The Pennsylvania State University
The Graduate School
Department of Ecosystem Science and Management
AN EXPERIMENTAL APPROACH TO EVALUATE EFFECTS OF NUTRIENTS
AND MEDIAN CONSUMERS ON STREAM TROPHIC DYNAMICS
A Thesis in
Wildlife and Fisheries Science
by
Rebecca Ann Eckert
 2012 Rebecca Ann Eckert
Submitted in Partial Fulfillment
of the Requirements
for the Degree of
Master of Science
August 2012
ii
The thesis of Rebecca Ann Eckert was reviewed and approved* by the following:
Hunter J. Carrick
Professor of Aquatic Ecosystems Ecology
Thesis Advisor
Jay R. Stauffer
Distinguished Professor of Ichthyology
Gregory A. Hoover
Senior Extension Associate
Ornamental Entomologist
Michael G. Messina
Professor of Forest Resources
Head, Department of Ecosystem Science and Management
*Signatures are on file in the Graduate School
iii
ABSTRACT
Studies of the simultaneous effects that consumers, nutrients, and other biotic/abiotic
factors have on intact food webs are rarely conducted in temperate streams. Such
knowledge may facilitate predictions about food-web structure regulation. Therefore, I
completed an in situ study using nutrient diffusing substrata with nitrogen (N) and
phosphorus (P) concentrations mimicking high productivity streams with a full factorial
design in three temperate, limestone streams in Pennsylvania across a trophic gradient
(mesotrophic, eutrophic, and hypereutrophic) during summer 2010 (low flow period) for
35 days. Biogeochemistry among the streams was similar, and all three supported “top
predator” fishes. I assessed differences in algal and macroinvertebrate biomass, density,
diversity, and taxonomic composition as influenced by amended nutrients across the
trophic gradient. All factors varied significantly among the streams (e.g., algal biomass
p=0.005, macroinvertebrate biomass p<0.001, algal diversity p=0.006, macroinvertebrate
diversity p<0.001, algal and macroinvertebrate guilds p<0.001) but did not follow simple
responses. Algal responses in the mesotrophic stream appeared to function as a three-step
chain (i.e., producer, primary consumer, secondary consumer) and showed the most
bottom-up influence while responses in the eutrophic and hypereutrophic streams
appeared to function as four-step chains; grazing seemed to be a strong factor preventing
nutrient response in the eutrophic and hypereutrophic streams. Brillouin’s Evenness
Index appeared most influenced by bottom-up factors (nutrient effect on algae and
macroinvertebrates p=0.021). Simple biomass and diversity responses were not seen; the
stream algal and macroinvertebrate responses seemed to most strongly depend on food
web configuration as mediated by median consumers.
iv
TABLE OF CONTENTS
List of Figures ........................................................................................................... vi
List of Tables ........................................................................................................... vii
Acknowledgements .................................................................................................viii
Background ................................................................................................................ 1
Introduction ................................................................................................................ 4
Methods...................................................................................................................... 7
Sites ................................................................................................................ 7
Experimental Design ...................................................................................... 8
Field Design ....................................................................................... 8
Laboratory Processing ..................................................................... 11
Data Analysis ............................................................................................... 13
Method Analysis .......................................................................................... 14
Results ...................................................................................................................... 16
Community Level Biomass and Density ..................................................... 17
Diversity Measures and Taxonomic Composition....................................... 22
Trophic Relationships .................................................................................. 43
Discussion ................................................................................................................ 45
Biomass Responses to Nutrient Enrichment ................................................ 45
Diversity Responses to Nutrient Enrichment ............................................... 48
Importance of Median Trophic Levels ........................................................ 53
Conclusions .................................................................................................. 56
v
References ................................................................................................................ 59
Appendix A: Algal taxa prescence/absence by stream ............................................ 69
Appendix B: Algal biomass by taxa in mesotrophic stream .................................... 71
Appendix C: Algal biomass by taxa in eutrophic stream ........................................ 74
Appendix D: Algal biomass by taxa in hypereutrophic stream ............................... 77
Appendix E: Algal density by taxa in mesotrophic stream...................................... 80
Appendix F: Algal density by taxa in eutrophic stream .......................................... 83
Appendix G: Algal density by taxa in hypereutrophic stream ................................. 86
Appendix H: Macroinvertebrate taxa prescence/absence by stream ....................... 89
Appendix I: Macroinvertebrate biomass by taxa in mesotrophic stream ................ 92
Appendix J: Macroinvertebrate biomass by taxa in eutrophic stream ..................... 97
Appendix K: Macroinvertebrate biomass by taxa in hypereutrophic stream ........ 102
Appendix L: Macroinvertebrate density by taxa in mesotrophic stream ............... 107
Appendix M: Macroinvertebrate density by taxa in eutrophic stream .................. 112
Appendix N: Macroinvertebrate density by taxa in hypereutrophic stream .......... 117
vi
LIST OF FIGURES
Figure 1. Representation of top-down effects in trophic cascades ............................ 2
Figure 2. Map of Mid-Atlantic region and three stream sites in Pennsylvania ......... 7
Figure 3. Individual NDS design and array of NDS bricks in stream ..................... 11
Figure 4. Algal and macroinvertebrate biomass distribution by stream/treatment .. 20
Figure 5. Lower trophic level biomass distribution by stream/treatment ................ 21
Figure 6. Algal group biomass distribution by stream/treatment ............................ 36
Figure 7. Macroinvertebrate guild biomass distribution by stream/treatment ......... 37
Figure 8. Algal group density distribution by stream/treatment .............................. 41
Figure 9. Macroinvertebrate guild density distribution by stream/treatment .......... 42
Figure 10. Biomass changes in predators, grazers, and algae by stream/treatment . 44
vii
LIST OF TABLES
Table 1. Biochemical characteristics of the three streams ........................................... 8
Table 2. Population of fishes in reach in each stream................................................ 10
Table 3. Physical stream conditions during experiment ............................................ 16
Table 4. Two-way ANOVA for algal, macroinvertebrate, and total biomass ........... 18
Table 5. Algal and macroinvertebrate biomass and density values ........................... 19
Table 6. Top ten algal taxa in mesotrophic stream .................................................... 23
Table 7. Top ten algal taxa in eutrophic stream ......................................................... 24
Table 8. Top ten algal taxa in hypereutrophic stream ................................................ 25
Table 9. Top ten macroinvertebrate taxa in mesotrophic stream ............................... 26
Table 10. Top ten macroinvertebrate taxa in eutrophic stream ................................. 27
Table 11. Top ten macroinvertebrate taxa in hypereutrophic stream ........................ 28
Table 12. Algal and macroinvertebrate diversity measures (Brillouin’s Indices) ..... 31
Table 13. Algal biomass by group ............................................................................. 33
Table 14. Macroinvertebrate biomass by group ........................................................ 34
Table 15. Two-way ANOVA for algal group biomass and density............................ 35
Table 16. Two-way ANOVA for macroinvertebrate guild biomass and density ....... 35
Table 17. Algal density by group ............................................................................... 39
Table 18. Macroinvertebrate density by group .......................................................... 40
viii
ACKNOWLEDGEMENTS
Thank you to my advisor, Hunter Carrick, without whom none of this would have been
possible as he gave me the opportunity to conduct my own project and taught me so
much about the aquatic stream world and its inhabitants. Thank you to my committee,
Greg Hoover and Jay Stauffer, who provided comments and feedback regarding my
project and taught me something new regarding their own aquatic specialties. Thank you
additionally to the Pennsylvania Department of Environmental Protection and Penn
State’s School of Forest Resources for funding.
Thank you also to my labmates, Keith Price, Melissa May, and Erin Cafferty, with whom
so much memorable time was spent, including lots of trips “climbin’ in yo’ streams,
snatchin’ yo’ substrates up, tryin’ to scrape ‘em so ya’ll need to hide ya rocks, hide ya
tiles, and hide ya razors cuz we scrapin’ everything in here”.
