an overview of algal monitoring and research in the u.s
TRANSCRIPT
An overview of algal monitoring and research in the
U.S. Geological Survey's National Water Quality
Assessment (NAWQA) Program
Julie A. Hambrook Berkman and Stephen D. Porter
Abstract
The National Water-Quality Assessment (NAWQA) Program of the U.S. Geological Survey uses algal data together with other physical, chemical, and biological variables to assess the quality of rivers and streams throughout the United States. Pilot NAWQA stud-ies (1987-1990) led to the development of methods for collecting algal samples from vari-ous instream habitats and to the publication of protocols for the identification and enu-meration of algal samples to ensure nationally consistent data. During the first decade of the Program, more than 7,000 algal samples were analyzed from more than 1,500 river reaches throughout the United States. These data from the NAWQA Program and related studies have been incorporated into more than 70 publications, 50 of which are cited in this paper. Determinations of algal biomass (algal biovolume, chlorophyll a, ash-free dry mass), autecological-guild and pollution-tolerance metrics, indicator taxa, physiognomy, and weighted-average optima and tolerance were used in evaluations of stream-water quality, depending onthe focus of each study. Relations of the biological datawith trophic condi- tion, hydrologic disturbance, geochemistry, and land use were examined. Although species optima and tolerance values for diatoms differ depending on the scale of assessment (na-tional and regional), species optima determined for indicators of alkalinity, water hardness, specific conductance, chloride, and sulfate were consistent with qualitative, autecological classifications for those taxa in western Europe. The ecological studies summarized here are part of nationally guided studies addressing selected water-quality issues, such as the effects of watershed urbanization and agricultural land use on nutrient enrichment and stream ecosystems. The algal data and related physical, chemical, and biological data are released (after quality reviews) and made available to the public online through the NAWQA website ht-tp : //water.usgs.gov/nawqa/.
Key index words : algae, autecology, disturbance, eutrophication, land use, water quality
Introduction
The U.S. Geological Survey (USGS) has been investigating algae in rivers, lakes, and reservoirs since the 1960s. These studies led to the publi-cation of methods for collection and analysis of samples (Greeson et al. 1977), a key to the phy-toplankton of North America (Greeson 1982) , and the addition of a biological laboratory for the identification and enumeration of aquatic or-
ganisms as part of the National Stream Quality
Accepted on September 2, 2004
Accounting Network (NASQAN) program. The
expertise in collecting and analyzing biological
samples during the 1970s and 1980s helped pro-
vide the foundation for developing the National
Water-Quality Assessment (NAWQA) Program.
The NAWQA program was established by the
U.S. Congress in 1986, with the following long-
term goals : (1) to provide a nationally consistent
description of current water-quality conditions for
a large proportion of the Nation's water re-
sources, (2) to define long-term trends (or lack
of trends) in water quality, and (3) to identify,
describe, and explain the major factors that af-
14 Julie A. Hambrook Berkman
feet observed water-quality conditions and trends
(Cohen et al . 1988 ; Hirsch et al . 1988). More information about the NAWQA Program can be found at http : //water.usgs.gov/nawqa/. Pilot studies for this program including algae monitor-ing (1987 — 90) were conducted in the Kentucky River Basin (Bradfield & Porter 1990, Stevenson & White 1995, Haag & Porter 1995), the Upper Illinois River Basin (Leland & Porter 2000) , and
the Yakima River Basin (Leland 1995 ; Cuffney et al . 1997). During the first decade of the fully
implemented NAWQA program, studies employ-ing chemical, physical, and biological components were conducted in 51 surface- and ground-water basins throughout the United States.
