Direct-count estimates of fungal and bacterial biovolume in dead leaves of smooth cordgrass (Spartina alterniflora Loisel.)

Two types of hyphal-extraction, direct-count methods of estimating fungal biovalume in standing-dead, autumn leaves of Spartina alterniflora were compared with a clearing+staining method which does not require homogenization. Bacterial biovolume also was estimated, by an acridine-orange direct-count method. Type of homogenization had little effect on measured fungal volume, but counts made using water-soluble-aniline-blue epifluorescence were consistently lower than those made using phase-contrast (by 6–10x). Clearing+staining could not be used to estimate hyphal lengths, but was of use in estimating total ascocarp volume (=0.06 mm³ per mm³ of leaf). Estimated fungal hyphal volume was approximately 0.27 mm³ per mm³ of leaf. Bacterial volume was <3% of fungal volume.     

The introduction and spread of smooth cordgrass (Spartina alterniflora) in South San Francisco Bay

Spartina alterniflora was first introduced into south San Francisco Bay in the 1970’s. Since that time it has spread to new areas within the south bay and is especially well established at four sites. The spread of this introduced species was evaluated by comparing its vegetative and reproductive characteristics to the native cordgrass, Spartina foliosa. The characters studied were intertidal distribution, phenology, aboveground and belowground biomass, growth rates, seed production, and germination rates. Spartina alterniflora has a wider intertidal distribution than S. foliosa and outproduced the native cordgrass in all aspects that were studied. These results indicate that the introduced species has a much better chance of becoming established in new areas than the native species, and once established, it spreads more rapidly vegetatively than the native species. Spartina alterniflora is likely to continue to spread to new areas in the bay and displace the native plant. In addition, this introduced species may effect sedimentation dynamics, available detritus, benthic algal production, wrack deposition and disturbance, habitat structure for native wetland animals, benthic invertebrate populations, and shorebird and wading bird foraging areas. *** DIRECT SUPPORT *** A01BY058 00013     

Impact of high herbivore densities on introduced smooth cordgrass,Spartina alterniflora,invading San Francisco Bay,California

Spartina alterniflora, smooth cordgrass, invading San Francisco Bay, California (USA), is attacked by high densities of a plant hopper, Prokelisia marginata, and a mirid bug, Trigonotylus uhleri. Both herbivores are sap-feeders. We investigated the impact of these herbivores on S. alterniflora's growth rate, vegetative spread, and seed production by manipulating herbivore densities in the field and in a greenhouse. Herbivore densities in the field peaked in early fall, with P. marginata averaging more than 300 individuals per mature culm of S. alterniflora (about 100,000 per m²) and T. uhleri densities exceeding 10 per culm (about 3,000 per m²). Field reductions of herbivore densities by approximately 70% with insecticidal soap did not result in greater vegetative growth rates or lateral spread of plants; plants grew vigorously with the highest densities of insects. In the greenhouse study, conducted with seedlings, herbivory significantly reduced plant mass and tiller number in some but not all replicate herbivory treatments. In both field and greenhouse, there were significant differences between some clones' growth rates independent of herbivory. Inflorescence production in the field was not affected by reduced-herbivory treatments. Seed set was low under conditions of both natural and reduced herbivory, averaging 0.4%. Despite densities of P. marginata and T. uhleri that are much higher than typically observed in areas where S. alterniflora is native, herbivory by these particular insects appears to have little impact and in unlikely to limit S. alterniflora’s spread through San Francisco Bay. *** DIRECT SUPPORT *** A01BY070 00009     

Measurement of cordgrass,Spartina alterniflora,production in a macrotidal estuary,Bay of Fundy

