLANDS The significant season by habitat interaction effect (P 0.05) among avian community characteristics was ascribed to the variable vegetative attributes (bare ground to dense grass/sedge cover) which seasonally attracted different species complexes and differences in associated habitats which influenced species present for some uplands. Among upland habitats only WSIU:MSU and WSIU:WSIDU exhibited signif i- cant differences (P 0.01) in avian community characteristics. No differences in individual characteristics were detected for these comparisons (Table 1) . Density maximized separation of upland habitats (Table 2), but the separation (r = 0.48) attained on this discrimina- tion was not strong. Seasonally, summer differed from fall and winter; number of species and density differed (P 0.05) in each comparison (Table 1). Winter and spring were different (P teristics (Table 1). 0.05) with regard to all charac- Density was the variable which contributed most to seasonal separation Cr = 0.69) of uplands (Table 2). Seasonal placement on the continuum from low to high density was winter, fall, spring, and summer. The migration of neotropic migrants, the prin- cipal species comprising the summer avifauna of most uplands, probably accounted for this pattern. Thus the pattern established was of migrants returning in spring, attaining peak numbers during summer residency (breeding), declining with the advent of fall migration and with winter representing a season of minimum use by birds. No winter influx of neararctic species to upland habitats comparable in number to summer levels of neotropic species was observed. Several species of raptors, northern shrikes (Lanius excubitor), western meadowlarks (Sturnella neglecta), and some permanent resident species typified winter avifauna of most uplands. Low densities are characteristic of some of these species, particularly raptors. The counterbalancing effect observed for marsh habitats wherein passerine migrants were replaced by waterfowl and shorebirds in a numerical sense was not observed for upland habitats with the exception of Y1SSU. Species com- position of MSSU least resembled WSIU and WSID(J avian communities (Table 3). Avian communities of JCSU and WSIDU were quite different also (Table 3). Seasonal differences in species composition were st apparent for suinmer:winter comparisons (Table 4). The significant season by habitat interaction effect (P c 005) determined for upland habitat characteristics was probably attributable to the variable vegetative features of the uplands. parison of upland habitats were significantly different (P All coni- 0.01). Each habitat characteristic with some exceptions, differed for each habitat comparison. Exceptions were percent cover of grass/sedge on JCS1J which was similar to amounts for MSU and WSIU and covereage of litter on NSU and WSIU (Table 7). differed (P Percent cover of tall grass/sedge 0.05) among all comparisons of MSIJ, WSIU, and WSIDU. IlihIe 1. 1*rceut cor for ead IiaUtae racrertatic kOat baud lalaid, H1112r SuL, and Jiw (5i.w Sar IItb I t Hatitat 8iswair fall Water Spring tiiari e HS e (Jia rae i era It Ic" Li L Utter& Ibsa/lidieii ugbai. Bite 1wwwr fall Whter Spring ftiau Iaiier 1131 Ui,ter Spdng f.ui Saitier friU WInter Spring taii &aIer IatLL WinUx Spring ttain 1.0 2.9 0.0 0.2 1.0$ 0.5 IL! 5.2 1.4 3.9 5.4$ lo.9 42.5 f 2.o$ 1.2 4.8 5.7 6.e 4.8 s.s$ 92.8 91.2 93.4 93.4 92.1$ (LU t r (J.o 0.3 13.1 20.5 38.2 112.5 Iu.5 30.0 13.43 1 0.0 0.0 0.0 0.0 o.o$ 11.8 52.5 85.5 15.2 56.2$ 0.0 (3.0 jJ t 0.5 1.0 0.5 0.5$ 33.8 112.5 23.8 iLl 14.8J 72.6 63.3 55.4 41.5 58.2$ 4.3 3.0 0.6 2.9 ti 15.1 40.5 49.5 41.5 43.3$ 14.5 20.5 36.6 26.4 24.5 6.1 2.4 0.8 0.2 2.1$ 23.8 7 5.2 13.8 1).O IlLS 9.9 5.2 6.1 1.9 1.9 3.9 12.1 8.2 6.3$ 15.9 2/.3 41.5 31.9 2.t 1/.4 22.3 21.3 25.6 21.4$ 34.7 15.9 ILL 11.4 20.5$ 25.6 38.9 5.2 3.9 14.1 wsitt4 1u 813.ta Were toeifoawaI (artn for analyate. jrce.it cowir of litter dtfferul for all eaenal eiparIenna eaeq.t fall:apring aid wtiter;apring. CVetce,t wait of fnra differed fOr alt senarsial c&riarieona exCpt a&aia,irIu11 ad taLpring. cor of graiwt/aedi differed for alt weaaonal creparlerne except uu,.,er:ti1l cor of ,aiet/Ltrieu waw nat cawildered for aierepring. aol tati:eprtiig. W1I3J coipailacIn. o.os iurcenL. (J3 C 31 Tall grass/sedge was the dominant cover of WSIDU; percent cover of tall grass/sedge was less on WSIU and MSU (Table 8). 0.95) among upland habitats A high degree of discrimination (r was attained on the basis of percent cover of moss/lichen. A second discriminated discrinjinant function, based on the variable bare grounds 0.91) between upland habitats. MSSU, JCSU, and WSIDU had a low incidence of moss/lichen (Table 7) and were distinctly separated from (r WSIU and MSU on the first discritniriant function. Separation of MSSU, JCStJ, and WSID[J was based on percent occurrence of bare ground; WSIDU (low incidence) and MSSU (high incidence) represented the extremes on this gradient. Observed differences in upland habitats, with respect to avian community characteristics, were associated with those uplands exhi- biting the most dense vegetative cover. differed (WSIU:MSU, WSIU:WSIDU) did so Further, those uplands that only in a tnultivariate sense; avian characteristics were not different univarlately. The influence of habitat characteristics on avian communities; although not readily apparent with respect to barren (MSSU) versus vegetated (MSU, WSIU, WSIDU) habitats, was quite apparent in species composition. Species observed in MSSU primarily were associated with aquatic (gulls, terns, waterfowl) or intertidal (shorebirds) substra- tes. Few passerines other than swallows, horned larks (Ereniophila alpestris), and common crows (Corvus brachyrhynchos) were observed on NSSU although passerines comprised a major proportion of the avifauna of other upland habitats. 32 Table 8. Percent cover of tall grass/sedgea ( 15 cm) for NSU, WSIDU, and WSIU, Lower Columbia River, 1978-79. Percent cover tall grass/sedge Sunimer b MSU WSIU WSIDU aD I Fall Winter Spring Mean 5.2 0.1 t t 1.5 11.8 7.6 1.4 0.1 5.3 76.1 37.6 8.7 12.4 33.7 were transformed (arcsin fiT). bBroken vertical lines indicate significance (P CTrace (< 0.05 percent). 0.05). 33 The importance of MSSU to various avian species appeared to stem from its juxtaposition to open water and marsh habitats and its lack of vegetative cover. The diversity index for MSSU "edge effect" (3.74) was higher than for any other upland habitat (Table 9). Western sandpipers, dunlins, gulls, particularly California gulls (Larus californicus), Caspian terns, and pintails (Anas acuta) were observed either roosting, loafing, and/or foraging on MSSU during fall, winter, and/or spring. Page et al. (1979), Gerstenberg (1979), and Gill and Jorgenson (1979) reported that shorebirds utilized a variety of habitats--barren sand dunes, sand and rock beaches, salt marshes, uplands, pastures, and freshwater marshes for roosting, loafing, and/or foraging during high tide cycles. These activities corresponded to the observed activities of western sandpipers and dunlins on MSSU. Birds normally associated with upland habitats were absent from MSSU which indicated that vegetative structure strongly influenced avian species composition. Use of ?ISSU by species not nor- mally associated with upland habitats accounted for the similarity of MSSU to other uplands with regard to avian community ctxaracteristics. For those upland habitats that differed from each other (WSIU:MSU, WSIU:WSIDU) individual avian community characteristics were not different. Density however, provided the most information with regard to discrimination among these upland habitats. Typically, 3 to 4 species were most abundant each season in WSIDU whereas WSIU and NSU had 2-3 numerically dominant species during summer and spring and none during fall and winter. Although several species present in MSU during winter were very common, their presence was attributed to the 34 Table 9. Diversity indicesa for "edge" of individual habitats, Lower Columbia River. Habitat WSIM WSIDU WSIU WSLB WSITS WSIT Total perimeter (meters) 10,296 1,260 7,213 6,444 7,038 6,462 Hectares DI 54.7 3.93 11.3 87.2 1.06 2.18 19.0 4.17 48.8 33.0 2.84 3.17 38.7 1.22 21.3 3.74 19.9 13.4 MSB 2,700 6,120 5,310 5,868 2,862 MSTS 9,468 34.1 3.36 4.52 2.48 4.57 JCSM JCSU JCSB 4,986 3,474 4,374 29.8 19.6 MSM MSSU MSU MSSB aDiversity = total perimeter/2i 10.6 27.2 (area) 2.58 2.21 2.37 35 proximity of other habitats or to transient species passing over the habitat. Elimination of these species from the analysis