Finally, thank you to my family for their support throughout my tenure as a graduate
student, in letting me follow my dreams and never holding me back.
Without any one of you, none of this would have been possible, and for that, I will
always be grateful.
1
Background
The debate as to the relative influence of consumer-mediated top-down effects as
opposed to resource-mediated bottom-up effects on food webs is longstanding in ecology
(Power 1992). Hairston et al. (1960) first proposed the idea of a “green world”, wherein
top-down influences are most important in controlling primary production. The total
number of trophic levels affects primary producers given that successive trophic levels
are thought to become alternately limited by resources or consumers (Fretwell 1987). For
instance, in a three-step chain (producer, primary consumer, secondary consumer),
secondary consumers may depress the biomass of primary consumers, allowing
producers to increase in biomass (Figure 1a; “green world”). Alternatively, in a four-step
chain, secondary consumers may be depressed by tertiary consumers, allowing primary
consumers to increase, and producers may then be depressed (Figure 1b). Bottom-up
processes operate under the assumption that there is a limiting resource regulating
production (e.g., White 1978). These resources can include nutrients (e.g., Dodds et al.
2002), space (e.g., Fonseca 1999), food/energy (e.g., Lindeman 1942), and also
environmental abiotic factors such as light (e.g., Hill et al. 2009) and temperature (e.g.,
Morin et al. 1999). Top-down and bottom-up processes were first described in detail by
Paine (1966) for intertidal food webs, wherein consumer effects were primarily
examined, but space was also addressed as a limiting factor of production.
These perspectives do not, however, take into consideration the many factors that
can complicate a straight food chain which may in reality function as a web. For instance
omnivory, defined as an organism feeding from more than one trophic level (Pimm and
Lawton 1978), is thought to be common in natural food webs (Diehl 1993; Křivan and
2
Diehl 2005; Thompson et al. 2007). Omnivory, when present, can confound relationships
among trophic levels by altering the distribution of energy thus creating a food web that
differs from a linear response (Diehl 1993; Thompson et al. 2007). The occurrence of
omnivory can complicate any assumptions made regarding interactions, especially since
theoretical and observational/experimental evidence is often contradictory
(HilleRisLambers et al. 2006). Additionally, other factors such as interspecific
competition can change the functioning of a food web (Katano et al. 2003). Even so,
aquatic systems provide habitats where top-down and bottom-up interactions can often be
clearly observed and experimentally tested (Gruner et al. 2008; Shurin et al. 2006).
A
2° Consumer
B
3° Consumer
2° Consumer
1° Consumer
1° Consumer
Producer
Producer
Figure 1. Representation of changes in biomass at alternating levels in trophic cascades
for (A) a three-level food chain and (B) a four-level food chain. Arrows to the left of the
trophic level indicate increases or decreases in biomass, as interpreted from top consumer
down.
Historically, the simultaneous effects of consumer and resource influences in
aquatic systems have been studied in lakes (e.g. Brett and Goldman 1997; Carpenter et al.
1985; Carpenter et al. 1987; McQueen et al. 1989) with less attention on streams (but see
Biggs et al. 2000; Deegan et al. 1997; Nyström et al. 2003; Rosemond et al. 1993). Much
3
of the stream research to date has focused on the influences of nutrients and
macroinvertebrate grazing on periphyton rather than entire food webs (Hillebrand 2002;
Rosemond et al. 1993; Sturt et al. 2011), on detrital based webs rather than algal based
webs (McIntosh et al. 2005; Rosemond et al. 2001), and some on the influence of fish
alone on lower trophic levels (Cheever and Simon 2009). Interestingly, all these studies
dealt with specific subsets of food webs, and therefore may have excluded important
feedbacks that can only be evaluated with ecosystem level research on relatively intact
communities (Peterson et al. 1993). Furthermore, the streams where ecosystem scale
studies have been done tend to be located outside the temperate zone, especially in the
Arctic (Deegan et al. 1997; Perrin and Richardson 1997; Peterson et al. 1993) and New
Zealand (Huryn 1998; McIntosh et al. 2005; Nyström et al. 2003). These results may not
directly translate to temperate streams (Deegan et al. 1997; Winterbourn et al. 1981),
presenting a gap in the knowledge of stream food webs as functioning natural
communities.
4
Introduction
Nutrient enrichment, particularly nitrogen (N) and phosphorus (P), has typically
promoted increased primary productivity in streams, and these effects have been
evaluated using both empirical (Biggs 2000; Dodds et al. 2002) and experimental
techniques (Fairchild et al. 1985; Rosemond et al. 1993). The relationship between
nutrients and benthic algal biomass in streams is, however, more complicated than the
relationship seen in lakes (Miltner and Rankin 1998) and may not be as strong as the
relationship between nutrients and phytoplankton (Francoeur 2001). Regardless,
excessive nutrient enrichment still leads to eutrophication (Smith et al. 1999) which can
have particularly detrimental effects on higher trophic levels through, for example,
additions of toxins and decreases in nocturnal dissolved oxygen availability (Miltner and
Rankin 1998; Wang et al. 2007). Increased biomass of primary producers has been shown
to propagate through the stream food web (Fretwell 1987; Wootton and Power 1993),
affecting both macroinvertebrates and fishes. For instance, nutrient additions have
increased macroinvertebrate biomass and density (Perrin and Richardson 1997; Rader
and Richardson 1992; Wang et al. 2007; Winterbourn 1990), have increased fish growth
(Peterson et al. 1993), and have been correlated to fish assemblage measures such as
salmonid catch and the fish index of biotic integrity (Wang et al. 2007).
While biomass exhibits a monotonic response along productivity gradients (such
as seen with nutrient enrichment in streams, e.g., Dodds et al. 2002), diversity has often
been found to show a unimodal response along productivity gradients (Liebold 1999;
Tilman 1982), although this idea has been subject to debate (e.g., Abrams 1995) and may
be dependent upon scale (Waide et al. 1999). Higher diversity is therefore often observed
5
with modest nutrient enrichment, and diversity decreases at high levels of nutrient
enrichment. This trend has been observed in benthic (Carrick et al. 1988) and pelagic
algal communities (Liebold 1999), macroinvertebrates and fish (Wang et al. 2007),
bacteria (Kassen et al. 2000), terrestrial plant communities (Hautier et al. 2009), and
aquatic systems in general (Paine 1966). It is likely that the response of diversity
measures along a productivity gradient is mediated by other ecosystem factors (e.g.,
temperature and oxygen availability for aquatic systems), and the strength of a given
factor’s influence depends upon the system being examined, e.g., lake versus wetland
versus desert (Waide et al. 1999).
These direct nutrient enrichment effects on biomass and diversity may be
mediated by community complexity and food chain configuration. Consumers regulate
lower trophic levels through both direct feeding activities and nutrient cycling (Huryn
1998; Rosemond et al. 1993; Small et al. 2009). For example, in aquatic ecosystems
trophic cascades have been identified as strong effects influencing primary productivity
from the top of the food chain (Shurin et al. 2006), and primary producer biomass
depends upon the number of functional trophic levels (Fretwell 1987). The strength of
consumer effects in streams can, however, be limited by multiple factors mediating these
effects from cascading to the primary producers. These can include abundance of drifting
prey available to fishes (Dahl 1998; Nakano et al. 1999), size selective predation
(Meissner and Muotka 2006), type of predator (Biggs et al. 2000), omnivory (Bascompte
et al. 2005; Diehl 1993; Katano et al. 2003; Power 1992; Thompson et al. 2007),
intraguild/interspecific interactions (Katano et al. 2006; Lancaster et al. 2005; Ruetz et al.