Design and summary of algae collection,
processing, and reports
Algae collection and sample processing Following the full implementation of the Pro-
gram in 1991, NAWQA has focused on the col- lection of periphyton samplesfrom erosional and depositional habitats in streams and rivers of
United States (Porter et al. 1993, Moulton et al. 2002) in order to characterize algal assemblages
in representative stream reaches. For NAWQA studies, a reach is defined as a portion of a stream whose length is approximately 20 times the width of the stream channel (Meador et al. 1993, Fitzpatrick et al . 1998). Quantitative sam-
plesfrom erosional habitats (RTH or richest- targeted habitat samples) were collected princi-
pallyfrom epilithic microhabitats in stream riffles andfrom epidendric microhabitats (submerged woody debris) in stream runs. About 67% of the 2,860 RTH samples collected through 2003 were from epilithic microhabitats and about 28% were from epidendric microhabitats. Quantitative sam-
ples (1,885 samples)from depositional habitats (DTH or depositional-targeted habitat samples, e. g. silt or sand) were collectedfrom stream pools or margins where water velocities were slow or not measurable (Porter et al. 1993, Moulton et
al. 2002). Algae samples were typically identified to species or variety for diatoms and genus for soft algae. Biovolumes were determined for all algal taxafrom morphometric measurements of representative individuals. Results were presented as standing crop (cells per square centimeter,
cells/cm') and biovolume (cubic micrometers per
and Stephen D. Porter
square centimeter,1.1m3/cm2) on the basis of laboratory protocols developed forthe NAWQA
program (Charles et al . 2002). Determinations of chlorophyll a (Arar & Collins, 1997) and ash- free dry mass data(American Public HealthAs- sociation et al. 1998) also are available for many samples, particularly those collected after 2000 and thosefrom selected synoptic studies prior to 2000 (e.g. Sorenson et al. 1999, Peterson et al.
2001).
Qualitative multi-habitat (QMH) algae samples were collected at monitoring sites during the first cycle ofthe NAWQA Program (1991-2001). Morethan 2,000 QMH samples were obtained
by collectingfrom each microhabitat (epilithic, epidendric, epiphytic, epipsammic, and epipelic)
prevalent in the stream reach and compositing equal amounts of biomassfrom each microhabi-
Lat. The preservative was 3 to 5% buffered forma- in (Porter et al. 1993, Moulton et al. 2002 http :
//water.usgs.gov/nawqa/protocols/OFRO2-150/in-dex.html). Algal taxa identified from RTH, DTH,
and (or) QMH samples from each site and date were combined into a master list of species for
each site. More than 500 phytoplankton samples (PHY)
were collected from reservoirs (Bauch & Malick 2003), large rivers (Kroening et al. 2003, Leland
& Frey in review 2004), and (or) small streams
(Stevenson & White 1995, Sorenson et al. 1999, Leland et al . 2001). These samples were taken from the same depth- and (or) width-integrated
samples collected for water chemistry. Subsam-
ples were processed for chlorophyll a determina-lions and (or) species identification and enumera-fion. Twelve workshops on harmonization of taxon-omy of diatoms (10) and soft algae (2) were
held to resolve differences in nomenclature and regional variations. During the first workshop a
list of 500 names and authorities, representing all diatom taxa at >1% abundance in NAWQA
samples, were examined in detail to facilitate data consistency among taxonomists and institu-
lions (ANSP 1999, http : //www.acnatsci.org/re-search/peer/research. html)
hops focused on reducing
ating a master list of 1,131
in 2004) with identification
image citations, establishing
Successive work-"unknown taxa"
, cre-taxa initially (>5,000
numbers and specific criteria for adopting
new names, incorporating the changes to the da-tabase (ANSP 2000a & b), and identifying the
problematic taxa within Navicula and Gompho-nema (ANSP 2001). At present (2004), one new
genus and nine new diatom species have been described on the basis of NAWQA collections
(Kociolek & Kingston 1999, Morales, 2002, Pota-
pova et al. 2003, Potapova & Ponader 2004).
Reports Algae samples collected and analyzed by the
U.S. Geological Survey scientists and associates have resulted in more than 70 publications since 1990. These publications are largely funded by the NAWQA Program but some represent related
projects using methods developed in the Pro-gram. These publications range from data sum-maries to interpretative reports, including discus-sions of community responses to land use (from forest to agriculture and urban), physical habitat
(e.g., as riparian shading, width, slope, current velocity), and water-quality (e.g., nutrients, salts, metals, pesticides, and organic matter).