might have increased similarity of MSU to '1SIU. The difference observed for WSIU:WSIDU appeared to be habitat related. Tall grasses typified WSIDU whereas WSIU was represented by a short grass/sedge community. The height and structural integrity of WSIDU tall grasses were suitable for long-billed marsh wrens (Cistothorus palustris) which were abundant during all seasons whereas this species was not observed in WSIU. Verner (1965) and Verner and Engelsen (1970) reported that long-billed marsh wrens typically occurred and nested in dense stands of emergent vegetation. Song sparrows (Melospiza inelodia), relatively abundant during fall and winter in WSIDU compared to WSIU, were probably attracted by the pre- sence of tall grass, dead Sitka spruce trees and invading willow and alder trees. Further, American goldf inches (Carduelis tristis) were observed foraging in WSIDU during late summer on achenes of Composites and concentrations of goldf inches caused a relatively high density estimate for this species. Densities of savannah sparrows (Passerculus sandwichensis) were remarkably similar for WSIU, WSIDU, and MSU (Appendices 5, 7, and 8). Starlings were quite abundant in WSIDIJ during winter and were observed foraging on the densely matted residual vegetation. The greater occurrence of organic debris asso- ciated with the densely vegetated diked upland compared to WSIU probably resulted in a greater availability of invertebrates and thus was more attractive to starlings. 36 Edaphic features of WSIDU may account for the dense vegetative cover present. Deposition of silt from the tiwaco Channel dredging operation in 1973 probably resulted in a more nutrient laden substrate for plant growth (N. Ellifrit, USFWS, pers. comm. 1981) than other upland habitats which were sandy in nature. Succession appeared to be more rapid for WSIDU as red alder and willow were pioneering the site during the study period and by 1981 considerable growth of these trees had occurred. TREE-SHRUB A significant (P ' 0.05) season by habitat interaction effect was determined and was attributed to seasonal differences in avian community characteristics among habitats. Since the seasonal differences contibutitig to the interaction effect were considered explainable biologically (see discussion, p. 40) and apparently were not attributable to random sampling error, I proceeded to test season and habitat effects. Tree-shrub habitats were not different with respect to avian community characteristics (Table 1). Species composition of all tree- shrub habitats was quite similar (Table 3) and exhibited considerable seasonal similarity (Table 4). Seasonal comparisons that exhibited differences were fall:surmner and fall:spring (Table 1). Evenness was different for both signifi- cant seasonal comparisons; number of species differed for the fall: spring comparison (Table 1). Discrimination among seasons for tree- shrub habitats was based on species equitability (Table 2). The degree of resolution (rc = 0.73) among seasons on this basis was high, 37 A gradient from low (fall) to high equitability (summer) was determined on this basis. Season placement on this gradient was fall, Fall and winter represented seasons when winter-spring, and summer. flocks of birds were encountered during censusing. Typically one or two species were numerically dominant, hence equitability was Breeding birds during spring and summer were dispersed, decreased. hence a higher equitability. A significant (P 0.05) season by habitat interaction effect was determined for tree-shrub habitats and was ascribed to the differing seasonal incidence of moss/lichen. This difference in pattern for percent occurrence of moss/lichen was apparent for WSIT where natural degeneration of forbs and grass/sedge led to increased exposure of the moss/lichen layer (Table 10). ignored. Consequently the interaction effect was Among tree-shrub habitats, MSTS differed from WSIT (P for all habitat characteristics measured. 0.05) Percent f orb and overhead cover of MSTS were similar to WSITS, otherwise habitat characteristics of these two habitats differed (P 0.05). Habitat characteristics of tree-shrub habitats on West Sand Island were different (P except for occurrence of litter and grass/sedge. 0.05) Canopy height (1.90 vs. 1.22) and canopy base height diversity (1.54 vs 0.58) and mean tree circumference (0.58 vs. than WS1TS. analysis. 0.40) of MSTS were greater (P c 0.05) Paucity of trees in WSIT precluded its inclusion in this The difference in tree characteristics for HSTS:WS1TS comparison was attributed to the presence of black cottoawoods In MSTS. Cottonwood typically exceeded heights and circumferences iSbIe 10. Innt ora va1ue tot eath hatd tat C oi.i,riat1c, tne-ahwb h.thlt&a, 1eE thioaubia &1wr, 1918-19. Hbitnt C6nrncter1stc Utter Mrf1iden Tree-6hnib awtat 6 wsfl WSIT I 1C Fall Wtltcrd SI)rtn4 1&un &I1u0r }rceuL or& Jlir 18U Winter Spring ttnO 10.5 10.5 9.3 ILl 1231 45.5 65.2 75.2 57.4 60.81 8.2 6.6 9.3 8.2 8.11 37.6 50.5 64.3 44.5 0.4 1.0 3.2 3.9 2.11 36.6 52.5 10.8 5/.4 Saaiere inaat 10.5 1.6 0.4 15.1 1.4 8.2 5.2 2.9 3.9 4'J.2 4.8 6.6 2.9 53 5.0 40.5 31.9 21.3 31.6 49.3 23.0 16.0 1.1 33.1 Io.Uf 50.4 I.2 15.9 21.3 re tnasfonanh arcain baiy perwit ccr of ead hahttar parater dittenal tot alt cu1parttaia except wtCe ot .aitsa notnd. Ch±rUnt cor of iitas/lldlen t5.rIilg awaar did not difter frtu tall and w1ztr Iniali. 4Lt C0lt of uuas/Iithen during wli&teti did not dilfer truu fall and aprtng lea1. cower of forb dudng aIsjIar did act differ ira fall nix! aprhig levels. coer of gras/aod dudng wliter did not ditler fran apring law3a. o'ir6ead cover was a lt for a nrapr1ng and faLleiuter cparaia. 68rdn tlnoi indiante aignificait ditferenan (F < 0.115). iu WUILet 8pu 11.1 45.5 41.3 iu.a si.i 32.8 64.3 343 31.9 no.4 41.8 10.4 o.8 5.9 11.8 5.2 5.01 39 normally attained by willow and red alder, which were abundant in both tree-shrub habitats. Percent overhead cover accounted for most separation (rc = 0.92) of tree-shrub habitats on the first discriminant function (Table 6). Percent cover of grass/sedge was the habitat characteristic primarily responsible for habitat separation (r (Table 6). = 0.77) for the second function Habitats ordered along the first gradient emphasized the upland nature of WSIT (few trees hence low overhead cover). West Sand Island tree-shrub was distinctly separate from MSTS on the second function because of the higher percent cover of grass than NSTS. Although no difference among tree-shrub habitats with regard to avian community characteristics was found, considerable differences in habitat characteristics existed. The presence of a tree-shrub coni ponent in WSIT attracted species associated with forested sites and emphasized the importance of the tree-shrub component on avian communities. Willson (1974) determined that the addition of trees was instrumental in attracting additional species and attributed the increase in number of species to increased environmental patchiness. The greater vertical diversity of NSTS compared to WSITS did not produce a significant increase in number of species. Wilison (1974) determined that few species were added after initial development of a canopy layer. Seasonal differences in pattern (season by habitat interaction) among tree-shrub habitats for number of species, density, and diversity were attributed to: 1) ecotonal nature of WSIT, 2) probable differential availability of cavities for nest sites (MSTS, WSITS), 40 3) associated habitats, 4) passage of neotropical migrants (particularly parulids), and 5) density of winter residents. The contrasting seasonal patterns (interaction) in avian conimunity characteristics observed for tree-shrub habitats indicated that MSTS differed from WSIT and WSITS at least during summer and fall. Differences were noted in number of species, density, and diversity during these seasons for MSTS contrasted to WSITS and JSIT. The eco- tonal nature of WSIT, which provided habitat suitable to species characteristic of both upland and tree-shrub habitats, probably accounted for the high number of species recorded there and the high density and diversity values. Thus only the contrast between the most similar tree-shrub habitats (structurally) remained unexplained. The mean number of species observed in MSTS during summer was similar to that recorded by Crawford and Edwards (1978) (7 and 8.7 respectively). Their estimated avian density for ?ISTS exceeded mine (8.5 versus 6.0 birds/ha, respectively) during summer yet remained well below the level recorded during this research for WSITS. Their estimated density for an unidentified tyrannid was excluded from the comparison as I attributed it to sampling anomaly. Consequently, the recalculated avian density for their summer data complemented the density estimate derived during this research and solidified my contention that MSTS differed from WSITS. More cavity nesting species were present (7 vs. 3, respectively) and they attained a higher density (4.2 vs. respectively) in WSITS than NSTS. 2.1 birds/ha Density of cavity nesting species in MSTS was 2.5 birds/ha during research by Crawford and Edwards (1978). 