2003; Tanabe and Namba 2005; Wissinger and McGrady 1993), and temperature (Kishi
6
et al. 2005). Trophic dynamics are therefore influenced by producers (Fretwell 1987;
Power 1992), by consumers (Paine 1966; Peterson et al. 1993), or a combination of the
two.
Although more than 50% of streams in the coterminous United States are located
in the eastern temperate forest ecoregion (Benke 1990), there is a lack of studies
considering consumer and nutrient effects simultaneously on intact stream food web
structure in temperate North America. This represents a major gap in knowledge that
should be addressed. Herein, I examined how benthic algae and macroinvertebrates
change with respect to nutrient enrichment along a stream trophic gradient. Specifically, I
examined benthic algae and macroinvertebrate biomass and diversity responses to
nutrient enrichment across this gradient to determine if the responses were simple and
linear or due to functional food web complexity (via trophic level analysis). This research
will contribute to the body of knowledge in temperate streams regarding relative
regulation in natural food webs by resources (nutrients) and consumers.
7
Methods
Sites
This experiment was completed in three limestone streams within the MidAtlantic region of the United States (Cooks Creek [mesotrophic], Penns Creek
[eutrophic], and Spring Creek [hypereutrophic]; Figure 2) representing a trophic gradient
with trophic state based on Dodds et al. (1998) benthic chlorophyll values; a gradient of
nutrient load, conductivity, and fish population also existed (Table 1). The three streams
are part of the Pennsylvania Department of Environmental Protection Water Quality
Network (WQN—see Table 1 for numbers) in Pennsylvania which monitors temporal
water quality trends. Experimental sites were chosen based on similar canopy cover,
stream width, and stream depth as well as security from vandalism. The eutrophic and
mesotrophic sites were located on private property (located at 40°51’32.4” N,
77°34’46.6” W and 40°35’7.4” N, 75°12’22.0” W respectively) while the hypereutrophic
site was on Pennsylvania State University property (located at 40°49’19.9” N,
77°50’12.8” W).
Map adapted from “Pennsylvania Counties Map”, AnonMoos, 25 April 2010, and “US Mid-Atlantic
states”, Grayshi and Roke, 13 March 2010. Both were uploaded to Wikimedia Commons under the
Creative Commons Attribution-ShareAlike license (CC BY-SA 3.0).
Figure 2. Location of Pennsylvania in the Mid-Atlantic region and three stream sites in
Pennsylvania.
8
Table 1. Biochemical characteristics of the three streams, where TN and TP are total
nitrogen and phosphorus, respectively, condo is conductivity, and chl-a is the
chlorophyll-a areal concentration from natural rocks.
Stream
Mesotrophic
(WQN 187)
Eutrophic
(WQN 229)
Hypereutrophic
(WQN 415)
TN*
(mg/L)
TP*
(mg/L)
Condo**
(μS)
pH**
(pH units)
Total
Alkalinity**
(mgCaCO3/m2)
Chl-a**
(mg/m2)
Fish
Population***
(number/m)
1.770
0.021
222.29
8.1
92.5
68.95
520.55
0.895
0.061
305.65
8.0
141.3
263.24
580.37
4.760
0.035
327.63
8.3
181.0
442.42
217.89
*Data from Carrick et al. 2009
**Data are means from year of sampling (2009-2010); pH, Condo, Chl-a n=8; Total Alkalinity n=4
***Data from preliminary three-pass depletion electrofishing of sites Leslie calculations (Aug 2010)
Experimental Design
Field Design- In-situ nutrient enrichment experiments were completed using methods
described by Fairchild et al. (1985) and subsequently by Godwin et al. (2009). Terracotta
clay flower pot saucers (15.24 cm diameter, 2.54 cm height, 1.20 cm depth) were used as
nutrient-diffusing substrata (NDS) for colonization by macroinvertebrates and
periphyton. The substrata were soaked for three days in deionized water and the open
side sealed with Plexiglas (0.159 cm thick, 17.78 cm by 17.78 cm squares) using silicone
to create a chamber with a diffusing surface area to volume ratio of approximately 1.39,
similar to Fairchild et al. (1985)’s surface area to volume. Agar containing the various
nutrient treatments was injected into a hole drilled in the Plexiglas. The hole was sealed
with silicone and electrical tape after the agar cooled. The Plexiglas additionally had two
Plexiglas strips (17.78 cm by 2.54 cm; 0.635 cm thick) attached with silicone in order to
create an interstitial space beneath the NDS that was adequate for macroinvertebrate
colonization (Fairchild and Holomuzki 2005). The NDS were attached to cement bricks
9
with silicone, with the Plexiglas strips oriented upstream and downstream to prevent
channelization of the flow (Figure 3a). Sixteen NDS were placed in each stream: four
controls, four 0.5 M nitrogen (N), four 0.05 M phosphorus (P), and four 0.5 M N + 0.05
M P saucers. Sodium nitrate (NaNO3) was used to provide N as nitrate, and sodium
phosphate monobasic (NaHPO4) was used to provide P as phosphate. I estimated the N
leaching rate as approximately 14 g/m2/day and the P leaching rate as about 1 g/m2/day
based upon data collected by Fairchild et al. (1985). This provided a molar N:P ratio of
about 15:1, close to the optimal (Redfield) ratio required for algal growth.
Prior to deployment, each reach was electrofished using a three-pass depletion to
determine the natural fish populations. Fish were identified in the field to species by R.
Taylor (M.S. candidate under J. Stauffer, ichthyologist at The Pennsylvania State
University); the exception was sculpin identified to genus (Cottus) with a few voucher
specimens returned to the Stauffer laboratory at The Pennsylvania State University.
Specific fish populations (and total fish population for the site) were calculated using the
Leslie method (Table 2) for length of reach electrofished. Upon deployment, the sixteen
NDS for each stream were randomly assigned a number of one through sixteen and
placed in the stream in an array as shown in Figure 3b. Five blank bricks were placed in
front of the NDS to account for any differential flow effects due to brick location in the
array. The array of bricks was placed in a run area of approximately the same depth in
every stream. Densiometer measurements were taken over the center of each array at
deployment.
10
Table 2. Population of fishes within electrofished reach (number per 25m), calculated
using Leslie method. Fish with * indicates population estimate could not be made and
value represents total number of fish caught in electrofished reach.
Common Name
Scientific Name
Mesotrophic
Eutrophic
Hypereutrophic
American eel
Anguilla rostrata
50.99
0.00
0.00
White sucker
Catostomus commersonii
16.03
1.67
3.00
Blacknose dace
Rhinichthys atratulus
307.58
43.42
30.81
*Bluntnose minnow Pimephales notatus
5.00
7.00
0.00
Central stoneroller
Campostoma anomalum
23.82
0.00
0.00
Cutlips minnow
Exoglossum maxillingua
1.00
0.00
3.00
*Longnose dace
Rhinichthys cataractae
4.00
0.00
0.00
Brown trout
Salmo trutta
1.00
21.69
35.27
*Rainbow trout
Oncorhynchus mykiss
0.00
2.00
0.00
Sculpin
Cottus spp.
0.00
559.47
188.15
*Bluegill
Lepomis macrochirus
0.00
1.00
0.00
*Pumpkinseed
Lepomis gibbosus
0.00
1.00
0.00
Smallmouth bass
Micropterus dolomieu
6.75
0.00
0.00
*Shield darter
Percina peltata
2.00
0.00
0.00
Tessellated darter
Etheostoma olmstedi
191.09
0.00
0.00
520.55
580.37
217.89
Total population
The NDS experiment was incubated in the streams for five weeks from midAugust to mid-September 2010 when flow was at its lowest to enable easier sampling
conditions and to avoid any scour events that could alter the results. Algal biofilm
colonization was expected to occur quickly, as typically new communities are similar to
communities on natural substrates in about four weeks in temperate streams (Biggs
1988). Insect colonization after spates occurs quickly (Matthaei et al. 1996; Miller &
Golladay 1996); with no scouring event prior to deployment, colonization was expected
to occur more quickly than post-spate rates. After five weeks, the NDS were collected
beginning at the downstream end of the array (#16) by lifting the bricks carefully out of
11
the water with a D-frame net placed downstream to catch any escaping
macroinvertebrates. The biofilms (with both macroinvertebrates and algae) present on the
NDS were removed with a razor and soft-bristle brush and washed into containers for
transport to the laboratory. Any macroinvertebrates caught in the net were added to the
container for the respective NDS. The entire surface of the pot was sampled in addition to
the exposed Plexiglas on the top and the Plexiglas on the bottom not directly attached to
the brick; a total surface area of 445.45 cm2 was sampled.