A one-year field study was conducted of the growth, mortality, and loss dynamics of aSpartina alterniflora low marsh in the Minas Basin, a macrotidal estuary at the head of the Bay of Fundy. Data were used to examine the suitability of four methods for estimating annual net aerial primary production (NAPP) of a marsh subject to energetic tidal flooding. Shoots start to grow in April and reach maximum height (about 0.5 m) and weight in October. Maximum shoot density (900–1,600 m^?2) occurs around June and drops thereafter due to the export of entire shoots. The average shoot produces about seven leaves and at least 2–3 are lost during the growing season. All remaining vegetation dies before the end of November. Methods based on harvesting vegetation underestimated NAPP, especially at lower elevations where export is greater due to more frequent and prolonged tidal flooding. The highest NAPP values, on the order of 500–600 g m^?2 y^?1, were obtained using methods based on the population dynamics of individual shoots. These methods are recommended for energetic tidal environments because they include the production of vegetation exported during the growing season.     

Epiphytic nitrogen fixation associated with standing dead shoots of smooth cordgrass, Spartina alterniflora

N₂ fixation associated with the epiphytic community on standing dead Spartina alterniflora shoots was examined in both a natural and transplanted salt marsh in North Carolina. Acetylene reduction (AR) assays were conducted over a 24-mo period to estimate N₂ fixation rates on standing dead stems and leaves. In the natural salt marsh, mean AR rates ranged from 0.5 nmol C₂H₄ cm^?2 h^?1 to 14 nmol C₂H₄ cm^?2 h^?1, while in the transplanted marsh mean AR rates ranged from 1 nmol C₂H₄ cm^?2 h^?1 to 33 nmol C₂H₄ cm^?2 h^?1. Diel AR activity of epiphytic communities in both marshes varied seasonally. Midday incubations yielded higher AR rates than nighttime incubations in the spring, while midday incubations in late summer and fall generally yielded AR rates equal to or lower than nighttime incubations. Desiccation during low tides occasionally repressed AR activity, although AR rates quickly rebounded with wetting. AR activity was localized in the epiphytic community, rather than in the underlying Spartina stem material. Based on the measured AR rates and the density of standing dead stems, the annual input of new N to the natural salt marsh via epiphytic N₂ fixation is estimated to be 2.6 g N m^?2 yr^?1. The estimate of annual input of new N to the transplanted marsh is 3.8 g N m^?2 yr^?1. These estimates should be added to previous estimates of N₂ fixation in marsh sediments to estimate the total contribution of new nitrogen to salt marsh nitrogen budgets.     

Loss and recycling of amino acids and protein from smooth cordgrass (Spartina alterniflora) litter

We investigated the source and composition of free and protein-bound amino acids during the decomposition ofSpartina alterniflora Loisel in laboratory percolators and in a field experiment in the Great Sippewissett Marsh (Falmouth, Massachusetts). In the percolator experiment, 50% of the nitrogen (N) could be extracted fromS. alterniflora litter in 16 d. This extract consisted of dissolved free amino acid N (28%), suspended protein amino acid N (16%), inorganic N (12%), and nitrogen from unidentified compounds (44%). Much of the free amino acid nitrogen was utilized by detrital microorganisms, resulting in a greater loss of suspended protein amino acid (SPAA) nitrogen from the biologically active percolator due to microbials biomass. Suspended microbial mass accounted for at least 50% of the SPAA washed out of the biologically active percolator. In the field study, 38% of the original litter nitrogen was leached fromS. alterniflora litter in litterbags during the first 13 d. After this initial leaching period, the concentration (41% to 69% of total nitrogen) and composition of most amino acids bound in the litter did not change over the 23-month period of the experiment. Increases in microbial protein did not account for increases in total nitrogen which occurred during the decomposition of the litter. Similarly, adsorbed ammonium did not appear to be responsible for this increase.     