41 Miller Sands, approximately 50 years old (Crawford and Edwards 1978), was characterized by relatively young trees that probably have not attained a suitable size and/or stage of decadence suitable for pro- duction of cavities, hence fewer opportunities for cavity nesting species. Apparently more snags and use of cavities by birds occurred in WSITS than MSTS. A 1939 aerial photo of West Sand Island confirmed the presence of trees there, while Miller Sands, visible in the background, appeared barren. Although larger trees were present ía MSTS, these were cottonwoods which were still growing vigorously and thus provided few cavities. Large cottonwoods in MSTS seemed more susceptible to blowdown with increasing size which may effectively remove them as potential snags. The presence of 3 fringillids (American goldfinch, house finch, and savannah sparrow) in WSITS that did not occur in MSTS was possibly attributed to the greater suitability of WSIT and WSIDU for foraging. These species utilized WSITS primarily for perch and loafing sites and foraged along the edges or in upland sites. Number of species present during fall in MSTS exceeded that of WSITS primarily because re parulids were observed. warblers observed in MSTS were not observed in WSITS. Five species of Warbiers were not observed during fall in FISTS by Crawford and Edwards (1978). The difference in number of warbler species may have reflected a difference in migratory patterns between FISTS and WSITS. Bagg (1969) reported that chickadees tended to funnel along shorelines upon encountering large water bodies until a shorter crossing point or land bridges were encountered. The decreased number of species during fall 42 in WSITS and WSIT was attributed to the seasonal exodus of neotropical migrants. Density was the characteristic which exhibited the most contrast between MSTS and West Sand Island tree-shrub habitats (WSIT, WSITS) during fall. The higher density of black-capped chickadees and ruby- and golden-crowned kinglecs in MSTS compared to WSIT and WSITS appeared most responsible for this observed difference. Black-capped chicka- dees were considerably more numerous during fall than summer in 1STS. Crawford and Edwards (1978) also reported higher densities of blackcapped chickadees during fall than summer. High estimated densities for both species of kinglets during fall as determined here were not reported by Crawford and Edwards (1978). Their fall sampling period occurred during an unseasonably dry period which may have delayed the migration of kinglets to lower elevation areas such as MSTS; however their data indicated a considerable increase in ruby-crowned kinglets with the onset of winter. The habitat factor responsible for these differing densities was not apparent. Differences in equitability among seasonal comparisons were attributed to the very high densities attained by a few species during fall. Black-capped chickadees and ruby- and golden-crowned kinglets flocked together during fall and typically were the most numerous birds. Consequently, equitability during fall was lower in contrast to spring and summer when flocking was less prevalent; reduction in flocking behavior was probably attributable to territoriality and absence of some species (e.g. kinglet spp.) that occurred with tue advent of spring and the breeding season. 43 BEACH Avian community characteristics of beach habitats were similar for habitat, season, and interaction effects (Table 1). Species com- position of beach habitats was quite similar although WSL3 exhibited a consistent difference (20 percent) from all other beach nabitats (Table 3). This difference probably reflected the occurrence of pelagic species at WSIB. re Seasonally, species composition of beach habitats was most different between summer and winter (Table 4). Although species composition exhibited a marked seasonal turnover, avian community characteristics were not significantly affected. As in marsh habitats, the migration of neotropical migrants was negated by migrant nearctic species. Bare ground, the only characteristic considered in comparisons of 0.025) on WSIB than on beach habitats, was significantly lower (P other beach habitats (Table 11). Accumulation of debris on WSIB was probably enhanced by its exposure to the main channel of the Columbia River; other beach habitats were not exposed to the main channel. This difference apparently was of minimal importance to birds. WEST SAND ISLAND The significant interaction effect (P ' 0.05) associated with ana- lysis of avian community characteristics was attributed to the diverse array of habitats considered and varying seasonal utilization of these habitats by birds. Avian community characteristics differed for vir- tually all habitat comparisons (P < 0.05) except WSIM:WSIU, WSIDU:WSIB, 44 Percent occurence of bare ground for 4 beach habitats: Table 11. MSSB, MSB, JCSB, and WSIB, Lower Columbia River, 1978-79. Percent bare grounda Summer Fall winter Spring Mean 97.1 97.4 94.8 98.5 96.9 MSB 99.2 93.8 91.8 94.8 94.9 JCSB 99.3 87.5 98.0 96.4 95.3 WSIB 92.3 93.4 76.1 95.2 89.2 Habitat MSSB b aData were transformed (arcsin JiI). bverticai lines connecting habitats indicate similarity; broken lines indicate significant difference. 45 WSIT:WSIB and WSIT:WSITS. Typically, species equitability and density were higher f or the tree-shrub habitats than for other habitats (Table 12). A continuum from low to high density formed the primary basis for separation (r = 0.69) of West Sand Island habitats (Table 2). Habitats were ordered along the first function beginning with tISIU, then WSIDU and WSIM, WSIB, and terminating with tree-shrub habitats (WSITS and WSIT). Equitability formed the basis for separation on the second discriminant function. This separation (r 0.48) enhanced the distinctiveness of the tree-shrub habitats from all other West Sand Island habitats. Species composition of West Sand Island habitats was relatively different for most comparisons (Table 13). Species composition of tree-shrub habitats differed from other West Sand Island habitats most, although species composition of WSIT was similar to uplands (Table 13). Considering that WSIT retained many similarities structurally of an upland habitat, some similarity in species composition to uplands was expected. Species composition of WSIM differed considerably from other West Sand Island habitats (Table 13). Seasonally, species composition during winter was quite different from summer; other seasonal comparisons were similar (Table 14). The changeover from summer to winter reflected the impact of migration on species composition. Tree-shrub habitats, the most advanced successional stage on West Sand, were readily distinguishable from more intermediate stages. Evenness and density were the characteristics that best separated T1 Table 12. Mean values of each avian community characteristics for habitats comprising each island, Lower Columbia River, 1978-79. Habitat complex Number of species Densitya (birds/ha) Diversity Evenness Miller Sands 175d MSMb,c 137b,c MSSUb 82b,c 1.28 MSUb 35b 1.36 7.2 13.5 068b,c 064C o.00c 1.38 O.6O' 1.62 0.68 MSSB 10.6 MSB 11.3 MSTSb,c 109b 12.4 1.84 JCSM 11.4 12.4 1.42 0.59 JCS1J 8.7 4.6 1.12 0.50 JCSB 11.5 41.5 1.39 0.58 161d 059e,f 7.Oc 078b,c Jim Crow Sand West Sand Island 152e,f,g 92e,f 10.3 112e WSITe 11.6 14.0 7.2 5,8,g 1.24 36f,g 1.43 8.4 1.52 lO.4f 1.90 12.3 1.93 0,66g 063g 074e 47 Table 12. (continued) aDensity data were transformed (square root). bMSM differed from MSSU, MSU, and MSTS. Number of species differed in each comparison; evenness differed for MSM:MSTS comparison. CMSTS differed from all other Miller Sands habitats. MSTS charac teristics which differed among habitat comparisons were number of species (MSM, MSSIJ), density (MSU, MSB), and evenness (MSM, MSSU, MSU, MSSB). dDiversity data were not utilized in analysis because of high correlation with evenness: Miller Sands (r = 0.74), West Sand Island (r = 0.76). eWSIM differed from WSIDU, WSIB, WSIT, and WSITS. Number of species was different from WSIDU and WSIB; evenness differed from WSIT and WSITS. WSIT differed from WSIDiJ, WSIU, and WSIM. from WSIDU and WSIM: Number of species differed density differed from WSIDU and WSIEJ; and evenness differed from WSIM and WSIDtJ. differed from WSIM (number of species), WSIDEJ (density, evenness), WSIU (densisty, evenness), and WSIB (evenness). 