A
Plexiglas
sheet
Interstitial
space
Plexiglas
strip
Clay pot
Flow
Brick
B
Flow
1
5
2
6
9
13
7
10
14
4
3
8
11
15
12
16
Figure 3. (A) Side view of an individual nutrient diffusing substrata as attached to
brick. (B) Field array of nutrient diffusing substrata as deployed in stream.
Laboratory Processing- Biofilms were returned to the laboratory and sieved with a 500
μm mesh screen over a plastic container to separate macroinvertebrates from the biofilm
mat (see below). The slurry separated from macroinvertebrates was placed in a beaker
12
and diluted to a known volume. A subsample of two milliliters was taken from the slurry
and vacuum filtered onto membranes (Whatman EPM 2000 filters). The filters were then
frozen until fluorometric analysis of chlorophyll-a as a proxy of biomass. Chlorophyll-a
was extracted with a 50:50 mixture of dimethylsulfoxide and 90% acetone and the
fluorescence measured on a Turner 10-AU Fluorometer (Turner Designs, Sunnyvale,
California, USA) before and after acidification to correct for phaeopigments (Carrick et
al. 1993). Sixty milliliters of the unfiltered slurry was then preserved in 1% formalin.
To identify soft algae, four drops of preserved algal slurry was placed in a Palmer
Maloney Counting Chamber and covered with a glass coverslip; cells were identified to
genus under 100x and 400x magnification (Carrick and Steinman 2001). The slide was
first scanned under 100x magnification to determine “large” countable cells. All of these
large cells were then identified and counted to a minimum of 100 cells at 100x (utilizing
either half or the entirety of the Palmer cell). The rest of the cell types were identified and
counted under 400x magnification in random fields to a minimum of 200 cells and 10
fields, until 400 cells total were reached under both 100x and 400x.
Macroinvertebrates captured on the sieve from each NDS were subsequently
washed into a container with 70% ethanol for preservation. Macroinvertebrates were
sorted to lowest possible taxonomic grouping, typically genus, according to Merritt et al.
(2008) and Peckarsky et al. (1990); Chironomidae were sorted to subfamily and tribe
according to head capsule shape following Simpson and Bode (1980). Approximately
10% of each chironomid subfamily or tribe from the most numerous Chironomidae
sample from each stream was mounted following standard procedures and identified to
genus for verification (Merritt et al. 2008; Simpson and Bode 1980). The
13
macroinvertebrates were sorted into guilds according to Merritt et al. (2008). Aquatic
worms were left at Oligochaeta and aquatic mites at Hydrachnidia. The
macroinvertebrate biomasses were determined using published length-dry mass
relationships (Baumgärtner and Rothhaupt 2003; Benke et al. 1999; Burky 1971; Butzler
2002; Dumont et al. 1975; Eckblad 1971; McMahon 1975; Meyer 1989; Ramsay et al.
1997; Sabo et al. 2002; Stoffels et al. 2003), based on an average length from the first
five organisms encountered of each taxon.
Data Analysis
Algal dry weight was calculated as 20% of cell biovolume based on taxonomic
assessment (O’Reilly and Dow 2006). Brillouin’s Diversity, Evenness, and Maximum
Diversity (Hmax) were calculated for each trophic level (Moore 1987). Higher
Brillouin’s Diversity values indicate greater diversity. Evenness values between 0.5 and 1
indicate even communities where calculated diversity is closer to the maximum.
Maximum diversity indicates taxa are distributed as evenly as possible. Population
growth rates and doubling times were calculated using standard equations and assuming
initial values of one cell/individual for population calculations and 0.01 mg for biomass
calculations. All data were log10 transformed or log10(x+1) transformed to meet
normality. A two-way analysis of variance (ANOVA) was conducted on biomass and
density for algae (chlorophyll-a) and macroinvertebrates to test for differences by stream
(trophic gradient) and nutrient treatment. The total biomass (mg dry weight) in each
stream was calculated and a two-way ANOVA conducted to test if variation existed
between specific trophic levels and/or between stream total biomass, and Tukey post-hoc
tests were conducted as appropriate.
14
In order to determine whether differences existed between functional groups, a
two-way multivariate analysis of variance (MANOVA) was conducted to test group
differences for algae and macroinvertebrates between stream and nutrient treatments,
with ANOVAs conducted as appropriate. Due to numerical deficiencies within groups,
algae of Euglenophyta, Rhodophyta, and Chrysophyta were categorized as “other” and
macroinvertebrates classified as parasites, scavengers, shredders, or non-feeding (i.e.,
pupae and adults) were also categorized as “other”, while piercers were grouped with
predators. Diversity differences were assessed by ANOVAs conducted on various
diversity measures. For all ANOVAs, Tukey’s post-hoc tests were conducted as
appropriate and an alpha value of 0.05 was used for the significance level, but results
with p-values between 0.05 and 0.1 may be considered to have biological significance
and may therefore be discussed. Pearson correlations were conducted to test for
relationships between algal and macroinvertebrate measures to infer connections between
trophic levels.
Method Analysis
A preliminary test of the NDS system was performed from May to June 2010 for
four weeks in the hypereutrophic stream. Duplicate nutrient treatments were placed in the
stream; two pots were randomly assigned to one brick and lacked Plexiglas strips to raise
them as in the final design (n=8 pots). The NDS were collected and treated in the same
manner as described above. Upon collection, the first brick (control and N+P treatment)
appeared scoured in comparison to the other three bricks which may have influenced the
resulting analysis. Macroinvertebrate colonization between NDS and natural substrates
15
(rocks collected in May) were examined for differences. A paired t-test between family
densities (normalized to area sampled) on rocks and control NDS revealed a significant
difference (n=25; p=0.041). During sampling, Trichoptera (caddisflies) were mostly
found clinging to the bottom of rocks (personal observation), so the analysis was repeated
with trichopterans removed and there was no significant difference (n=18; p=0.092). This
suggested the NDS design may have lacked suitable habitat for these organisms
(Fairchild and Holomuzki 2005). The final design was then modified by adding two
Plexiglas strips (as described above; Figure 3A) to raise the pot off the surface of the
brick to accommodate Trichoptera.
16
Results
Physical conditions
The physiochemical parameters measured during the experiment showed little
variation (Table 3). For example, stream discharge values were well below historical
averages for August and September (71% of historical average for mesotrophic, 58% for
eutrophic, and 53% for hypereutrophic). While an episodic rain event on August 22, 2010
caused increased discharge in the eutrophic and hypereutrophic streams, the streams
quickly returned to base flow conditions as seen in the median flow value of 2.46 m3/s for
the eutrophic stream and 0.59 m3/s for the hypereutrophic stream. Conductivity was high
in all three streams, while chlorophyll-a measured on rocks showed an increase from
collection at time of deployment to collection at retrieval. In all streams pH was greater
than seven and total alkalinity values were greater than 100 mg CaCO3/L, indicative of
the limestone geology underlying the streams. Water temperature decreased over the
course of the study by about 2°C in all streams, with warmest temperatures seen in the
mesotrophic stream and coldest in the hypereutrophic stream.