The relationship of smooth cordgrass (Spartina alterniflora) to tidal datums: A review

An analysis of data relatingSpartina alterniflora Loisel. to tidal elevations along the Atlantic and Gulf coasts demonstrated that although this species is primarily confined to the intertidal zone, its elevational limits. of occurrence do not correspond to a consistent elevation relative to a tidal datum in all marsh locations. The variation in the vertical distribution of this species reported among marsh studies was attributed primarily to differences in mean tide range (MTR). A positive correlation between MTR and elevational growth range (r=0.91) demonstrated that theSpartina alterniflora zone expands with increasing tidal amplitude. Differences in MTR among marsh locations accounted for 70 and 68% of the statistical variation in the upper and lower limits, respectively, ofS. alterniflora growth. Among marshes of similar tidal amplitudes, the upper limit of occurrence ofS. alterniflora in northern marshes was significantly lower than that in marshes at lower latitudes. These results, in combination with regional differences in plant species distribution across the upper intertidal zone, suggested that some of the variation in the upper limit was due to latitudinal differences in growth conditions and/or differences in interspecific competition. Local and regional differences in other factors such as salinity, nutrients, or physical disturbance may have also contributed to the variation in the limits of growth relative to a tidal plane within and among marshes.     

The effects of crude oil and remediation burning on three clones of smooth cordgrass (Spartina alterniflora loisel.)

Burning has been employed as an oil spill remediation technique in coastal marshes, even though the combined and interactive effects of oil and burning on vegetation are poorly understood. Variation among clones of perennial marsh grasses in response to these perturbations is not known. We performed a greenhouse experiment designed to assess the effects of Venezuelan crude oil alone and of oil followed by burning on three clonal genets ofSpartina alterniflora. The fully-crossed 6-mo experiment involved five dosages of oil (0 l m⁻², 4 l m⁻², 8 l m⁻², 16 l m⁻², and 24 l m⁻²) and two burn treatments (burned or unburned) applied to ramets from three clones. All oil-only dosages reduced survival, but burning after oiling (oil + burn treatments) increased survival relative to oil-only groups in all except the highest two oil dosages. Higher oil-only treatments also reduced ramet densities and inhibited density increases over 6 mo. Burning after treatment with the 16 l m⁻² oil concentration allowed increased production of new ramets, but burning exacerbated the negative impacts on ramet density at the oil concentration of 24 l m⁻². At some intermediate oil dosages, burning remediated the negative effects of oil on aboveground biomass production and growth in height. There was a significant effect of oil-only treatments on numbers of flowering ramets produced, in which two clones responded with decreased flower production and one exhibited increased flowering. There was no main effect of oil + burn on flowering. There were significant among-clones differences in all response variables to one or both treatments. Our experiment demonstrates that burning of oiledS. alterniflora marshes may have little measurable effect at low levels of Venezuelan crude oil, can remediate the effects of oil at intermediate oil concentrations, but can increase the negative impacts at high concentrations of oil. These results indicate that oil spills have the potential to adversely affect genetic diversity inS. alterniflora populations by eliminating some sensitive clonal variants or changing the relative dominance of genets. These results suggest certain clones may be better suited for phytoremediation or restoration planting following oil spills.     

Aeration,nitrogen and salinity as determinants of Spartina alterniflora Loisel. Growth response

A greenhouse experiment was conducted to examine the effects of salinity, nitrogen, and aeration on the growth of Spartina alterniflora Loisel. The experiment was conducted in a factorial arrangement of treatments with salt marsh substrate at three salinity levels (15, 30, 45‰), at two nitrogen levels (0 and 168 kg/ha) and at two aeration levels (zero and oxygen saturation). The maximum biomass was found in the low salinity, nitrogen enhanced, aerated treatment which had 11 times more biomass than the highest (45‰) salinity, nitrogen poor, unaerated treatment. the average effect of nitrogen over the three salinity levels was a 2.01, 1.47, 1.25, and 1.52 times increase in aerial biomass, density, height, and belowground biomass of the plants, respectively. The main effect of aeration was a 2.49, 2.01, 1.57, and 1.85 times increase in the same variables. The combination effect of aeration and nitrogen additions enhanced biomass by 453%. An increase in salinity from 15‰ to 45‰ decreased biomass, density, height and belowground biomass of S. alterniflora by 66, 53, 38, and 61%, respectively. The effect of salinity was more pronounced between 30 and 45‰ than it was between 15 and 30‰. N, P, K, Ca, Mg, Na, Fe, Mn, Zn, Cu, and S concentrations in the aerial living biomass were also examined. There was no evidence to suggest that elemental concentrations (on a per gram basis) were consistently correlated with increased or decreased growth. In relation to salinity, correlations between growth and elemental concentrations were negative while for nitrogen enhanced and/or aerated systems, the correlations were positive.     