The periphyton was the principal subject of most of the reports (85%), but sixteen included information on the phytoplankton. Seven of these reports focused on large rivers, lakes, or reser-voirs, where phytoplankton was the primary com-
ponent of plant production. Publications on phy-toplankton range from data reports (Averett et al. 1993) to seasonal and spatial analyses of species
/ environment relations (Stevenson & White 1995). Nine studies included phytoplankton as
well as the benthic algae in order to account for whole stream production in response to gains and losses in nutrients. Analysis of algal biomass as chlorophyll a, ash-free dry mass, or biovol-ume, and the resultant metabolic rate of the community as estimated by diurnal dissolved oxygen was used to evaluate varying land use
and riparian cover conditions (Sorenson et al. 1999, Peterson & Porter 2002). In addition, two
papers on diatoms from reservoir sediment cores evaluated changes in environmental conditions and land-use practices over recent decades
(Bradbury & Van Metre 1997, Juracek 2003).
Results and Discussion
During the first decade of the NAWQA pro-
gram, more than 7,000 algae samples were ana-lyzed from more than 1,500 river reaches throughout the United States (Fig. 1). The algal data from the NAWQA Program and related studies have been incorporated into more than 70 publications, 50 of which are cited in this pa-
per. Estimates of algal biomass (total algal bio-volume , chlorophyll a, ash-free dry mass) ,
autecological-guild and pollution-tolerance metrics, indicator taxa , physiognomy , and weighted- average optima and tolerance calculations were all used to evaluate the condition of streams and rivers.
16 Julie A. Hambrook Berkman
Indicators of trophic condition Many approaches mentioned above have been
used to evaluate the trophic condition of streams and rivers. For example, streamflow in the semi-arid southwestern United States often is domi-nated by point-source discharges of wastewater effluent with high loads of nitrogen and phos-
phorus, and riparian shading along southwestern desert streams generally is sparse or absent. A combination of excessive nutrient loading, rela-lively stable stream hydrology, and abundant light availability has resulted in accelerated eu-trophication as indicated by chlorophyll a values exceeding 300 mg/m2 in effluent-dependent streams
and values ranging from 4.8 to 45 mg/m2 in streams not influenced by wastewater discharges
(Gebler 1999). Periphyton biomass and productiv-ity in the Yellowstone River basin (Northwestern Great Plains ecoregion) were influenced by nonpoint-source (agricultural) nutrient enrichment as well as by natural, geothermal discharges of dissolved nitrogen near Yellowstone National Park (Peterson & Porter 2002). Results from this study showed close correspondence among vari-ous indicators of periphyton biomass and stream trophic condition, including microalgal chlorophyll
a, ash-free dry mass, total algal biovolume, macroalgal dry mass, estimates of system pro-ductivity and several autecological metrics (e.g.,
percentage of eutrophic diatoms and nitrogen- uptake metabolism) (Peterson et al . 2001). In New England coastal basins (northeastern United States) nutrient and chlorophyll concentrations were determined by Riskin et al. (2003) from reference streams, moderately impaired streams, and impaired streams all with and without tree canopy (shading). Algal biomass increased with nutrient impairment and was greater in all stream classes (reference-impaired) without tree canopy, where more light was available for plant
growth. Methods for establishing threshold con- centrations of nitrogen and phosphorus were pre- sented where the allowable or desired concentra- 1 tions of nutrient (nutrient criteria) and chloro-
phyll a could vary by the designated use or ex-isting quality of a water body. 11
Autecological-guild and pollution-tolerance met-
were used in many NAWQA and related re- containing algal results . Examples of
autecological-guild approaches included the use
and Stephen D. Porter
of qualitative species classifications by van Dam et al . (1994), Bahls (1993), VanLandingham
(1982) , Lange-Bertalot (1979) , Lowe (1974), Palmer (1969), and other researchers. The per centage of algal species associated with each au-