Table 3. Physical stream conditions measured during the experiment.
Mesotrophic1
Eutrophic2
Hypereutrophic3
1
Areal
pH
Alkalinity
Temp Condo
Chl-a
(pH
(mg
(°C) (μs/cm)
2
(mg/m )
units) CaCO3/L)
36.27 17.80 290.60 8.19
110
128.76 13.80 378.50 7.96
170
202.89 11.90 404.35 7.54
240
Mean
Std
Min
Max
Flow
Dev
Flow
Flow
(m3/s) (m3/s) (m3/s) (m3/s)
1.28
0.12
1.02
1.56
2.94
1.44
1.78
8.10
0.65
0.21
0.51
1.78
USGS 01446776 Bushkill Creek at Tatamy, PA
USGS 01555000 Penns Creek at Penns Creek, PA
3
USGS 01546400 Spring Creek at Houserville, PA
Areal chlorophyll-a (Chl-a) was determined using biofilms scraped from natural rocks collected at the time
of deployment and retrieval. Temperature (temp) and conductivity (condo) were determined in stream with
a handheld YSI-85 meter, while pH was determined upon arrival at the lab with a benchtop VWR pH
meter. Total alkalinity was calculated using color titration method. Stream discharge was retrieved from
USGS’s waterwatch.usgs.gov website for sites as close as possible to the experimental site.
2
17
Community Level Biomass and Density
Algal biomass (as chlorophyll-a) showed a significant difference between streams
(two-way ANOVA, p=0.005; Table 4). Tukey post-hoc testing indicated significantly
higher algal biomass in the mesotrophic compared to the hypereutrophic stream
(p=0.005) and compared to the eutrophic stream (p=0.060). Algal dry weight was
significantly correlated to chlorophyll-a (r=0.510, p<0.001, n=47) and also showed
significant differences by stream (two-way ANOVA, p<0.001; Table 4); post-hoc testing
showed the mesotrophic stream had significantly more biomass than both streams
(p<0.001 for both comparisons).
Macroinvertebrate biomass likewise varied by stream (two-way ANOVA,
p<0.001; Table 4), with Tukey post-hoc tests indicating the mesotrophic stream had
significantly less biomass than both the eutrophic (p=0.002) and the hypereutrophic
stream (p<0.001). Total dry biomass (algal + macroinvertebrate; Table 4) showed the
same pattern as algal biomass (two-way ANOVA, stream p<0.001), with the mesotrophic
stream having significantly higher biomass than the eutrophic (Tukey post-hoc test,
p=0.002) and hypereutrophic stream (p<0.001). Similarly, two-way ANOVAs on algal
and macroinvertebrate density showed significant differences between streams
(F2,35=28.982, p<0.001; F2,35=35.505, p<0.001, respectively), no algal nutrient treatment
differences (F3,35=1.936, p=0.142), nor macroinvertebrate nutrient treatment differences
(F3,35=2.342, p=0.090). Tukey post-hoc tests indicated algal density was significantly
higher in the mesotrophic stream than the other two (p<0.001 for both comparisons). In
contrast, the mesotrophic stream had significantly lower macroinvertebrate density than
18
the other two streams (p<0.001 for both). Non-significant slight differences (p=0.078)
were seen between N (lowest density) and P (highest density) in Tukey post-hoc tests.
Overall patterns revealed that dual nutrient enrichment (N+P) in all streams
produced the highest algal biomass (chlorophyll-a; Table 5). The most pronounced
differences among treatments were seen in the mesotrophic stream (Figure 4a).
Macroinvertebrate biomass, however, was highest on the P treatments and lowest on the
N treatments except in the mesotrophic stream. Instead, in the mesotrophic stream N
treatments were greater than the P treatments, but highest biomass was on the N+P
treatments similar to the chlorophyll-a pattern (Table 5, Figure 4c). Dry algal biomass
reflected a pattern intermediate between the chlorophyll-a biomass and the
macroinvertebrate biomass results (Figure 4b). The majority of biomass in each stream
was comprised of algae, with approximately 30% of lower trophic level biomass in the
macroinvertebrates (Figure 5); by stream, a greater amount of biomass was found in the
macroinvertebrates in the eutrophic and hypereutrophic stream (34.0% and 40.3%,
respectively) than in the mesotrophic stream (12.3%).
Table 4. Two-way ANOVA results for algal biomass (mg Chl-a/m2), algal dry weight
(mg DW/m2), macroinvertebrate biomass (mg DW/m2), and total biomass (mg DW/m2)
showing F values with p-values in parentheses.
Factor
Stream
Nutrient Treatment
Stream *Nutrient
Treatment
Algal Biomass
(Chl-a)
6.183 (0.005)
1.255 (0.305)
0.437 (0.849)
Algal Biomass
(Dry weight)
37.241 (<0.001)
2.091 (0.119)
1.839 (0.120)
Macroinvertebrate
Total Biomass
Biomass
(Dry Weight)
(Dry weight)
11.054 (<0.001)
14.635 (<0.001)
0.423 (0.738)
0.798 (0.503)
1.110 (0.377)
0.729 (0.630)
19
Table 5. Algal and macroinvertebrate (macro) biomass and density among streams and
nutrient treatments. Numbers represent means ±1 standard deviation; N=nitrogen
treatments, P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Biomass
(mg Chl-a/m2)
Algal Density
(cells/cm2)
Macro Biomass
(mg DW/m2)
Macro Density
(number/m2)
Total Biomass
(mg DW/m2)
43.58
(6.99)
2.68x106
(1.22x106)
204.3
(83.9)
1470
(437)
1611.0
(394.4)
N
71.80
(45.98)
3.33x106
(1.19x106)
229.8
(128.2)
1072
(368)
1715.6
(392.9)
P
70.04
(16.32)
2.60x106
(3.17x105)
166.3
(31.2)
1325
(286)
1698.2
(237)
N+P
81.56
(15.03)
3.01x106
(1.01x106)
270.1
(209.4)
1527
(609)
1856.0
(641.6)
Stream Average
66.74
(27.25)
2.91x106
(9.43x105)
217.6
(123.2)
1348
(434)
1720.2
(404.7)
46.98
(22.37)
1.24x106
(7.62x105)
338.7
(221.6)
2357
(988)
1257.2
(58.0)
N
44.08
(15.64)
1.23x106
(2.56x105)
313.7
(80.7)
2554
(831)
1101.2
(244.2)
P
39.53
(16.22)
7.14x105
(2.11x105)
543.1
(222.6)
4737
(1180)
1019.2
(275.9)
N+P
53.97
(34.37)
1.47x106
(5.28x105)
416.3
(158.5)
2688
(945)
1480.9
(211.2)
Stream Average
46.14
(21.62)
1.16x106
(5.25x105)
403.0
(185.9)
3084
(1333)
1214.7
(264.1)
35.81
(17.74)
9.44x105
(3.05x105)
514.6
(147.8)
3216
(621)
1088.9
(271.1)
N
39.30
(9.00)
1.11x106
(1.73x105)
348.2
(135.9)
2711
(1196)
934.3
(123.3)
P
30.27
(10.27)
1.11x106
(7.17x105)
516.9
(213.5)
3405
(1274)
1160.2
(406.5)
N+P
47.88
(27.63)
1.25x106
(5.07x105)
357.8
(156.5)
2514
(940)
1113.9
(522.9)
Stream Average
38.85
(17.47)
1.10x106
(4.10x105)
428.9
(165.7)
2932
(972)
1068.6
(329.9)
Mesotrophic
Control
Eutrophic
Control
Hypereutrophic
Control
20
A
140
Chlorophyll-a (mg/m2)
120
100
Control
80
N
60
P
40
N+P
20
0
Mesotrophic
Algal Biomass (mg DW/m2)
B
Eutrophic
Hypereutrophic
2500
2000
Control
1500
N
1000
P
N+P
500
0
Mesotrophic
Hypereutrophic
900
800
700
600
500
400
300
200
100
0
Macroinvertebrate Biomass
(mg DW/m2)
C
Eutrophic
Control
N
P
N+P
Mesotrophic
Eutrophic
Hypereutrophic
Figure 4. Distribution of biomass between streams and nutrient treatments. Values
shown are means with error bars representing ±1 standard deviation; N=nitrogen
treatments, P=phosphorus treatments, N+P=dual nutrient treatments. A) Algal biomass
distribution using chlorophyll-a as a proxy B) Algal biomass distribution using dry
weight C) Macroinvertebrate biomass distribution using dry weight.