The influence of subsurface hydrology on nutrient supply and smooth cordgrass (Spartina alterniflora) production in a developing barrier island marsh

The supply of nutrients from surface and subsurface water flow into the root zone was measured in a developing barrier island marsh in Virginia. We hypothesize that high production of tall-formSpartina alterniflora in the lower intertidal zone is due to a greater nitrogen input supplied by a larger subsurface flux. Individual nitrogen inputs to the tall-form and short-formS. alterniflora root zones were calculated from water flow rates into the root zone and the nutrient concentration corresponding to the source of the flow. Total dissolved inorganic nitrogen (DIN) input (as ammonium and nitrate) was then calculated using a summation of the hourly nutrient inputs to the root zone over the entire tidal cycle based on hydrologic and nutrient data collected throughout the growing season (April–August) of 1993 and 1994. Additionally, horizontal water flow into the lower intertidal marsh was reduced experimentally to determine its effects on nutrient input and plant growth. Total ammonium (NH₄ ⁺) input to the tall-formS. alterniflora root zone (168 μmoles 6 h^?1) was significantly greater relative to the short-form (45 μmoles 6 h^?1) during flood tide. Total NH₄ ⁺ input was not significantly different between growth forms during ebb tide, and total nitrate (NO₃ ^?) and total DIN input were not significantly different between growth forms during either tidal stage. During tidal flooding, vertical flow from below the root zone accounted for 71% and horizontal flow from the adjacent mudflat accounted for 19% of the total NH₄ ⁺ input to the tall-formS. alterniflora root zone. Infiltration of flooding water accounted for 15% more of the total NO₃ ^? input relative to the total NH₄ ⁺ input at both zones on flood tide. During ebb tide, vertical flow from below the root zone still accounted for the majority of NH₄ ⁺ and NO₃ ^? input to both growth forms. After vertical flow, horizontal subsurface flow from upgradient accounted for the next largest percentages of NH₄ ⁺ and NO₃ ^? input to both growth forms during ebb tide. After 2 yr of interrupted subsurface horizontal flow to the tall-formS. alterniflora root zone, height and nitrogen content of leaf tissue of treatment plants were only slightly, but significantly, lower than control plants. The results suggest that a dynamic supply of DIN (as influenced by subsurface water flows) is a more accurate depiction of nutrient supply to macrophytes in this developing marsh, relative to standing stock nutrient concentrations. The dynamic subsurface supply of DIN may play a role in spatial patterns of abovegroundS. alterniflora production, but determination of additional nitrogen inputs and the role of belowground production on nitrogen demand need to also be considered.     

Factors controlling aboveground Spartina alterniflora (Smooth cordgrass) tissue element composition and production in different-age barrier island marshes