48 Table 13. Habitat comparisons of species composition values (MacArthur's Difference) for habitats grouped by island, Lower Columbia River, 1978-79. Habitats Island/habitat Miller Sands MSM MSSU {SU MSSB MSB MSSU 1.16 MSU 1.40 1.45 MSSB 1.16 1.10 1.42 MSB 1.16 1.23 1.37 1.20 MSTS 1.82 1.87 1.52 1.89 1.70 JCSM JCSU WSIU WSIB WSITS Jim Crow Sands JCSU 1.57 JCSB 1.16 1.32 WSIM WSIDU West Sand Island WSIDU 1.49 WSIU 1.52 1.37 WSIB 1.53 1.63 1.45 WSITS 1.69 1.59 1.65 1.75 WSIT 1.57 1.37 1.39 1.62 1.13 49 Table 14. Seasonal species composition values (MacArthur's Difference) for combined island habitats, Lower Columbia River, 1978-79. Habitats group by island Season Habitat complex Summer Fall Winter Miller Sands Fall 1.40 Winter 1.58 1.34 Spring 1.27 1.36 1.31 Jim Crow Sands Fall 1.64 Winter 1.84 1.32 Spring 1.44 1.39 1.37 West Sand Island Fall 1.35 Winter 1,64 1.22 Spring 1.15 1.26 1.45 50 tree-shrub habitats from other West Sand Island habitats. Concentra- tions of waterfowl, shorebirds, or gulls commonly occurred in JSIM or WSIB which tended to depress equitability. A tendency for more spe- cies to occur in WSIM than other West Sand Island habitats, except tree-shrub habitats, was probably associated with edge effect. The high elevation, which precluded inundation and resulted in habitat similar to uplands, was probably attractive to upland species MILLER SANDS ISLAND Terrestrial habitats on Miller Sands (MSsU, MSU, MSTS) were dif- ferent from MSM, whereas avian community characteristics for beach habitats were similar to MSM (Table 12). Another pattern that deve- loped concerned MSTS, which differed from all other habitats on Miller Sands (Table 12). Number of species differed for each significant comparison except beaches (MSB and MSSB):MSTS and MSIJ:MSTS (Table 12). Avian density for MSTS differed from that determined for MSU and MSJ3 (Table 12). Evenness was higher for MSTS for all comparisons except MSB:MSTS (Table 12). Number of species best described habitat separation (r for the first discrinilnant function (Table 2). = 0.62) A habitat continuum was developed with upland habitats (fewest species) and MSM (most species) most distant from each other. Habitats were scaled from low diversity to high, i.e. uplands, beaches, marsh, and tree-shrub along the second function. Species composition of MSTS differed considerably from that recorded for other Miller Sands habitats (Table 13). All habitats 51 subject to tidal inundation and MSSU and MSSB were quite similar in species composition (Table 13). Seasonally, species composition during winter was most different from summer (Table 14) and reflected the impact of migration on composition. Two patterns were evident from comparisons among Miller Sands habitats: 1) MSTS differed from all other habitats, and 2) MSM dif- fered from terrestrial habitats on Miller Sands but not beach habitats which were subject to the same tidal influence. An increase in avian community characteristics associated with a more advanced successional stage seemed apparent for Miller Sands habitats. Species equitability for MSTS was greater in most significant comparisons (4 of 5) than for other Miller Sands habitats. Large, often monotypic flocks, of either waterfowl, gulls, or shorebirds commonly occurred in marsh and beach habitats. Such aggregations tended to depress equitability. Miller Sands spit upland resembled beach and marsh habitats regarding aggregations of birds. Low densities of birds during fall and winter in MSU often were dominated by one species, hence equitability was low. Miller Sands marsh differed from terrestrial habitats associated with Miller Sands apparently because of its attractiveness to aquatic associated species and certain terrestrial species. Shorebirds, gulls, waterfowl, and swallows, species which utilized MSM at varying stages of tidal inundation and seasons, formed the basis for the higher species richness components recorded for MSM than was recorded for terrestrial habitats. These species complexes also frequented beach habitats which were similar to MSM with regard to avian com munity parameters. 52 JIM CROW SANDS Habitats on Jim Crow Sands were similar in avian community characteristics and only JSCM:JCSU comparison exhibited any notable difference in species composition (Table 13). Species composition changed from summer to winter reflecting the influence of migration on community composition (Table 14). The structural simplicity of Jim Crow Sands habitats, related to their early successional stages and age possibly accounted for the lack of observed differences among these habitats. CONCLLTS ION Structural nature of the vegetation for the habitats investigated influenced avian species composition and community parameters. Differences were apparent for comparisons of various habitats and among habitats. Later successional stages, typified by greater structural complexity, tended to support more species and attain higher densities and equitability. Seasonal changes in avian com- munity parameters associated with the influx and exodus of neotropic and neararctic migrants were also of importance. MANAGEMENT RECOMMENDATIONS Management decisions, with respect to these islands, should be based not only on gross habitats present, but the structural nature of each habitat and its juxtaposition to other habitats. Changes in spe- cies composition and parameters were quite apparent as vegetative 53 structural qualities changed. Thus a manager has the capacity through vegetative manipulation to alter avian species composition and parameters on these islands. iiowledge of the species present and their associated community parameters can aid in predicting the outcome for events, such as dredged material deposition, that alter these habitats. Differential seasonal use of habitats by birds points out the necessity for managers to consider these habitats on a seasonal basis. Thus management decisions can be made apriori with regard to species composition and community parameters desired. 54 LITERATURE CITED Adams, C. 1908. The ecological succession of birds. Audubon Field Notes 23:4-12. Bagg, AM. 1969. The changing seasons. Barnes, D. 1971. Anatomy of a spoil island. I Univ., Kingsville, Texas. Bond, R.R. 1957. Auk 25:109-153. Texas A & M.S. Thesis. 70 pp. Ecological distribution of breeding birds in the upland forests of southern Wisconsin. Ecol. Monogr. 27:35l-382. Buckley, P.A. and F.G. Buckley. 1975. 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Chinook, Wash., and the head of Sand Island. Committee on Rivers and Harbors, U.S. House of Representatives, Document No. 50. 11 pp. A comprehensive study of suc- 1977. Coastal Zone Resource Corporation. cessional patterns of plant and animals at upland disposal areas. Tech. Rep. D-77-2. Vicksburg, Miss. Army Engineers Waterways Exp. Sta., U.S. 359 pp. Cohen, E. and P. Burns. Document No. 413 (Rev. A), Northwestern variance and covariance. Univ. SPSS-MANOVA--Multivariate analysis of 1977. Evanston, Iii. 155 pp. Crawford, J.A. and D.K. Edwards. 1978. Habitat development field investigations, Miller Sands marsh and upland habitat development site, Columbia River, Oregon. Appendix F: Postpropagatioa assessment of wildlife resources on dredged material. Tech. Rep. D-77-38, U.S. Army Engineer Waterways Exp. Sta., Vicksburg, Miss. 68 pp. DesGranges, J.-L. 1980. Avian community structure of six forest stands in laNauricie National Park, Quebec. Papers No. 41. Occas. can. Wildi. Serv. 34 pp. Dunstan, F.M., and R.R. Lewis, III. 1974. Avian utilization and plant succession on dredged material islands in Tampa Bay, Florida. DuPue, 3. Unpub. Rep. 1974. Coastal Zone Resources Corp. 73 pp. Nesting and reproduction of the black skimmer on four spoil islands in the Laguna Madre. Univ., Kingsville, Texas. M.S. Thesis, Texas A & I 56 Mi analysis of avian communities on a dredged 1979. Edwards, D.K. M.S. Thesis, Oregon State Univ., Corvallis, OR. material island. 47 Pp. Edwards, J.M. and M.A. Edingtoa. partition in breeding Spatial patterns and habitat 1972. birds of an upland wood. J. Anim. Ecol. 41:331-357. Habitat utilization by wintering and 1979. Gerstenberg, R.H. migrating shorebirds on Humboldt Bay, California. F.A. Pages 33-40 in Pitelka (ed.), Shorebirds in Marine Environments. Cooper Ornithological Society. Avian Biology No. 2, Gill, R., Jr., and Paul D. Jorgenson. 