21
A
3000
Dry Weight (mg/m2)
2500
2000
1500
1000
500
0
B
Control
N
P
N+P
Control
N
P
N+P
Control
N
P
N+P
3000
Dry Weight (mg/m2)
2500
2000
1500
1000
500
0
C
3000
Dry Weight (mg/m2)
2500
2000
1500
1000
500
0
Figure 5. Total lower trophic level biomass distribution for each stream by treatment.
N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments, algal
dry biomass is light gray, and macroinvertebrate dry biomass is dark gray. A)
Mesotrophic stream B) Eutrophic stream C) Hypereutrophic stream.
22
Diversity Measures and Taxonomic Composition
A total of 51 algal taxa was identified among all three streams, comprised of 19
Bacillariophyta, 17 Chlorophyta, 12 Cyanobacteria, and 3 “other” (Euglena, Synura,
Bactrachospermum; full list in Appendix A). The most taxa were found in the
mesotrophic stream, with 44 taxa total, comprised of 19 Bacillariphyta, 12 Chlorophyta,
11 Cyanobacteria, and 2 “other” (Table 7). In the eutrophic stream, 41 taxa were found,
with 18 in Bacillariophyta, 14 Chlorophyta, 7 Cyanobacteria, and 2 “other” (Table 8). A
total of 38 taxa was found in the hypereutrophic stream, with 19 Bacillariophyta, 11
Chlorophyta, 6 Cyanobacteria, and 2 “other” (Table 9). A total of 62 macroinvertebrate
taxa was identified among the three streams, with 15 collector-gatherers, 12 collectorfilterers, 15 scrapers, 10 predators, and 10 “other” (full list in Appendix H). Of these, 34
were found in the mesotrophic stream, with 10 collector-gatherers, 7 collector-filterers, 8
scrapers, 6 predators, and 3 “other” (Table 9). In the eutrophic stream, 33 taxa were
found, comprised of 12 collector-gatherers, 5 collector-filterers, 4 scrapers, 5 predators,
and 7 “other” (Table 10). A total of 41 taxa was encountered in the hypereutrophic
stream, with 13 collector-gatherers, 9 collector-filterers, 8 scrapers, 4 predators, and 7
“other” (Table 11).
23
Table 6. Top ten algal taxa in mesotrophic stream by biomass and density. Percent
represents total from average of samples. For entire taxa lists, see Appendices B and E.
Division
Genus
Percent of Total Sample
Biomass
Bacillariophyta
Naviculoid (large)
22.44
Bacillariophyta
Cocconeis
21.31
Bacillariophyta
Nitzschia
13.61
Bacillariophyta
Achnanthes-like
12.94
Bacillariophyta
Gomphonema
9.38
Bacillariophyta
Synedra
4.09
Bacillariophyta
Melosira
1.98
Bacillariophyta
Rhoicosphenia
1.94
Bacillariophyta
Diatoma
1.80
Bacillariophyta
Naviculoid (small)
1.63
Total
91.14
Density
Bacillariophyta
Achnanthes-like
23.85
Cyanobacteria
Blue-green balls
23.58
Cyanobacteria
Lyngbya sp1
8.05
Cyanobacteria
Lyngbya sp2 (thin)
7.89
Bacillariophyta
Naviculoid (large)
6.87
Bacillariophyta
Gomphonema
4.96
Bacillariophyta
Naviculoid (small)
4.64
Bacillariophyta
Nitzschia
4.62
Bacillariophyta
Cocconeis
3.87
Cyanobacteria
Microcystis sp
3.60
Total
91.93
24
Table 7. Top ten algal taxa in eutrophic stream by biomass and density. Percent
represents total from average of samples. For entire taxa lists, see Appendices C and F.
Division
Genus
Percent of Total Sample
Biomass
Bacillariophyta
Naviculoid (large)
22.31
Bacillariophyta
Nitzschia
18.68
Bacillariophyta
Cocconeis
16.13
Bacillariophyta
Cymbelloid
13.23
Bacillariophyta
Achnanthes-like
9.03
Bacillariophyta
Surrirella
6.24
Bacillariophyta
Diatoma
3.58
Bacillariophyta
Gomphonema
2.47
Bacillariophyta
Naviculoid (small)
1.62
Cyanobacteria
Blue-green balls
1.41
Total
94.70
Density
Cyanobacteria
Blue-green balls
35.12
Bacillariophyta
Achnanthes-like
22.47
Bacillariophyta
Naviculoid (large)
9.22
Bacillariophyta
Nitzschia
8.57
Bacillariophyta
Naviculoid (small)
6.24
Cyanobacteria
Lyngbya sp2 (thin)
5.77
Bacillariophyta
Cocconeis
3.95
Bacillariophyta
Cymbelloid
2.43
Cyanobacteria
Phormidium
1.95
Bacillariophyta
Gomphonema
1.76
Total
97.48
25
Table 8. Top ten algal taxa in hypereutrophic stream by biomass and density. Percent
represents total from average of samples. For entire taxa lists, see Appendices D and G.
Division
Genus
Percent of Total Sample
Biomass
Bacillariophyta
Naviculoid (large)
15.66
Bacillariophyta
Nitzschia
13.80
Bacillariophyta
Cocconeis
13.75
Bacillariophyta
Achnanthes-like
13.51
Bacillariophyta
Cymbelloid
11.54
Bacillariophyta
Gomphonema
11.43
Bacillariophyta
Diatoma
4.61
Chlorophyta
Schizomeris
3.26
Bacillariophyta
Surrirella
1.65
Bacillariophyta
Naviculoid (small)
1.60
Total
90.82
Density
Cyanobacteria
Blue-green balls
29.62
Bacillariophyta
Achnanthes-like
27.93
Bacillariophyta
Gomphonema
6.78
Cyanobacteria
Lyngbya (thin)
6.38
Bacillariophyta
Naviculoid (large)
5.38
Bacillariophyta
Nitzschia
5.26
Bacillariophyta
Naviculoid (small)
5.13
Cyanobacteria
Lyngbya
4.67
Bacillariophyta
Cocconeis
2.80
Cyanobacteria
Phormidium
2.21
Total
96.17
26
Table 9. Top ten macroinvertebrate taxa in mesotrophic stream by biomass and density.
Percent represents total from average of samples. Under family, (L) represents larva, (P)
represents pupa, (A) represents adult. Under genus, S.F. represents subfamily, T.
represents tribe. For entire taxa lists, see Appendices I and L.
Order
Family
Genus
Guild
Percent of
Total Sample
Biomass
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
22.81
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
14.21
Diptera
Chironomidae (P)
“Other”
13.77
Amphipoda
Gammaridae
Gammarus
Scraper
10.15
Plecoptera
Perlidae
Agnetina
Predator
9.15
Trichoptera
Polycentropodidae
Polycentropus
Predator
6.05
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
5.26
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
4.21
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
2.80
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
1.90
Total
90.30
Density
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
37.42
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
18.71
Diptera
Chironomidae (P)
Trichoptera
Hydropsychidae
Diptera
“Other”
9.61
Ceratopsyche
Collector-filterer
6.72
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
6.51
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
4.34
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
3.72
Trichoptera
Polycentropodidae
Polycentropus
Predator
2.89
Diptera
Tipulidae
Antocha
Collector-gatherer
1.34
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
1.34
Total
92.61
27
Table 10. Top ten macroinvertebrate taxa in eutrophic stream by biomass and density.