Aboveground production and tissue element composition of Spartina alterniflora were compared in bareier island marshes of different age off the Eastern Shore of Virginia. The marshes were also characterized by physical and chemical parameters of the substrate. The results suggest that sediment nutrient stock do not directly control the spatial pattern of element content or production of S. alterniflora between these marshes. Elevated salinity likely limits the nitrogen uptake capability of S. alterniflora in the high marsh, which, in turn, controls leaf tissue nitrogen content of plants within individual sites. Low substrate redox potential may control the spatial pattern of nitrogen uptake between the different-age marsh sites, loading to more favorable growing conditions at the low stations of the young marsh sites where values of tissue nitrogen and production are highest. Tissue phosphorus did not differ between, or within the marsh sites. The result of a fertilization experiment suggest that nitrogen, and not phosphorus, is the primary limiting nutrient in this sytem. This indicates that nutrient limitation and other stresses work in conjunction to control tissue element content and macrophyte production at these marsh sites. Spatial variability of factors that control leaf tissue nitrogen and production is likely related to topography and grain size of an individual marsh, which is a function of marsh age. Most studies in different-age marshes have compared transplanted marshes to older, natural marshes. This work is one of few studies comparing developing and mature natural, marshes on barrier islands.     

Productivity of cordgrass,Spartina alterniflora,estimated from live standing crops,mortality, and leaf shedding in a Virginia salt marsh

Net annual productivity of tall and medium form cordgrass,Spartina alterniflora, was estimated by a new clip sampling method in a sloping foreshore salt marsh at Wallops Island, Virginia. This method measured live standing crops only, to avoid problems of measuring dead biomass inherent in other methods. Losses from live standing crops by shoot mortality and by leaf shedding were estimated from these measurements and added separately to production of live tillers and of live culms. This allowed quantification of various components of production.Spartina tillering in different zones of the marsh produced 62 to 211 g dry weight per m² per yr. Tiller mortality removed 37 to 106 g per m² per yr from live standing crops. Culms produced 348 to 1,132 g per m² before flowering and die-back. Culm mortality removed 28 to 246 g per m² before flowering. Leaf shedding removed an additional 83 g per m² in tall formSpartina. Altogether, net annual productivity These estimates are much higher than previous estimates of productivity and standing crops inSpartina marshes nearby.     

Growth dynamics of cordgrass,Spartina alterniflora loisel., on control and sewage sludge fertilized plots in a Georgia salt marsh

Seasonal plant growth dynamics were followed for a year in undisturbed plots of tall and short formSpartina alterniflora Loisel. and in plots of short formS. alterniflora which were enriched with sewage sludge at a rate of 100 g dry sludge m^?2wk^?1, corresponding to a nitrogen enrichment of 2 g N m^?2wk^?1. Monthly determinations of aboveground live and dead biomass, density of live stems, the ratio of number of young shoots to total number of shoots, and belowground mass of macro-organic matter to a depth of 30 cm were made for each area. Sludge fertilization increased the live biomass of the short formS. alterniflora by up to 150% of the control live biomass, but had little effect on the dead biomass, stem density, or proportion of young shoots. There was a trend of increased amount of belowground macro-organic matter in fertilized compared to control plots during the last 6 months of the study. In all areas, there was a marked decrease in the proportion of young shoots from winter to early summer, followed by a rapid increase in the percent of young shoots from late summer to fall. Sampling of plots 7 and 20 months after termination of sludge enrichment showed higher plant biomass and % N content in surface soils, but no difference in N content of live plant tissue, in fertilized compared to unfertilized marsh. After 20 months, about half of the sludge nitrogen remaining in the soils of the fertilized plots had disappeared.     

Mechanisms of expansion for an introduced species of cordgrass, Spartina densiflora, in Humboldt Bay, California