1979. Studies in 261 pp. A preliminary assessment of timing and migration of shorebirds along the northceritral Pages 113-123 in F.A. Pitelka (ed.), Shorebirds Alaska Peninsula. Studies in Avian Biology No. 2. in Marine Environments. Cooper Ornithological Society. Haapanen, A. 1965. Bird fauna of the Finnish forests in relation to forest succession: Jefferson, C.A. 1974. I. Ann. Zool. Fenn. 2:153-196. Plant communities and succession in Oregon coastal salt marshes. OR. 261 pp. Ph.D. Thesis. Oregon State Univ., Corvallis, 192 pp. Johnston, D.W. and E.P. Oduin. 1956. Breeding bird populations in relation to plant succession on the Piedmont of Georgia. Ecology 37:50-62. Karr, J.R. 1968. Habitat and avian diversity on east central Illinois. Condor 70:343-357. strip-mined lands in 57 Karr, J.R. and R.R. Roth. 1971. Vegetation structure and avian Am. Nat. diversity in several new world areas. northern lower Michigan. Kohout. Kim, J. and F.J. communities Bird populations and biotic 1948. Kendeigh, S.C. 105:423-435. Ecology 29:101-114. 1975. Analysis of variance and covarlailce: subprograms ANOVA and 0NE.(AY. Pages 398-433 in H.H. Nie, C.H. Hull, J.G. Jenkins, K. Steinbrenner, and D.H. ieut, 675 pp. New York. 1975. Discriminant analysis. Page 434-467 in H.H. Nie, C.H. Hull, J.G. Jenkins, K. Steinbrenner, eds. Statistical Package for the Social Sciences. 0.1-i. Bent, NcGraw-1-1ill, 675 pp. New York. Landin, M.C. eds. McGraw-lUll, Inc., Statistical Package for the Social Sciences. I<lecka, W.R. in 1978. National perspective of colonial waterbirds nesting on dredged material islands. 43:89-99. Resour. Conf. MacArthur, R.H. Population ecology of some warbiers of 1958. northeastern coniferous forests. 1965. Trans. N. Am. Wildi. Nat. co1ogy Patterns of species diversity. 1977. north Florida. Cambridge Philos. 40:510-533. Soc. Biol. Reviews. NcElveen, J.D. 39:599-619. The edge effect on a forest bird community in Proc. Ann. Conf. Southeast Assoc. Fish Wildl. 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Inventory of riparian habitats and associated wildlife along the Columbia River (from McNary Dam to the mouth of the Columbia River). Final Report, U.S. Army Corps of Engineers, NPD Contract No. DACW68-73C-0186. Oregon State Univ. Oregon Historial Society. 1980. 771 pp. Columbia gateway, a history of the Columbia River estuary to 1920. PNRBC, Vancouver, WA. Page, G.W., L.E. Stenzel, and C.M. Wolfe. 1979. 65 pp. Aspects of the occurrence of shorebirds on a central California estuary. Pages 59 15-32 in F.A. Pitelka (ed.), Shorebirds in Marine Environments. Studies in Avian Biology No. 2. Cooper Ornithological Soceity. 261 pp. edge." A diversity index for quantifying habitat 1975. Patton, D.R. 3:171-173. Wildi. Soc. Bull. Peters, C.F., R.O. Richter, D.A. Manuwal, and S.G. Herman. 1978. Colonial nesting sea and wading bird use of estuarine islands in Tech. Rep. the Pacific Northwest. Waterways Exp. Sta., Vicksburg, Miss. birds. 179 pp. Some aspects of the ecology of migrant shore- 1966. 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Anderson, and R.H. Strand. 1975. Dominant pat- terns iii bird populations of the eastern deciduous forest biome. Pages 90-95 in Smith, D.R., Tech. Coord., Proceedings of the symposium ott management of forest and range habitats for nongame birds. Soots, R.F., Jr. Tech. Rep. USDA For. Serv. Gen. and J.F. Parnell. 1975. W0-l. 343 pp. Ecological succession of breeding birds in relation to plant succession on dredge islands in North Carolina estuaries. Raliegh, N.C. N.C. Sea Grant Pubi. UNC-SG-75-27, 91 pp. Stauffer, D.F. and L.B. Best. riparian communities: J. Wildi. Manage. 1980. Habitat selection by birds of Evaluating effects of habitat alterations. 44:1-15. Thompson, D.H. and M.C. Landin. 1978. An aerial survey of waterbird colonies along the Upper Mississippi River and their relationship to dredge material deposits. Tech Rep. D-78-13. Waterways Exp. Sta., Vicksburg, Miss. Verner, J. Condor 1965. U.S. Army Eng. 67 pp. Breeding biology of the long-billed marsh wren. 67:6-30. Verner, J. and G.H. Engelsen. 1970. Territories, multiple nest building, and polygyny in the long-billed marsh wren. Auk. 87:557-567. Wilison, M.F. ture. 1974. Ecology Avian community organization and habitat struc55:1017-1029. 61 Woodward-Clyde Consultants. Sands Complex. 1976. Terrestrial ecology of the Miller U.S. Army Corp. of Engineers, Portland District. Yeaton, R.1. and M.L. Cody. sparrow populations. 1975. Competitive release in island song Theor. Pop. Eiol. 5:42-58. S aDIaNdcW 1. Dowttcher app. Ijeatern sandpIper Sanderling Northern phularope Glaucous gull Glaucous-winged gull Western gull flerrtog gnU California gull hung-billed gull Hew gull Bonaparte's gull Casplan tecu Belted kingfisher Veux'a swift black-b11ted plover Spotted sandpiper Greater yellowlege Lesser yelluwlega Least sandpiper Dunlin K1tt&ieer Scmipaltaated plover Anerican coot Utd eagle Coiwaon mergauaer Red-breasted rganser 24.5 4.0 39.8 43.3 4.8 10.9 6.6 158.6 21.3 4.2 4.8 July 15.1 5.3 12.4 34.2 2.6 2.8 June 1.9 101.1 0.8 1.6 30.2 33.4 3.2 15.9 235.8 23.7 5.1 6.5 40.0 11.0 2.8 11.4 14.6 59.7 9.9 6.4 0.7 1.9 Septewber 230.6 5.3 August 1.8 29.0 4.9 1.0 14.6 2.11 34.7 3.9 28.9 6.4 24.7 5.0 4.6 0.7 1.9 22.4 1,409.5 397.9 5.0 2.8 9.5 0.1 5.0 94.4 9.9 109.0 20.1 86.1 5.0 43.11 6.4 October Nuvewber 8.8 1.4 5.6 5.6 3.8 5.0 5.0 15.1 130.7 20.2 13.2 35.4 35.8 13.0 1.6 29.3 9.3 5.0 2.4 25.5 143.11 7.3 19.0 132.7 0.4 3.2 3.0 3.2 1.6 3.9 0.6 March 38.2 8.9 1.6 6.4 2.11 10 ebreary 0.2 2.5 1.4 3.2 bceaber January Mouth Species den1ty (blrde/l00 ha), June 19711-Hay 1979, mIter Saw.Iu Harsh, Cotuebte tUver, Oregon. Uorned grebe Western grehe Døuble-crcted coraoraut Crest blue heron Whistling swan Canada goose Mallard Gedeell Ptntati Green-winged teal American wigeon Greeter eceup Lesaer ecaup BufCLehead Species Appendix 00.1 35.7 13.2 20.2 i.s 10.3 135.4 752.5 9.5 0.4 4.0 Apr.L 2.6 4.5 4.5 12.0 21.9 106.1 430.6 3.11 25.5 11.8 5.3 19 tay cy 1. (contInued) Rveoneaa In 250.3 2.06 0.14 19 486.8 2.29 0.78 11 651.2 1.48 0.62 13 499.8 2.03 0.79 13 621.1 1.65 0.64 Llen8ity (birds/ha) Olveratty 35.4 Number of species 3.2 25.5 4.8 47.1 4.8 0itober 12.7 Septuwber 159.2 August 33.9 July 111.9 319.1 9.1 June 85.8 31.0 70.7 Violet-green wa1low Tree swallow Ram swallow Cliii swallow Common crow Rrowa-headed cowbird Song sparrow SpecIes Appendli (1.46 1.31 2,235.9 17 7.2 Novesbet 40.2 2.09 0.91 It) Decewber Month 301.2 1.95 0.74 14 20.1 Januaty 79.8 1.62 0.74 9 February 405.5 1.56 0.69 10 254.6 54.1 March In 721.2 4.57 0.57 17 4.8 80.4 2n.0 flay 2,392.9 1.85 0.48 16.1 72.7 Q75.0 104.8 Apr11 Appendix 2. lpeclea density (birda/lOO ha), June 1978-Muy 1919, Jtiu Crow Sands Harsh, Coluwbta River, Oregon. Houth Species June July August SepLewbur October Novewber Conenu loon Oecaber January 1.6 Double-crested coranrant Great blue heron 0.4 Whistling swan Canada goose tbtury 7.1 Piutati Green-winged teal Macrican wigeon Northern hoveier 11.7 5.6 50.7 12.9 31.5 93.9 1.6 6.5 3.2 31.8 163.5 1.6 Surf acoter 3,9 0.8 47.4 0.8 18.4 17.7 15.8 12.7 31.8 3.2 35.4 3.2 70.3 462.6 4.4 260.7 681.8 83.9 4.4 4.9 12.6 1.6 6.5 7.1 2.4 5.4 Bonaparte's gull 69.7 241.1 Casptau tern Violet-green swallow 120.5 11.8 2.4 35.9 18.0 20.2 0.8 60.5 9.0 6.6 3,351.9 61.7 19.9 33.6 3.0 6.5 70.5 1,274.5 8.8 329.4 44.2 13.2 2.8 2.8 3.9 8.8 6.6 3.9 246.2 13.1 8.1 7.3 5.1 66.2 14.7 63.9 1.8 5.4 4.4 225.8 6.4 6.5 2.4 26.9 42.1 4.0 49.7 12.7 15.1 21.4 39.0 39.8 20.i Lb 11.1 327.2 0.8 hO.! 2,931.2 8.8 24.3 1,340.5 40.7 2.6 2.2 52.0 4.2 Cons&on crow 3.2 52.4 6.5 Western sandpiper Sauderitag Northern phalarope Glaucous-winged gull Western gull Cliff swallow 0.6 7.9 2.5 27.5 I3unlin Tree swallow Barn swallow 5.6 4.0 5.4) Coajan iaerganaer Red-breasted merganser Merlin Semtpslinated plover Cossion tern 8.9 0.6 24.5 Greater acaup Duitleheati New gull Hay 21.2 62.9 Mallard herring gull California gull King-billed gull Aptil 13.5 32.5 Snow goose KilIdeer Spotted sandpiper Greater yellowlegs Least sandpiper Hatch 9.0 2. (continued) Evenness Nuather of species Denolty (birds/ha) Dtveretty Long-billed tharab wren Water pipit Aiaerican goldfinch Savannah sparrow Song aperrow Spectea Appendix 259.4 1.89 0.82 10 June 14 19.9 Auguat 543.2 1869.2 1.64 1.59 0.62 0.62 13 JuLy 263.0 2.09 0.79 14 38.2 SeLember 0.71 1.411 303.1 8 35.4 12.7 5.068.L 0.94 0.41 10 October Novebr 13 719.5 1.42 0.56 9 January 134.2 1.30 0.59 