Percent represents total from average of samples. Under family, (L) represents larva, (P)
represents pupa, (A) represents adult. Under genus, S.F. represents subfamily, T.
represents tribe. For entire taxa lists, see Appendices J and M.
Order
Family
Genus
Guild
Percent of
Total Sample
Biomass
Diptera
Chironomidae (L)
Diptera
Chironomidae (P)
S.F. Tanypodinae
Predator
56.49
“Other”
14.87
10.43
Ephemeroptera Heptageniidae
Maccaffertium
Scraper
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
5.18
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
4.53
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
1.82
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
1.34
Gastropoda
Physidae
Scraper
0.79
Collector-gatherer
0.61
“Other”
0.53
Ephemeroptera Baetidae
Diptera
Baetis
Chironomidae (A)
Total
96.60
Density
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
50.18
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
20.97
Diptera
Chironomidae (P)
Diptera
Chironomidae (L)
Diptera
Chironomidae (L)
Acariformes
Hydrachnidia
Trichoptera
Hydropsychidae
Collembola
Diptera
“Other”
8.46
T. Chironomini
Collector-gatherer
5.96
S.F. Orthocladiinae
Collector-gatherer
4.69
“Other”
3.69
Ceratopsyche
Collector-filterer
1.23
Poduridae
Podura aquatica
Collector-gatherer
0.73
Tipulidae
Antocha
Collector-gatherer
0.59
Maccaffertium
Scraper
0.50
Ephemeroptera Heptageniidae
Total
97.00
28
Table 11. Top ten macroinvertebrate taxa in hypereutrophic stream by biomass and
density. Percent represents total from average of samples. Under family, (L) represents
larva, (P) represents pupa, (A) represents adult. Under genus, S.F. represents subfamily,
T. represents tribe. For entire taxa lists, see Appendices K and N.
Order
Family
Genus
Guild
Percent of
Total Sample
Biomass
Trichoptera
Hydropsychidae
Diptera
Chironomidae (P)
Amphipoda
Gammaridae
Diptera
Ceratopsyche
Collector-filterer
28.90
Non-feeding
20.11
Gammarus
Scraper
16.66
Chironomidae (L)
S.F. Tanypodinae
Predator
15.89
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
8.26
Diptera
Tipulidae
Antocha
Collector-gatherer
2.27
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
1.69
Collector-gatherer
1.16
Collector-gatherer
0.87
Non-feeding
0.86
Ephemeroptera Baetidae
Diptera
Chironomidae (L)
Diptera
Simuliidae (P)
T. Chironomini
Total
96.68
Density
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
39.28
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
19.01
Diptera
Chironomidae (P)
“Other”
12.72
Diptera
Tipulidae
Antocha
Collector-gatherer
7.35
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
6.34
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
4.05
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
2.89
Gastropoda
Planorbidae
Menetus dilatatus
Scraper
1.32
Acariformes
Hydrachnidia
“Other”
0.81
Amphipoda
Gammaridae
Scraper
0.71
Gammarus
Total
94.48
29
Algal and macroinvertebrate species richness exhibited significant differences
among the streams (two-way ANOVA, F2,35=8.166, p=0.001 and F2,35=10.738, p<0.001,
respectively) but not nutrient treatments (F3,35=0.508, p=0.679 and F3,35=0.418, p=0.741,
respectively). Tukey post-hoc tests revealed that the mesotrophic stream had significantly
more algal taxa than the hypereutrophic stream (p=0.001) but not more than the eutrophic
stream (p=0.080). Macroinvertebrate taxa again showed a different pattern, with the
hypereutrophic stream supporting more macroinvertebrate taxa than the mesotrophic or
eutrophic streams (p=0.003 and <0.001, respectively). Algal and macroinvertebrate
diversity varied by stream (two-way ANOVA: F2,35=5.955, p=0.006; F2,35=20.297,
p<0.001, respectively) but not by nutrient treatment (F3,35=2.580, p=0.069; F3,35=2.459,
p=0.079, respectively). Tukey post-hoc tests revealed significantly higher algal diversity
in the mesotrophic stream from the eutrophic (p=0.008) and hypereutrophic (p=0.030)
streams; the eutrophic stream had the lowest macroinvertebrate diversity (p<0.001 for
both). Slight treatment differences were found in algae between the P (lowest diversity)
and control (highest diversity) treatments (p=0.060), while the largest macroinvertebrate
treatment differences were between the P and N+P treatments, with P having lower
diversity (p=0.054). The Hmax values for Brillouin’s Diversity showed that the algal and
macroinvertebrate maximum diversity varied significantly by stream only (two-way
ANOVA, F2,35=7.509, p=0.002; F2,35=14.102, p<0.001, respectively); Tukey post-hoc
tests showed opposite patterns between the two. Algal Hmax values were higher in the
mesotrophic stream than the hypereutrophic stream (p=0.002), while macroinvertebrate
Hmax values were highest in the hypereutrophic stream (mesotrophic to hypereutrophic:
p<0.001; eutrophic to hypereutrophic: p=0.003). Two-way ANOVAs on Brillouin’s
30
Evenness for algae and macroinvertebrates indicated significant differences by both
stream (F2,35=3.960, p=0.028; F2,35=18.083, p<0.001, respectively) and treatment
(F3,35=3.673, p=0.021; F3,35=3.666, p=0.021). Algal evenness was significantly higher in
the mesotrophic stream than the eutrophic stream (Tukey post-hoc test, p=0.023), and the
control treatments were significantly higher than both N (p=0.026) and P (p=0.032).
Macroinvertebrate evenness was significantly different between all three streams
(mesotrophic to eutrophic: p<0.001; mesotrophic to hypereutrophic: p=0.050; eutrophic
to hypereutrophic: p=0.004; eutrophic<hypereutrophic<mesotrophic), and N treatments
were significantly more even than P treatments (p=0.013).
31
Table 12. Algal and macroinvertebrate (macro) diversity measures (Brillouin’s Indices;
description in methods) among streams and nutrient treatments. Numbers represent
means ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments,
N+P=dual nutrient treatments.