The dominant plant in Humboldt Bay salt marshes in Spartina densiflora, a species of cordgrass apparently introduced from South America. At several salt marshes and restoration sites around Humboldt Bay, distribution of this plant has increased significantly. We investigated the relative contributions of vegetative tiller production and seed germination to the establishment and expansion of S. densiflora. Lateral spread of plants surrounded by competitors were compared to areas without competing plant species. Plants growing in areas without competitors had significantly higher rates of vegetative expansion (p<0.0001). Viable seed production, germination rates, seedling survivorship, and growth of adult plants were measured in six salinity treatments. Approximately 1,977±80 viable seeds are produced per plant (0.25–0.5 m²). The number of germinating seeds was inversely related to increases in salinity. Salinity treatments between 19‰ and 35‰ produced significantly lower germination rates than salinities of 0–18‰ (p<0.0001). Seedling survivorship was 50% at ≤4‰ and 8–14% at ≥11‰. Lateral expansion of adult, greenhouse-grown plants occurred in all salinity treatments, with modest decreases in the highest salinity treatments (p<0.05). Our findings indicate that S. densiflora expands primarily by vegetative expansion, and lateral tillers are produced by throughout the year. Spartina densiflora produces prolific amounts of seed, but recruitment in mature salt marshes may be limited by competitors and higher salinities. At restoration sites, planting of native species such as Salicornia virginica, Distichlis spicata, or Jaumea carnosa may prevent monospecific stands of S. densiflora from developing.     

Vertical Distribution and Migration of Decapod Larvae in Relation to Light and Tides in Willapa Bay,Washington

We investigated the occurrence of behaviors that maximize predator avoidance and seaward transport in estuarine decapod zoeae by collecting larvae from discrete depths in a partially mixed estuary, Willapa Bay, Washington, USA, and relating their abundance and vertical distribution to a suite of environmental variables. Abundances of first zoeae of Neotrypaea californiensis and Pinnotheridae were associated with tidal phase, diel phase, and water height. Both taxa were most abundant during ebb tides, and abundances increased with water height, suggesting behaviors that enhanced seaward transport. Additionally, N. californiensis were both shallower and more abundant at night, indicative of behaviors to avoid visual predators. Our results suggest that both tidal transport and predator avoidance are important and sometimes interactive selective forces shaping larval decapod behavior.     

Factors influencing spatial and annual variability in eelgrass (Zostera marina L.) meadows in Willapa Bay, Washington, and Coos Bay, Oregon, estuaries

Environmental factors that influence annual variability and spatial differences (within and between estuaries) in eelgrass meadows (Zostera marine L.) were examined within Willapa Bay, Washington, and Coos Bay, Oregon, over a period of 4 years (1998–2001). A suite of eelgrass metrics were recorded annually at field sites that spanned the estuarine gradient from the marine-dominated to mesohaline region of each estuary. Plant density (shoots m^?2) of eelgrass was positively correlated with summer estuarine salinity and inversely correlated with water temperature gradients in the estuaries. Eelgrass density, biomass, and the incidence of flowering plants all increased substantially in Willapa Bay, and less so in Coos Bay, over the duration of the study. Warmer winters and cooler summers associated with the transition from El Niño to La Niña ocean conditions during the study period corresponded with this increase in eelgrass abundance and flowering. Large-scale changes in climate and nearshore ocean conditions may exert a strong regional influence on eelgrass abundance that can vary annually by as much as 700% in Willapa Bay. Lower levels of annual variability observed in Coos Bay may be due to the stronger and more direct influence of the nearshore Pacific Ocean on the Coos Bay study sites. The results suggest profound effects of climate variation on the abundance and flowering of eelgrass in Pacific Northwest coastal estuaries.     

Genetic effects of a large-scaleSpartina alterniflora (smooth cordgrass) dieback and recovery in the northern Gulf of Mexico