19.9 79 becewbet Month 0.80 1.84 12.o 1(1 Iebcuaty 10 359.8 1.24 0.54 Match 3,981.0 1.01 0.43 12 25.5 April 0.21) 32.0 0.52 14 May th Appendix 3. Species density (birds/lOU ha), June 1978-May 1919, WeSt Sand tslsnd flaroh, Coluabia Flyer, Oregon. liontim Species Common Loon Double-crested cormorant Pelagic corworatit June July 1.4 5.3 2.2 19.9 1.6 Great blue heron Mallard l'intail 4.2 AuguSt 1.4 September 1.0 1.9 0.4 22.2 0.8 Green-winged teal EuropeSu wigeon &uiartcan wigeon Northern shoveler Greater aceup Buff lehead Red-breasted mnerganser Rough-legged hawk Marsh hawk Seamipalamated plover Kilideer Black-bellied plover 2.2 October Novewber 1.8 22.2 3.2 0.8 12.4 6.4 23.9 51.7 January February March April 4.8 31.4 46.9 0.9 1.7 1.3 0.7 17.1 1.0 0.6 0.8 17.5 10.3 1.4 16.1 0.8 66.5 24.1 2.1 2.8 24.5 1.1 19.9 38.2 0.4 4.4 2.2 0.2 0.8 6.4 3.2 9.5 6.4 0.8 12.7 8.8 6.8 2.6 65.1 3.2 2.6 155.9 417.8 234.7 16.2 6.4 811.3 406.8 44.2 1.4 Greater yellowlega Least sandpiper 2.2 11.2 204.1 135.5 Dumilin Dc,wttcher app. Western aandplper Herring gull California gull Ring-billed gull 41.7 12.9 213.7 32.5 23.0 17.4 1.6 11.9 Rufous hwaaiogbtrd Violet-green swallow Tree swallow 97.3 8.6 161.1 378.8 481.9 4.3 258.6 1.5 0.7 1o3.9 15.6 32.3 287 .6 245.2 3.2 2.4 21.1 2.8 0.8 4.2 3.2 15.3 1.8 0.2 9.1 19.6 318.3 10.8 60.0 1.4 21.5 28.7 1.4 23.4 1.4 4.3 4.1) 7.9 Mew gull Bonaparte's gull Caspian teru Belted kingfIsher 1.4 ho Whimbrel Sanderling GIaucou-winged gull Western gull Hay 1.4 1.4 113.9 21.5 1.6 2.8 Oecmbec 1.9 21.0 5.3 8.0 3.2 1.9 318.3 70.9 53.3 318.3 04.5 40.3 (contInued) Eve nueaa Number o spectea Density (birds/ha) Diversity American goldfinch Savaunab sparrow Song sparrow flouae finch Red-winged blackbird Common ylowtbroat St ant eg Long-billed uiarah wren Water pipit Comun crow barn awallow Cliff wmiIow Spectea 3. - AppendLX S59.5 1.58 0.56 19.9 5.7 6.4 39.8 8.8 June 11.9 4.8 20 868.1 16 329.3 2.09 0.75 11 517.6 2.03 0.72 14 564.1 1.64 0.62 0.67 1.99 i2.7 37.4 7.2 0ctobr 101.9 35.4 19.9 Seplember 28.7 August 39.8 79.6 25.5 9.9 39.8 39.8 5.7 30.9 0.4 4.8 19.9 July 1.53 0.57 1.55 0.59 7 161.6 0.77 0.40 13 43ó.3 0.66 0.26 14 655.4 15 ebruary 834.4 3.2 January 12.7 286.5 beceaber 35.4 9.7 November Hoth U.3 1.46 10 sOh.5 12.7 March 20 2,415.9 2.14 0.00 50.9 k2.7 3.2 17.7 april 1.ng 0.73 .5 13 0.5 10.1 2.4 9d.5 May 0' -4 Appendix 4. Species density (birds/100 ha), June 1978-May 1979, Miller Sands Spit Upland. Guheubta 1(iver, Oregon. Month Species June July Doubl-creuted cormorant August 1.4 Canads goose September October November 1.4 6.0 Snow goose 0.4 3.9 2.2 35.4 Common iaerganaer Bald eagle Semipalmated plover 1,170.3 280.5 110.6 13.9 308.2 28.3 139.5 631.4 Water pipit Diversity Evenness 31.2 43.6 9.5 11.9 6.8 1.0 8.8 242.1 18.1 4.2 4.2 42.7 103.7 113.2 7 1,317.5 1.44 0.74 3.9 67.') 1.0 62.2 11.2 44.8 1.9 12.7 1.4 35.4 2o .1) 9.9 6.4 39.2 11.0 9.9 24.9 15.9 263.6 1.5 4.2 4.2 I.9 9 12 496.6 3,357.8 1.46 0.88 0.66 0.35 9 10.9 1.48 0.68 1.8 4.4 6.3 33.4 14.4 8.4 59.1 51.3 119.4 3,565.8 370.4 2.1 1.6 Common crow Density (birds/ha) 0.1 11.1 2654.4 Common raven Number of Species 3.9 8.8 6.4 139.7 Mew gull Cliff swallow 1.0 91.3 Dun liii Morned lark Violet-green swallow Tree awallow barn swallow May 0.4 Least sandpiper Caspian tern AprIl 15.11 79.6 Whimbrel California gull tng-Stlied gull larch 5.7 50.9 0.1 0.1 Marsh hawk Herring gull ebivary 331.4 Crecu-winged teal Greater scaup buff 1ehed Western sandpiper Sauderling Northern phalarope Glaucous-winged gull Western gull 3.8 January 0.3 Pintali Kilideer black-befli ad plover December 3.2 3.1 0.4 0.1 0.8 245.6 145.6 109.2 56.6 18.2 36.3 11.7 35.4 9 4 142.7 0.83 0,60 1,844.3 1.01 0.46 1 10 4 8 21.8 1.23 0.63 213.8 1.69 0.74 67.5 1.21 0.87 100.2 1.65 0.79 9 10 651.6 4,u91.b 1.63 0.8/ 0.74 0.38 S. brown-hedd cowbird Starling Yellow-romped warbler Townsend's warbler Wileon'a warbler Western weadowlark Cedar wawtng Violet-green swallow Tree swallow Rant swallow Cliff swallow Common crow block-capped chickadee American robin Golden-crowned kiuglet Ruby-crowned kinglet Rufous huiai ngbird Common f licker Common nighthawk CaSpiaft tecn flew gull Piniall American wtgeoD Cooper's hawk Red-tailed hawk bald eadle Peregrine falcon Merlin American kestrel Common snipe Dunlin Dowiichter app. Western sandpiper Sanderling Glaucous-winged gull Western gull California gull Ring-billed gull Mal lard 26.5 8.0 12.7 19.1. 9.5 318.3 12.1 1.6 June 8.8 74.7 19.9 62.9 19.1 0.4 10.5 July Species density (birds/LOU ha) Crest blue heron Species Appeadtc 79.6 2.6 8.8 143.2 11.0 32.6 10.4 2.6 AuguSt 42.6 114.0 19.9 41.4 31.6 Septesber 1.0 25.5 12.1 1.0 12.7 25.5 [(.1 6.6 3.2 5.0 Noveabnr 12.7 22.0 65.9 25.5 12.1 3.9 2.2 13 0 October Month 3.9 1.4 12.1 151.2 2.4 Oeceldber 2.6 13.0 3.1 12.19 1.5 144.4 211.1 0.8 13.0 January 15.7 3.1 February tune 1918-May 1919, Miller Sands Uici. Coinebia lUver, Oregon. 5.3 2.6 8.8 5.3 15.6 81.5 March 623.3 140.8 bO.3 25.5 59. 1 3.9 35.4 35.4 19.5 April 21.1 35.4 bl.2 218.5 5.3 3.2 u.S Nay Appendix 5. (contInued) tIe at Species June July Aiserican goldfinch Savanoah sparrow Dark-eyed Junco White-crowned sparrow Song sparrow 7.9 448.2 358.1 Number of utpeetea Density (birds/ha) Diversity Evenness 60.3 August 3.9 89.1 6.5 537.1 38.2 IL 12 10 924.2 1.33 616.2 1.58 0.56 0.63 872.1 1.28 0.56 Septeather 35.0 October Nuveutber beceinber January Februaty Harct 33.7 6.4 toe.o Li.? 11.7 Hay 211.o 39.8 392.6 1.15 U 180.5 1.91 0.90 0.80 1 April 8 117.0 1.80 0.87 6 195.3 0.74 0.41 10 811.0 1.47 0.64 6 18.8 0.45 0.65 129.8 1,08 0.60 11 1211.7 1.65 0.69 8 630.8 1.36 0.ob 6. Cliff swallow Common raven Common crow Ruby-crowned kinglet Water piptt Starling Orange-crowned warbler Red-winged blackbird brown-headed cowbird House finch AmerIcan goldfinch Rufous-sided towhee White-crowned sparrow Song sparrow bgtii swallow Tree awllow American kestrel glildeer Spotted sandpiper Dunlin Dowitcher app. Claucous-witiged gull Western gull Herring gull California gull Ring-billed gull Mew gull Ceaptan tern Rufous hutsalngbtrd Downy woodpecker Western flycatcher Horned Lark Violet-green swallow Karb hawk 0.8 7.9 20.5 13.1 0.8 59.1 25.5 12.1 127.3 38.2 6.5 6.4 3.2 9.5 July 6.4 35.4 12.1 136.9 12.7 32.6 15.6 19.6 June 39.8 14.8 1.9 12.7 6.4 684.4 3.8 5.0 3.9 August 3.2 5.0 46.2 26.5 9.4 62.4 2.2 10.1 19.9 3.9 2.6 3.9 Septesiber 8.8 6.5 76.3 99.5 October Houth 636.6 '2.0 3.2 9.9 Oeceaber 5.0 13.0 2.8 *2.1 0.4 January ' 144.2 5.0 1.4 0.4 Lebruaty Upland. Coluwbia giver, Oregon. Novesber SpecIes density (birds/tOO ha), June 1918-Hay 1919, JIsi Crow Saad Canada gooae Snow goose Mallard Pintati Greater ucaup Red-breasted isergauser Species AppendIx u.S 5.1 19.9 5.9 '23.0 3.? 3.? ltarcli 6.5 lob.0 119.4 5.3 8.8 2.8 8.8 13.2 14.9 114.3 45.1 2.7 11.8 24.4 13.0 15.9 April 39.8 05.0 179.0 122.5 59.1 0.1 1.4 3.8 110.8 19.9 41.0 41.0 00.9 41.0 35.4 19.9 Hay A1ped1x 6. (coiittnud) Mouth July Steabur Spectea June Number of species Density (bIrds/he) DiversIty Evenness it 11 10 12 417.9 1.95 0.81 246.1 1.65 0.69 184.3 0.62 0.21 otl.9 1.18 0.48 AuMuat October 4 191.9 0.96 0.70 November 3 642.4 0.06 0.05 Decewber January 1 5 9.9 0 83.3 0.82 0.51 I) FeD&uary -4 bI.0 0.22 0.16 March 1 69.1 1.bS 0,86 Att1 ay lb 20 573.4 113e.b 2.Uo 2.30 0.74 0.77 N.) Appendix 7. Spccies density (blrds/l00 he), June 1978-May 1919, West Saud Island Uled Upland. Coluwbia 1ver, Oregon. Month Spedee Double-crested cormorant Great blue heron Mallard Red-tailed hawk Marsh hawk Bald eagle Merlin ICilideer June Rufous hu.waingbicd 0.8 Septeuber October Number of specIes Density (birds/ha) Diversity Evenness 1.6 Decesber 3.2 1.6 January February March april Play 2.4 6.5 2.4 15.1 1.6 3.2 13.0 1.6 2.4 0.8 5.0 8.8 16.4 1.4 7.1 61.3 2.8 19.9 3.2 4.7 0.8 12.7 35.4 5.9 2.8 17.9 2.8 10.0 6.5 10.0 88.4 3.2 79.6 636.6 19.9 1.0 19.9 2.6 1.3 1.3 11.9 19.9 36.2 22.3 114.6 7.9 4.8 159.2 3.2 79.6 10.3 238.7 3.2 2.2 59.7 19.9 1.3 39.8 8.8 389.0 i41. 