tecological guild (e.g., percentage of eutrophic, nitrogen-autotrophic, nitrogen-heterotrophic, halo-
philic, and other diatom guilds) was calculated, and these metric values were compared among sites (e.g., Peterson & Porter, 2002, Wynn et al. 2001, Deacon & Spahr 1998). An "index of bio-logical integrity" (IBI) approach for diatoms also was used in several studies (e.g., Scudder & Stewart 2001). Bahls' (1993) IBI approach in-volves calculation of the Shannon-Wiener diver-sity index, a classification of pollution-tolerant species (cf. Lange-Bertalot 1979), and a "silt in_- dex" based on the sum of the percentages of Navicula, Nitzschia, Surirella, and Cylindrotheca species and varieties. Bahls' silt index was used
to compare algal status in a number of streams
sampled between 1996 and 1998 (NAWQA Sum-mary reports 2000, http : //water.usgs.gov/nawqa
/nawqa_sumr.html). Although the relations be-ween the silt index and indicators of stream sil-
tation processes (e.g., substrate embeddedness
and suspended-sediment concentrations) were
relatively weak for stream sites within study ar-
eas, correspondence was close between the in-
dex and sites characterized by elevated concen-
trations of dissolved nutrients. Many of the
Javicula and Nitzschia taxa that strongly influ-
enced metric values also are indicators of eu-
rophic conditions.
The relative abundance of nitrogen-fixing algae,
blue-green algae with heterocysts (Bold &
Wynne 1985) and diatoms in the family Epithe-
miaceae known to contain endosymbiont blue-
green algae (Geitler 1977, Fairchild et al. 1985), vas found to be a useful autecological indicator
or water-quality metric reflecting low concentra-ions of dissolved nitrogen in streams and rivers.
The abundance of nitrogen fixers often was rela-
tively large in forested or rangeland streams
with low concentrations of dissolved nitrogen and
low Nitrogen/Phosphorus (N/P) ratios in the
western United States (Leland 1995, Cuffney et
al. 1997, Munn et al. 2002, Peterson & Porter
2002), and weighted-averaged optima and toler-
nce values for dissolved inorganic nitrogen were
Algae and the USGS
very low (Munn et al . 2002). Therefore, domi-nance of nitrogen fixers in western forested and rangeland streams frequently can be an indicator of low nutrient concentrations and good water
quality. A similar water-quality interpretation was attributed to northern streams (Wisconsin) domi-nated by nitrogen fixers, specifically Calothrix pa-
rietina (Scudder & Stewart 2001). In the southern United States, nitrogen-fixing
taxa were equally abundant in agricultural and forested basins (Petersen & Femmer 2003), and although the abundance of nitrogen-fixing dia-toms (Epithemiaceae) increased as ambient con-centrations of dissolved nitrogen decreased, the abundance of Calothrix parietina was unrelated to concentrations of nitrogen and phosphorus. Similarly, populations of Calothrix , but not nitrogen-fixing diatoms, were predominant in nearly all urban streams sampled in eastern New
Jersey (Kennen & Ayers 2002). However, the abundance of Calothrix was not correlated with concentrations of dissolved nutrients. In nutrient-rich streams and rivers, nitrogen-fixing algae
often become dominant only after extended peri-ods of stable, low streamflow (East et al. 1998, Leland et al. 2001), possibly reflecting seasonal
depletion of dissolved nitrogen from algal and macrophyte uptake. In those cases, dominance of nitrogen-fixing algae most likely indicates low N/ P ratios in the water rather than low nutrient concentrations and near-pristine water quality.
Nitrogen-fixing diatoms rarely were found in abundance in urban streams (Leland & Porter 2000, Kennen & Ayers 2002) or in Midwestern agricultural streams with high nutrient concentra-tions (Porter 2000, S.D. Porter, unpublished data). Because dissolved nitrogen generally is detect-able in water samples from these streams during sampling-index periods, eutrophic algal species
probably are strong competitors for space and other resources in these streams. Conversely, the studies along a gradient of disturbance from forest (western and northern United States) or rangeland to increasing human activity, found
that the dominance of nitrogen fixers can be an indicator of low nutrient concentrations and good water quality.