A large-scale dieback event struck marshes along the northwestern Gulf of Mexico coast during summer 2000, in apparent response to a prolonged and severe drought. Along the Louisiana coast, large areas of the dominant marsh species,Spartina alterniflora, turned brown, followed by death of at least the aboveground structures of entire plant mortality. Key ecological and genetic measures were studied in a dieback-affected marsh in southwest Louisiana (C83 marsh, Sabine National Wildlife Refuge), for which existed predieback ecologic and genetic datasets. Effects on genetic diversity only were studied in a second set of sites in southeastern Louisiana (near Bay Junop), where the dieback was more widespread. We hypothesized that stem density, live aboveground biomass, and genetic diversity would be significantly reduced compared to predieback conditions and to nearby unaffected marshes. Stem densities and biomass levels approached predieback conditions 14 months after first observance of the dieback in the Sabine marsh and were similar to or exceeded the same measures for a nearby unaffected marsh. DNA extracted from leaf samples in the Sabine and Bay Junop sites was used to construct genotype profiles using AFLPs and analyzed using the complement of Simpson’s Index (1-D), the richness measure G/N, average heterozygosity <H>, and the estimated proportion of polymorphic genes <P>. Genetic diversity was relatively unaffected by the dieback at either the Sabine or Bay Junop sites. Evidence from field observations and the results of the genetic analyses suggest that seedling recruitment is an important factor in the recovery of both the Bay Junop and C83 sites, although re-growth from surviving below-ground rhizomes appeared to dominate recovery at the latter site. Survival of below-ground structures, leading to the rapid recovery observed, indicates a high level of resilience of the Sabine marsh to drought-induced stress. Still, the genetic diversity ofS. alterniflora-dominated marshes may be promoted by occasional disturbance events, which produce open areas in which seedling recruitment can occur.     

Use of phytoplankton species indicators to track the origin of phytoplankton blooms in Willapa Bay, Washington

We focus on the question of whether high phytoplankton production events observed in a United States Pacific Northwest estuary consist of estuarine species blooms fueled by oceanic nutrient input or reflect offshore oceanic blooms that have advected into the estuary. Our approach is to use certain phytoplankton species as indicators associated with water mass origin, either estuarine or oceanic, to help resolve this question in Willapa Bay, Washington. We used species analysis and primary production data from 10 selected dates in May–September of 1998 and 1999, representing periods of high through low productivity. Out of 108 phytoplankton species identified from Willapa Bay, nine were selected and tested as indicators of oceanic species, six as estuarine, and two as surf zone. Our test results demonstrated the oceanic and estuarine species to be satifactory indicators of source waters. The prevalence of these species indicators in our samples revealed that the highest primary production and the appearance ofPseudo-nitzschia spp. were associated with oceanic intrusions of phytoplankton biomass into Willapa Bay. While the largest blooms were oceanic in origin, numerous medium-sized production events were from either oceanic, surf zone, or estuarine sources, indicating a complex situation.     

Heavy-mineral distribution in modern and ancient bay deposits,Willapa Bay,Washington, U.S.A.

Analysis of heavy-mineral distribution in modern sediments of Willapa Bay, Washington, indicates a dominance of two mineralogic assemblages, one with approximately equivalent amounts of hornblende, orthopyroxene and clinopyroxene, the other dominated by clinopyroxene. The hornblende-orthopyroxene-clinopyroxene suite is derived from the Columbia River, which discharges into the ocean a short distance south of the bay. The clinopyroxene suite is restricted in modern sediments to sands in rivers flowing into the bay from the east. The heavy-mineral distributions suggest that sand discharged from the Columbia River, borne north by longshore transport, and carried into the bay by tidal currents accounts for most of the sand within the interior of Willapa Bay.Three heavy-mineral assemblages are present in the surrounding Pleistocene deposits; two of these are identical to the modern assemblages described above. These heavy-mineral assemblages reflect the relative influence of tidal and fluvial processes on the Late Pleistocene deposits; their relative influences are consistent with those inferred on the basis of sedimentary structures and stratigraphic relations in about two-thirds of the samples examined. The anomalies can be explained by recycling of sand from older deposits. The persistence of the two heavy-mineral assemblages suggests that the pattern of estuarine sedimentation in Late Pleistocene deposits closely resembled that of the modern bay.The third heavy-mineral suite, dominated by epidote, occurs in a few older Pleistocene units. On the north side of the bay, the association of this suite with southwest-directed foresets in crossbedded gravel indicates derivation from the northeast, perhaps from an area of glacial outwash. The presence of this suite in ancient estuarine sands exposed on the east side of the bay suggests that input from this northerly source may have intermittently dominated bay deposition in the past.