75.2 84.4 2.o 8.8 4.2 3.9 0.8 19.7 59.1 9.9 58.5 5.3 19.8 71.5 3.1 233.9 6.2 2.6 2.6 75.2 IIue finch AmerIcan goldfinch Savannah sparrow .White-crowned sparrow Song sparrow Novewber 0.8 0.8 A,aerlcan robin Nottbern shrike Starling Western meadowlark Red-winged blackbird August 0.8 flack-bellied plover Glaucouw-winged gull Western gull California gull Ring-billed gui1 Mew gull Short-eared owl Coamon flicker Downy woodpecker horned lark Violet-green swallow Tree swallow Barn swallow ClIff swallow Purple martin Common crow Long-billed marsh wren July 3.2 62.9 397.9 12 314.4 601.9 12.8 10 721.0 14.8 239.7 212.2 3.2 37.4 39.0 8 1,415.9 1,113.8 1,368.7 1.50 1.24 1.32 0.60 0.5 0.63 12 :193.6 1.54 0.62 ii 272.8 1.88 0.78 298.1 32.5 11 520.6 (.06 0.44 3 34.8 7 123.1 1.08 0.55 14.9 8 267.6 0.54 0.26 39.8 7 193.6 0.81 0.42 20.0 7 212.9 1.25 0.64 119.0 229.2 39.0 11 449.1 1.11 0.49 6 592.1 1.45 0.81 8. Yellow-ruisped warbler Common yetlowthroat Starling Cedar waxwing Townsend's solitaire black-capped chickadee American robin Common crow Purple matttn Horned lark Violet-green awallew Tree swallow barn swallow Common fLicker Caspian tCrn bonaparte's gull Mew gull California gull Ring-billed gull Lesser ycilowlegs Sandarling Ciaucous-wlnged gull Western gull Herring gull Common snipe Whimbrel bald eagle Asierican keatrel Kilideer Marah hawk Sliarp-ahiiuied hawk Cooper's hawk Red-tailed hawk bufflehaad Pintail 10.1 21.5 38.6 15.9 44.6 11.8 5.6 19,9 35.4 17.0 2.8 31.4 June 17.7 4.0 31.8 19.1 19.1 9.5 1.3 10.5 14.0 2.8 0.2 26.0 1.6 July 19.9 2.6 3.2 251.1 6.4 2.2 11.3 48.1 6.9 1.0 3.9 1.9 3.6 Auguat 31.4 8.1 3.2 0.3 32.2 9.4 Sepi.iaber 3.9 6.6 0.4 9.0 6.4 19.9 13.0 3.2 5.3 5.0 October Nuvewber Month 9.4 4.2 u.S 0.8 1.4 5.0 12.5 0.8 I.! 0.2 Deceabet January iebruary Species density (blrd/I00 h.i), June 1978--May 1979, Weat Sand laland UplCnd, Coluwbia River, OreRwi. Double-crested cormorant Pelagic cormorant Mallard Common loon Species Appendix b.8 63.1 21.5 34.8 1.1 4.6 0.8 March 2.6 12.1 10.5 3.0 2.8 34.8 39.0 19.9 8.8 13.0 ApriL 23.1 34.2 4.2 29.6 86.7 1.9 35.4 6.4 ay 8. (cont:tnucd) Nwuber of species Density (birds/ha) Diversity Evenness Wilson's warbler Western meadowlarit House finch American goldfinch Savannah sparrow White-crowned sparrow Song sparrow Lapland iongspur SPeCIeS Appendix 0.50 1.42 16 765.8 1.38 0.50 17 688.0 381.1 16 2.1 15.7 184.1 1.70 0.77 9 6.5 0.10 36.2 1.13 S 3.2 October 55.0 457.6 5.0 54.5 Sepiewber 21.1 6.5 August 15.7 July 965.9 1.93 0.70 19,5 159.2 86.5 437.7 June 94.2 1.98 0.90 9 6.5 26.0 Novesber 12.0 0.52 0.75 2 2.6 Uccuwbct Month 0.83 0.47 6 70.2 2.6 54.7 January 9 16s.O 1.70 0.11 1 19.9 .5 hatch no.0 1.27 0.65 5.0 31.6 february 1.31 0.49 591. 15 2.7 6.5 4W.0 6.5 April 1.91 0.71 bTh.6 Ii Lj.0 59.1 193.6 May cormorant House finch Water pipit Conimon crow Cliff swallow Rough-winged swat low 8am swallow Horned lark Violet-green swallow Tree swallow Ronaparte'a gull CapJan tern Hew gull Herring gull California gull Ring-billed gull Northern phalarope Claucous-wtnged gnU Western gull Sande tl.tng Western sandpiper Llunitn DowttchCr ap. Kilideer illack-beltfed plover Ratrd'a aan4ptpar Least sandpiper Semipaluasted plover Raid eagle HarIi hawk White-fronted goose Canada goose Great blue heron I)ouble-creutsd 80.8 222.0 5.7 643.0 2.6 79.6 6.6 14.1 26.0 39.8 0.5 June 123.8 12.7 198.9 318.3 1.5 22.6 8.5 19.5 14.9 14.4 134.4 July 22.6 3,481.5 8.8 169.8 2.6 18.5 32.5 11.9 6.5 50.9 0.4 August 2.8 11.1 35.0 12.8 7.6 1.9 5.3 Septeauber 1.1 9.4 11.4 5.0 5.0 272.8 35.4 59.7 35.0 19.9 b.6 253.1 7.5 3.2 8.11 October 0.5 11:15.3 1,144.1 54.1 43.1 November Motit Ii 5.3 4.2 3.2 23.7 140.11 2.6 15.0 10.4 4.1 3.6 20.4 20.3 103.1 0.1 0.5 Veccsber January 5.1 3.2 13.5 12.n 13.2 79.t 0.1 1.9 February 9. SpecieS density (his/1011 ha), June 19711-Hay 1919, Hitter Sand Spit Iteah, C1utsh1a Rivet, Oon. Red-throsted loon Species Appendix 100.1 377.5 14.4 29.0 9.7 36.1 5.0 Hard, 82.0 100.5 620.1 26.3 210.1 5.0 14.13 03.1 209.9 1.6 0.4 April 1.4 191.0 23.e 2.6 2.2 19.9 13.0 1,021.0 75.0 691.4 1.0111.5 0.5 21.0 May -3 Appendix 9. (conLinued) HonLIi Species Number of Species Density (birds/ha) Diversity Evenness June It 1,120.7 1.35 0.56 Juiy 13 August 13 885.4 3,921.3 0.55 1.18 0.69 0.22 Septeisber 1 83.1 1.58 0.81 October 15 930.7 1.81 0.67 November 1 2,860.4 1.02 0.52 December 1 198.9 1.02 0.53 Jauuary 9 184.7 1.42 0.64 Februsry 1k) 112.2 f.ôi 0.11 Hatch 1 b18.0 1.42 0.51 April 13 Hay lb 2,148.4 3,loi.o 1.12 1.Sb (J.b7 0.b9 10. rganaer Rufous hummingbird belted kingfisher Bonaparte'S gull Caspian tern Mew gufl California gufl King-billed gull Herring gull Spotted sandpiper Least sandpiper Dunlin Dowitcher app. Western sandpiper Marbled godwit Sonderling Northern phalarope Glaucous-winged gull Western gull Iitildeer Semipalmated plover American kestrel Merlin Red-breasted urganser Bald eagle Coawon Ruddy duck American wtgeon Lesser scaup Greater aesop Bufflehead Green-winged teal Pintati Haltatd Wlntte-ttonted goose Canada goose Whistling swan Horned grebe Western grebe Double-created corworaint 70.7 18.4 56.6 105.8 69.4 June 35.4 25.5 11.7 29.1 241.3 5.0 3.2 7.1 July Species density (blrds/100 ha) ked-throated loon Spectea Appendix 181.1 21.8 21.8 ÔJ.4 49.4 2.6 3.9 35.4 19.1 August 3.2 8.0 5.0 5.0 1.8 3.8 8.8 Septesber June 1978--May 19/9k 23.7 3.5 22.0 42.1 U.3 3.9 8.8 October 1.4 3.8 8.1 316.5 22.6 7.9 5.0 Novewber Mouth 8.1 1.4 168.4 6.7 26.0 1.0 24.3 20.7 6.5 6.4 19.9 Uecewbcr 29.5 27.3 27.4 11.0 1.6 42.1 12.7 3.9 o.4 15.7 39 4.9 January 87.0 5.8 29.3 23.9 8.8 13.0 lebruary Millet Sands ticach, Coluahia River, 0reon. 318.3 58.7 13.5 4.7 14.9 15.7 9.9 24.9 19.9 March 3.9 No.7 14.8 23.2 37.9 2.6 129.9 n.5 7.3 22.3 7.2 730.1 l24j.1 32.5 U.) 39.8 May 9o3.8 08.2 19.9 5.0 5.0 2.6 19.9 39.8 April Appendix 10. (continued) Mouth June Specie.s Violet-green swallow Tree wa11ow barn swallow Clitf swallow Camta crow 66.3 26.4 July 35.4 437.1 99.5 18.8 August Septewbet 59.1 11.1 1.5 1.4 October Noveabet Dece4aber 2.8 January 1.4 February 9.9 M.rch 11.3 AprIL 144.7 38.4 34.8 21.1 19.8 79.6 21.0 35.4 35.4 19.6 Savannah sparrow Song sparrow (bttds/ha) Btverity Eveirnema 1.1 3.9 Amrtean goldfinch Density 12.1 127.3 79.6 hewick's wren Starling Brown-headed cowbird House finch Number of specIes May 7 413.6 1.83 0.94 13 11 LI 953.5 1.63 0.63 463.8 1.89 0.79 io9.1 1.67 0.10 12.8 9 154.6 1.85 0.84 1 365.3 0.59 0.30 12 14 323.1 1.ö6 0.67 261.3 2.15 0.8! 7 177.7 1.53 (1.79 10 491.1 1.34 0.58 19 2I)23.0 1.n4 0.56 In 260t.1) 1.59 0.57 II. Water ptpit Common crow Cliii swallow Casplan tern Rufous huaidagbtrd Horned lark Violet-green swallow Tree swallow Rough-winged awllw Barn swallow Co,sn tern Bonaparte's gull Mew gull California gull Ring-billed gull Herring gull 278.5 59.1 5.7 39.8 8.9 92.2 6.1 3.0 15.1 1.9 59.7 13.0 28.3 1.9 318.1) 159.2 2.2 6.5 80.6 8.5 269.7 123.0 54.6 5.0 6.5 17.1 12.1 2.4 46.3 11.0 14.1 103.9 2.1 4.4 10.4 44.4 2,217.2 23.9 168.5 5.3 12.3 247.6 5.7 114.0 9.4 26.0 1.7 IU.4 0ctbcr 6.0 0.9 13.2 50.8 0.4 Sep esber 126.3 79.0 August Northern phalarope Glaucous-winged gull Western gull 12.1 141.5 July Sander1ing Spotted sandpiper Baird's sandptpet Lcaat sandpiper Dunli Western sandpiper KIlideer Black-bellied plover Coman nrganser Scaipalmated plover Green-winged teaL American wigeon l'lntatl White-fronted goose Mallard Canada goose 9.9 June 2.8 2.1 124.3 105.0 1.9 1.0 00.5 3.9 3.8 1.9 39.8 November Month 14.1 3.6 3.3 8.1 233.6 5.3 1.5 7.1 33.2 Deceabet 7.1 28.1 35.4 17.5 115.18 1,340.8 3.2 January 1.4 141.5 1.8 3.4 15.5 20.0 25.5 February Species density (birda/i00 ha), June 1978-N..y 1979, JIm Crow Snd5 beach, Columbia River, Oregon. Great blue heron Horned grebe Common loon Species Appendix ).1 5.0 17.0 7.4 5.0 7.3 hi,rcli 4.2 12.) 43.2 18.4 27.4 10.2 21.0 32,506.2 b,43o.0 21.0 12.7 58.5 39.8 AFEII 7.1 191.0 t.9 )J.8 35.4 il.) 25.5 19.9 0.0 42.1 318.5 232.9 63.2 13.0 50.9 19.9 2.0 Hay ii. (coiiiuueJ) Nuwbar of apecieö Denatty (birda/ha) Diveraily Evenueaa Savannah aparrow Whtte-crowned aparrow Snow bunting Aertcan goldfinch Houae finch Specea Appendix 13 9.5 July 613.0 3,033.5 1.02 1.13 0.40 0.70 12 June 1.56 0.71 9 9511.1 2311.7 59.7 Augual 2.04 0.72 Il 8611.4 11 OcLobur 441.4 1.51 0.63 241.6 Sptewber (1.50 937.7 1.24 12 54.7 Nuvuwbur Mouth 328.2 1.13 0.49 10 l.4 Ueeeaber 1,554.3 0.57 0.29 7 January 0.66 1.31 334.5 8 119,4 Februat.y 14 Ajrii 19 141.5 59.7 May 41.9 39,327.1 1,341.6 1.67 0.51 2.38 0.93 0.19 (1.81 March Appendix 12. Species denaity (blcda/100 ha), June i918hay 1919, West Sand latand ileach, Colusbia kiver, Oregon. Species June August Steuther October Noveiuber Deceabet January 1ebruary March 10.5 PelagIc coraorant Great blue heron 2.6 42.6 8.6 14.9 0.1 35.4 24.2 176.8 44.2 8.11 9.9 8.8 5.0 17.1 99.5 5.0 50.9 $allard 3.9 Greater seaup Surf scoter Black scoter llldeer 8.8 2.6 4.6 12.7 18.8 1.9 9.s 13.0 Black-bellied plover 11.5 kuddy turnstone 2.8 Whirubrel 19.6 22.1 9.9 0.2 141,4 65.8 10.4 6.5 9.0 6.0 13.7 93.8 15.3 21.6 37.2 26.2 146.6 2.0 5.3 319.5 503.1 899.6 64.8 419.9 2.3 3.4 259.0 446.9 50.9 70.8 Hew gull Bonaparte's gull Heermann's gull Black-legged kittiwake Barn awallow 11.1 3.2 3.9 3.8 44.2 3.2 Harsh hawk Caspian tern Belted kIngfisher Violet-green swallow 3.1 8.8 7.9 White-winged acoter king-billed gull 12.7 14.7 Coeison goldeneye Bufflelread