NAWQA Program 17
Indicators of hydrologic disturbance and
grazing The physical structure of periphyton communi-ties, including size, stature, and mode of attach-ment of predominant algal species (physiog-nomy) in the community, has been used as an indicator of hydrologic disturbance (Hambrook et al . 1997, Porter 2000) , differences in stream ve-locity (Scudder & Stewart 2001), and the influ-ence of algal grazers (Petersen & Femmer 2003). Coles et al. (2004) reported that the percentage of filamentous, stalked, and erect diatoms in-creased greatly with urban intensity and stream depth in coastal New England streams, whereas the percentage of Achnanthidium minutissimum, a small prostrate species, was much larger in shallow streams with low urban intensity. Porter
(2000) reported a dominance of small Achnanthes, Achnanthidium, and Fragilaria species in upper
Midwest streams and rivers subjected to re-
peated episodes of rainfall, runoff, and high streamflow. Hambrook et al. (1997) found that
the percentage of stalked algal cells in periphy-ton communities decreased significantly in Big Darby Creek, Ohio, following storms and that
prostrate diatoms and algae with holdfasts (e.g., Audouinella hermannii) were relatively resistant to washout. The prominence of prostrate diatoms in the periphyton community structure collected after storm events (Hambrook et al. 1997, Porter 2000) may provide a useful screening tool as an indicator of hydrologic (or grazing) disturbance for investigators who wish to ensure that they have collected an undisturbed periphyton sample
(e.g., for developing chlorophyll and nutrient re-lations).
Indicators of geology and land use
The presence and abundance of diatom taxa that are tolerant to or have water-quality require-ments for dissolved minerals, especially for major ions such as calcium and magnesium (alkaliphilic species) or chloride and sulfate (halophilic spe-cies) can be a useful indicators of natural, geo-chemical variability across ecoregions of the United States (e.g., Leland 1995, Brown & Short 1999, Potapova & Charles 2002, 2003), as well as an indicators of urban land use (Coles et al . 2004, Leland & Porter 2000) or agricultural land use (Cuffney et al. 1997, Carpenter & Waite
18 Julie A. Hambrook Berkman
2000, Leland et al. 2001, Munn et al. 2002). At a national scale, Potapova & Charles (2003) re-
ported that specific conductance and components of alkalinity (calcium, carbonate, and bicarbonate) explained the largest proportion of variation in diatom data. The national distribution of indicator taxa (e.g., alkaliphilous, acidophilous, and circum-neutral taxa ; hard-water and soft-water taxa) was associated with major differences in geology and
geochemistry (e.g., carbonate bedrock in eastern regions in contrast with shale, sandstone, or
granitic lithology in western regions). Similarly, geochemical sources of chloride and sulfate vary regionally, in association with natural landscape features of the humid southwest, Mississippi delta, and soft-water coastal rivers in the south-eastern United States, as well as in regions influ-enced by mineral extraction, notably coal mining. Increases in human activities such as urbaniza- tion (Groschen et al. 2004, Fischer, et al. 2004, Robinson et al. 2004, Rowe et al. 2004) and in-
tensive agricultural practices (Carpenter & Waite 2000) also increase the loads and concentrations of dissolved salts in streams and rivers, as indi-cated by increases in specific conductance values and the percentage of halophilic diatoms (Coles et al. 2004, Leland et al. 2001, Cuffney et al. 1997). Although diatom-species optima and tolerance values differ in relation to scales of assessment, both nationally (Potapova & Charles 2003) and regionally (e.g. Leland & Porter 2000, Leland et al. 2001, Munn et al. 2002), species optima for indicators of alkalinity and water hardness, and specific conductance, chloride, and sulfate were consistent with qualitative, autecological classifica-tions for those taxa in western Europe (e.g., van Dam et al. 1994, Lange-Bertalot 1979). Potapova & Charles (2003) suggested that weighted-
average optima and tolerance values derived for taxa, nationally, could be assigned to categories
according to their water-chemistry affinities to improve understanding of species autecology in the United States relative to existing European classification systems.