Herring gall California gull 5.0 50.9 39,11 Aruerican wigeon Western gull 5.0 141.5 Brant Glaucous-winged guLl flay 3.8 aed-throeted loon bowitcher app. Western sandpiper Sanderltng Northern phalarope iprtt 0.4 Coiswon loon Western grabs Doubie-eteated corworaut July 13.2 s8.5 1.4 5.2 6.6 41.1 2.3 4.11 63.2 201.4 £19.5 415.9 509./ 5.1 470.4 362.3 240.9 119.4 6.0 126.3 1.4 4.1 2.2 278.5 159.2 5.2 5.3 0.8 3.2 190.1 11.8 255 12. (continued) Number of species Density (birds/ha) DIversity Evenness Purple martin Cowman crow American robin Ruby-crowned kiuglet Water plpit Starting Rouse finch American goldfinch Savannah sparrow White-crowned sparrow Song sparrow Speclee Appendix 13 79.6 19.9 6.5 July 9 3.8 Auguat 1,047.5 1,325.2 1,299.2 1.55 1.10 1.15 0.76 0.60 0.50 10 14.3 23.6 35.4 June 1,033.5 1.20 0.47 13 2.1 2.8 September 0.74 1.99 977.5 15 39.0 203.7 4.2 October 681.6 1.29 0.54 11 2.8 Nuvewber Month 674.4 1.45 0.56 13 159.2 131.5 318.3 December 0.62 1.341 403.0 9 79.6 165.7 January 301.o 1.76 0.11 10 February 1/7.2 1.56 0.15 8 3.9 2.8 March 12 23.6 4.2 Hay 647.6 2,451.0 1.bZ 1.60 0.08 0.64 11 119.1 7.9 Apr11 13. Eveuneaa DiversIty l)eueity (birds/ha) Number of species Fox sparrow Song aparrew kutous-ided towhee Awericau goldfinch Brown-headed cowbird Black-throated gray warbler Cheataut-sided warbler Wjlaon's warbler Orange-crowned warbler Yellow warbler Starling Golden-crowned klnglet Ruby-crowned kinglet Varied thrush Swainon'a thrush Black-capped chickadee Winter wren bawick's wren American robin Coaaan crow Tree swallow Barn swallow Western flycatcher Downy woodpecker 0.74 1 467.1 1.44 1 676.6 1.84 0.95 119.4 1 159.2 19.9 12.1 219.4 59.7 127.3 12.1 179.1 3.9 August 135.8 59.1 99.5 35.4 July 649.5 1.71 0.91 119.4 114.6 42.4 99.5 70.) 99.5 3.9 June 1.32 0.55 1,1133.3 11 99.5 40.4 79.6 79.6 79.6 70.7 19.9 86.4 14.9 1,237.9 19.9 September 2.01 0.79 2,174.5 14 35.4 109.4 119.4 159.2 324.9 108.3 59.1 701.4 378.0 16.4 58.5 13.0 19.9 5.0 1.84 0.77 3,214.0 11 238./ 70.7 318.3 1,034.5 813.5 6.5 318.3 278.5 89.1 26.0 19.9 13 0.19 1.71 0.14 1.98 0.11 /19.0 10 617./ 10 136,9 1.83 15 1,544.6 2.17 0.80 278.5 79.6 170.2 59./ 59.7 26.0 9.9 11.7 8.8 49.5 106.1 76.4 25.5 35.4 214.1 198.9 3.2 10.5 35.4 216.5 167.1 39.8 2.2 u.4 19.5 53.1 3.8 March Febru,ry 99.5 106.1 35.4 6.4 218.8 59.7 2.2 35.4 3.2 January 39/.9 6.4 164.7 2/.9 22.3 99.5 159.2 79.6 2.6 35.4 31.6 3.2 Oscesaber Month free/Shrub. Columbia giver, Oreuzi. October Novewber Species density (birda/IQO ha), June l9/8-Mny 1979, Miller SaUd Bald eagle Merlin Glaucous-winged gull Western gull Rufous hisruingbird Coinaon flicker Harsh hawk Mallard Cooper's hawk Red-tailed hawk SpecIes Appendix 15 3u3.1 25.5 19.b 119.4 32.5 70.7 )ti.8 13.0 127.3 5.0 13.0 May 1,283.8 2.23 1,88 0.83 (1.78 1,021.9 11 84.0 308.4 10.0 /9.6 20.0 191.0 19.9 237.1 19.5 19.9 b.0 april 14. 21.5 59.9 19.1 216.5 179.0 19.9 42.7 25.5 318.3 79.6 127.3 139.3 218.8 39.8 50.9 139.3 19.9 106.1 39.8 159.2 141.4 99.5 119.4 1.4 39.8 106.1 70.7 July 83.7 59.7 318.3 June 99.5 39.8 19.9 59.7 12.7 12.7 59.7 232.1 25.5 3.2 8.8 5.9 5.9 79.6 August 75.3 14.6 212.2 /0.3 77.3 13.0 2.6 8.8 Septuaber 159.2 79.6 12.7 13.0 1.0 2/8.5 6.4 32.5 October 28.1 19.5 159.2 12.7 9.9 6.5 November Month 238.1 143.2 106.1 119.4 19.9 2.6 6.5 9.5 December 391.9 119.4 31o.3 159.2 424.4 198.9 /9.6 59.7 79.6 35.4 Jdnuary 198.9 10.1 33.4 30.8 419.4 3.2 Eebruary 19.o 19.1 212.2 26.5 53.1 123.4 35.4 25.7 159.2 26.0 12.7 March Species deneity (birds/lOU ha), June 1978-Hay 1979, West Sand Island Tree/Shrub, Cotuabia giver, Ore8ut,. Cooper's hawk Red-tailed hawk Harsh hawk Western gull California gull Rufous hummingbird Common flicker Downy woodpecker Western flycatcher Olive-sided flycatcher Violet-green swallow Tree swallow Barn swallow Steller's jay Common crow Black-capped chickadee Winter wren Rewick's wree American robin Varied thrush Swainson's thrush Golden-crowned kinglet Kuby-ccowned kinglet Cedar waxwtng Starling Rutton' wire.) Yellow-ruisped warbler Yellow warbler 8lack-thrnted gray warbler Wilson's warbler Brown-headed cowbird House finch American goldfinch Rufous-sided towhee White-crowned sparrow Golden-crowned sparrow ?ox sparrow Song sparrow Species Appendix 1o4.2 10./ 19.9 35.'. 79.6 57.3 35.4 4.9 10.1 44.2 26.5 05.0 305.6 2.2 50.9 391.9 19.9 ApriL 189.0 50.9 59.7 318.3 97.3 411. 31.8 139.3 8.8 8.8 58.5 0.5 195.0 20.4 1.4 1t.5 63.1 e. 79.0 May Ln ppendtx 14. (contInued) Month Spece Number of lpecles Deusity (btrd/ha) Dlver8tty Eveiinese June July 18 12 1,905.6 1,286.0 2.47 2.32 0.94 0.85 AuguL 14 665.0 2.06 0.18 Sepu*ber October 8 8 49.0 587.9 1.31 0.66 1.65 0.19 Noveibec Ueceiber January b 8 1.25 654.7 1.54 0.74 29.5 0.70 10 1,812.4 2.04 0.89 'ebruary MarcL b 11 462.4 1.43 116.8 2.03 0.80 u.8S ArII nay 18 20 2.31 0.82 2.32 0.78 i550.0 1,862.) 15. Golden-crowned kinglet Ruby-crowned kinglet Cedar wazwlng Northetn shrike StarlIng Hutton's vireo Orange-crowned warbler Yellow warbler Black-throated gray warbler Swatnoa's thrush Long-billed marsh wren American robin Varied thrush HermIt thrush Iiewick's wren tree swallow Barn swallow Cliff swallow Purple martin Common crow Stellar's jay Black-capped chickadee Winter wren Violet-green wailow Sharp-shinned hawk Cooper's hawk Red-tailed hawk Marsh hawk Peregrine falcon Kilideer Glaucous-winged gull Western gull California gull Caspian tern Short-eared owl Rufous hummingbird Common flIcker Downy woodpecker e.reen-..,Ineed teal 19.9 70.7 11.1 [.6 65.0 198.9 12.7 5.0 6.4 19.9 13.0 6.5 5.0 October 50.9 19.9 10.7 19.5 19.9 6.5 September 9.9 5.0 35.4 39.8 25.5 278.5 39.8 35.4 4.2 12.7 8.8 14.9 17.1 0.8 August 238.7 198.9 19.9 119.4 59.7 139.3 2.8 [9.6 25.5 2.8 198.9 45.5 305.9 318.3 318.3 35.4 2.6 1.4 11.8 July 2.6 June 15.5 318.3 12.1 389.0 59.7 6.5 Noveiber Month 12.7 - 19.6 2.2 6.6 December 99.5 495.1 218.5 99.5 59.7 12.7 9.5 4.4 19.6 1.9 3.2 3.9 January 9.5 35.4 32.0 28.9 35.4 19.6 99.5 33.2 1ebrusry 25.5 19.5 17.7 1.6 154.0 14.2 159.2 o.5 5.0 Mwrch SpecIes density (birds/IOU ha). June l918-t4y 1979, West Sand Island Transition, coluabia giver, Cun. Common loon Great blue heron Species Appendix 19.6 9.5 10.1 26.0 8.8 116.8 12.7 2.9 141.9 19.9 23s.1 3.2 1.9 April 35.4 22.3 19.9 45. 4.9 115.6 160.1 12.1 12.8 79.6 8.8 May 15. (continued) Density (birds/ha) Diversity Evenness Ruber of upecteS Wilson's warbler Northern oriole Western meadowiark brown-headed cowbird Rouse finch American goldfinch Rufous-aided towhee Savannah sparrow Datk-eyed junco White-crowned sparrow Golden-crowned sparrow ox sparrow Song sparrow Specie8 AppendiX 42.1 358.1 311.1 22.1 1/6.8 127.3 224.9 557.0 295.2 24.3 124.4 20 21 2,472.6 2,706.1 2.52 2.36 0.84 0.77 19.9 July 19.9 June 14 701.9 2.08 0.79 6 382.3 1.16 0.65 226.1 635.8 2.05 0.08 141 179.0 61.9 104.8 59./ 17.7 19.9 October 238.7 Se1.tewber 19.8 37.6 79.6 17.7 Auguet 1.35 0.69 941.0 1 139.3 November Month 0.18 1.87 984.9 11 19.6 103.4 39.8 35.4 37d.O i.o Decemauer 4,316.6 1.94 0.72 15 83.5 59.7 19.9 January 066.9 2.02 0.84 14 242.6 354 3b.4 FebLUSry 522.6 1.06 0.12 10 119.4 March lb 139.3 19.9 321.1 id.2 Hay 2,081.6 1,690.8 1.vb 2.46 0.73 0.12 19 159.2 1s9.2 141.5 71.1 19.9 3.2 A4jttL Appendix 16. Mean percent covera of each habitat characteristic for habitats comprising each island, Lower Columbia River, 19 78-79. Percent cover Habitat complex Moss! lichen Litter Bare ground Forbs Grass! sedge Miller Sands MSM MSSU MSU MSSB MSB MSTS 0.5 00b 43.3 00b 19.4 55C 585d 2.4 92.7 1.0 24.5 2.1 15.6 2.5 96.9 0.5 949 0.2 7.4 00b 12.1 60.8 2.2 0.0 2.5 95.3 5.5 84.5 14.8 58.2 2.8 36.8 22.7 2.3 79e 0.2 5.0 Jim Crow Sands JCSB JCSM l.5f csu 07g 01g 3.8 13.0 West Sand Island WSIM WSIDU wsIu WSIB WSIT WSITSI 25h 0.0 63h 0.0 21h 81h 56.2 t 346k 5.4 38.2 29.1 21.4 20.5 14.1 12.2 89.2 0.0 0.0 0.5 10.0 30.4 0.2 5.0 49.3' 90 Appendix 16. (continued) aData were transformed (arcsin IPT). bpercent cover of moss/lichen was not considered for MSSU:MSS1, NSSU:MSB, and NSSB:MSB comparison. CMS and MSB were similar. dBare ground was eliminated from analysis because of high correlation with moss/lichen (r -0.81) and litter (r = -0.81). epercent cover of grass/sedge for NSM was similar to MSU. percent cover of moss/lichen for JCSM was similar to JCStJ. cover of f orbs for JCSM was similar to JCSR. hpercent cover of moss/lichen was similar for WSIU:WSITS and WSIM:WSIT comparisons. Percent cover of litter ws similar for WSIN:WSIU, WSIDU:WSIT, and WSIT:WSITS comparisons. 3Percent cover of bare ground was similar for WSIM:WSIU comparison. kpercent cover of grass/sedge was similar for WSfl4:WSIDIJ, WSIM:WSITS, WSIM:WSIT, WSIDU:WSIT and WSIT:WSITS comparisons.