Indicator species and rare taxa Algal species associated with low-nutrient, near-
pristine stream conditions have been identified in a number of NAWQA reports ; however, the indi-
and Stephen D. Porter
cator species appear to vary regionally across the United States. For example, Hannaea arcus is found in abundance in high quality streams in the western United States (Cuffney et al. 1997, Brown & Short 1999, Cox-Lillis 2000, Mize & Deacon 2001) but is rarely reported east of the Mississippi River. Similarly, Cymbella delicatula has been cited as an indicator of excellent water
quality in limestone regions of the semi-arid southeastern and temperate midwestern United States (Bradfield & Porter 1990, Petersen & Femmer 2003) ; however, this taxon is rarely re-
ported from other regions. Certain nitrogen-fixing taxa (e.g., Nostoc spp., Calothrix parietina, Epi-themiaceae) and red algae (Audouinella herman-nii) often have been interpreted as oligotrophic, clean-water taxa (Cuffney et al. 1997, Brown & Short 1999, Scudder & Stewart 2001, Peterson & Porter 2002) but these taxa also can be abun-
dant in shaded, agricultural and urban streams of the midwestern and eastern United States
(Porter 2000, Kennen & Ayers 2002, Petersen & Femmer 2003). The water-quality-indicative prop-erties of other taxa such as Diatoma mesodon
(Carpenter & Waite 2000), Nitzschia frustulum var. perminuta (Cuffney et al. 1997), and Prasi-
ola or Pseudulvella (Brown 1996, S.D. Porter, un-
published data) appear to be basin-specific and (or) associated with pristine, high-elevation sites.
Potapova and Charles (2004) observed that are (but locally abundant) diatom species often
ire ignored as biological indicators, and they hy-
pothesize that rare or "native" American diatom species, particularly those associated with pristine
habitats, could be useful as indicators of stream
quality. They concluded that the number and abundance of rare species in the NAWQA da-
aset tended to be larger in streams that are less influenced by human activities ; however,
hey found regional distributions of rare, often indescribed taxa that may be endemic to the
southeastern coastal plain or western montane regions of the United States but that do not nec-
essarily indicate pristine water-quality conditions. Furthermore, the distribution of certain "native"
diatoms such as Gomphonema kobayasii (Ko-ciolek & Kingston 1999) appears to be wide-spread in eutrophic streams and rivers. There-ore, the presence of rare or "native" diatom
species is in itself not a reliable indicator of
water quality. These initial findings from selected sites
across the United States are included in the
planning of nationally guided regional and na-tional synthesis reports. The algal data and re-lated physical, chemical, and biological data that have been verified as accurate will be released and made available to the public through the USGS Data Warehouse which can be reached through the NAWQA website hap //water.usgs.
gov/nawqa/. In the second decade of the pro-
gram (2001-2011), biological-community data are to be obtained at selected sites for long-term trend analysis. Ecological studies also will be
part of nationally guided studies addressing se-lected water-quality issues, such as the effects of nutrient enrichment (Munn & Hamilton 2003) and urbanization (Couch & Hamilton 2002) on aquatic ecosystems.
Acknowledgments
The authors of this report thank the following U.S. Geological Survey reviewers for their contri-butions. Harry V. Leland and Charles R. Demas contributed thoughtful technical and editorial re-view comments. Daniel T. Button generated the figure. C. Michael Eberle reviewed the accuracy and suitability of the report, and provided valu-able editorial comments.
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Julie A. Hambrook Berkman : U.S. Geological Survey, 6480 Doubletree Ave., Columbus, OH 43229— 1111 U.S.A.
Stephen D. Porter : U.S. Geological Survey, WRD, P.O. Box 25046 MS 406, Lakewood, CO 80225— 0046 U.S.A.