Algal
Diversity
Algal
Hmax
Algal
Evenness
Macro
Diversity
Macro
Hmax
Macro
Evenness
2.060
(0.092)
3.225
(0.101)
0.639
(0.011)
1.691
(0.133)
2.203
(0.097)
0.768
(0.056)
N
1.882
(0.195)
3.224
(0.111)
0.583
(0.041)
1.600
(0.106)
2.014
(0.114)
0.796
(0.062)
P
1.930
(0.060)
3.227
(0.060)
0.598
(0.009)
1.535
(0.126)
2.120
(0.061)
0.725
(0.068)
N+P
2.014
(0.062)
3.227
(0.073)
0.624
(0.016)
1.723
(0.267)
2.177
(0.239)
0.791
(0.062)
Stream Average
1.972
(0.126)
3.226
(0.079)
0.611
(0.030)
1.638
(0.171)
2.128
(0.149)
0.770
(0.063)
1.961
(0.135)
3.184
(0.105)
0.616
(0.041)
1.315
(0.095)
2.088
(0.227)
0.638
(0.108)
N
1.759
(0.187)
3.166
(0.061)
0.555
(0.052)
1.510
(0.043)
2.194
(0.141)
0.690
(0.037)
P
1.683
(0.092)
3.102
(0.180)
0.544
(0.031)
1.321
(0.154)
2.340
(0.102)
0.566
(0.073)
N+P
1.869
(0.264)
3.139
(0.142)
0.594
(0.058)
1.466
(0.238)
2.241
(0.254)
0.652
(0.040)
Stream Average
1.818
(0.196)
3.148
(0.120)
0.577
(0.052)
1.403
(0.162)
2.216
(0.195)
0.636
(0.079)
1.853
(0.053)
3.028
(0.103)
0.613
(0.026)
1.683
(0.104)
2.435
(0.121)
0.693
(0.072)
N
1.891
(0.046)
3.134
(0.071)
0.604
(0.021)
1.786
(0.109)
2.412
(0.077)
0.742
(0.058)
P
1.820
(0.052)
2.995
(0.050)
0.608
(0.007)
1.636
(0.144)
2.378
(0.082)
0.687
(0.040)
N+P
1.796
(0.148)
3.138
(0.151)
0.573
(0.043)
1.754
(0.078)
2.407
(0.060)
0.730
(0.037)
Stream Average
1.841
(0.087)
3.079
(0.113)
0.599
(0.030)
1.720
(0.112)
2.410
(0.081)
0.715
(0.054)
Mesotrophic
Control
Eutrophic
Control
Hypereutrophic
Control
32
Algal (Table 13) and macroinvertebrate (Table 14) group biomass varied among
streams (MANOVA: Wilks’ Lambda=0.219, F8,64=9.078, p<0.001; Wilks’
Lambda=0.067, F10,62=17.716, p<0.001, respectively). Further two-way ANOVA analysis
indicated significant differences in Bacillariophyta, Chlorophyta, and Cyanobacteria algal
groups between streams (p<0.001 for Bacillariophyta and Cyanobacteria, p=0.013 for
Chlorophyta; Table 15, Figure 6). Tukey post-hoc tests revealed significantly more
Bacillariophyta in the mesotrophic stream than the other two (p<0.001 for both
comparisons) and more in the eutrophic stream than in the hypereutrophic (p=0.027),
while Cyanobacteria was significantly higher in the mesotrophic stream than the other
two (p≤0.001 for both). Chlorophyta was significantly higher in the mesotrophic and
hypereutrophic than the eutrophic stream (mesotrophic>eutrophic, p=0.042;
hypereutrophic>eutrophic, p=0.020). Two-way ANOVA indicated all macroinvertebrate
guild biomass but scrapers varied by stream (Table 16; Figure 7). Tukey post-hoc tests
showed all three streams were significantly different from each other in collector-gatherer
biomass, with lowest biomass in the mesotrophic stream and highest in the
hypereutrophic stream (mesotrophic to both eutrophic and hypereutrophic: p<0.001;
eutrophic to hypereutrophic: p=0.010). Collector-filterer biomass was significantly lower
in the eutrophic stream than both the mesotrophic stream (p=0.002) and the
hypereutrophic stream (p<0.001). Predator biomass was highest in the eutrophic stream
(p<0.001 for both comparisons). All three streams were significantly different in
macroinvertebrate “other” biomass, with the mesotrophic stream having the least biomass
(p<0.001 for both comparisons), and the hypereutrophic stream having the highest
biomass (mesotrophic to hypereutrophic p<0.001; eutrophic to hypereutrophic p=0.046).
33
Table 13. Algal biomass (mg DW/m2) for each group by stream and treatment. “Other”
includes Rhodophyta, Euglenophyta, and Chrysophyta. Numbers represent means ±1
standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient
treatments.
Bacillariophyta Chlorophyta Cyanobacteria “Other”
Mesotrophic
Control
N
P
N+P
Stream Average
Eutrophic
Control
N
P
N+P
Stream Average
Hypereutrophic
Control
N
P
N+P
Stream Average
6575.00
(1351.61)
7008.99
(1557.52)
7424.09
(1376.19)
7105.98
(1617.06)
7028.52
(1360.36)
208.38
(345.78)
96.60
(118.96)
65.80
(19.57)
304.06
(497.74)
168.71
(293.06)
249.06
(200.56)
323.50
(100.92)
169.55
(95.04)
519.60
(673.38)
315.43
(347.22)
0.72
(1.16)
0.09
(0.10)
0.05
(0.09)
0.04
(0.08)
0.22
(0.60)
4422.70
(1232.08)
3849.53
(1127.97)
2311.24
(494.52)
5100.11
(653.58)
3920.89
(1350.06)
73.37
(90.00)
14.10
(20.04)
24.49
(26.47)
75.38
(137.54)
46.83
(80.30)
96.37
(72.49)
73.71
(13.68)
44.94
(32.70)
145.23
(151.44)
90.06
(85.59)
0.06
(0.07)
0.19
(0.09)
0.10
(0.07)
2.27
(3.72)
0.66
(1.93)
2660.99
(848.56)
2793.53
(794.65)
3004.38
(1305.15)
3331.49
(1546.90)
2943.81
(1057.71)
140.21
(159.39)
61.65
(40.72)
148.80
(90.98)
311.85
(514.28)
166.75
(270.37)
70.05
(27.95)
72.49
(15.19)
63.10
(56.17)
136.81
(135.73)
87.11
(74.76)
0.11
(0.08)
2.69
(5.07)
0.18
(0.31)
0.16
(0.24)
0.83
(2.62)
34
Table 14. Macroinvertebrate biomass (mg DW/m2) for each group by stream and
treatment. “Other” includes parasites, pupae, adults, scavengers, and shredders. Numbers
represent means ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments,
N+P=dual nutrient treatments.
Mesotrophic
Control
N
P
N+P
Stream
Average
Eutrophic
Control
N
P
N+P
Stream
Average
Hypereutrophic
Control
N
P
N+P
Stream
Average
Collector-gatherer
Collector-filterer
Scraper
Predator
“Other”
11.74
(5.00)
11.72
(9.07)
17.17
(6.91)
19.65
(11.59)
15.07
(8.40)
62.83
(55.51)
61.88
(30.60)
19.28
(9.31)
42.95
(32.44)
46.73
(36.97)
20.74
(18.32)
88.47
(114.23)
11.67
(18.32)
35.04
(28.99)
38.98
(62.10)
73.63
(27.16)
35.49
(18.16)
91.79
(28.12)
135.06
(173.50)
83.99
(88.09)
29.50
(16.79)
32.22
(11.43)
21.91
(4.93)
37.37
(15.96)
30.25
(13.10)
29.40
(23.79)
27.73
(11.16)
54.25
(10.69)
37.02
(19.90)
37.10
(18.91)
7.67
(15.34)
19.64
(16.53)
19.29
(21.54)
42.67
(77.80)
22.32
(39.71)
56.63
(108.61)
31.53
(21.37)
39.75
(73.14)
58.92
(67.36)
46.71
(67.59)
193.60
(156.09)
157.46
(45.15)
343.48
(155.82)
223.08
(27.10)
229.40
(124.45)
47.72
(8.89)
76.30
(33.06)
79.96
(38.55)
54.52
(28.14)
64.62
(29.87)
86.04
(30.29)
51.34
(18.17)
70.18
(14.18)
47.48
(22.23)
63.33
(25.93)
172.88
(81.81)
78.35
(38.59)
110.08
(85.54)
134.95
(140.49)
125.00
(91.55)
47.94
(59.85)
79.01
(84.06)
146.22
(204.77)
47.83
(36.43)
75.85
(100.33)
73.47
(38.60)
66.83
(35.91)
78.48
(63.11)
60.79
(37.60)
69.32
(38.90)
131.67
(24.74)
70.91
(24.89)
111.81
(49.78)
64.53
(15.75)
93.59
(39.30)
35
Table 15. Two-way ANOVA results for algal group biomass (mg DW/m2) and density
(cells/cm2) showing F values with p-values in parentheses.
Factor
Biomass
Stream
Nutrient Treatment
Stream *Nutrient
Treatment
Density
Stream
Nutrient Treatment
Stream*Nutrient
Treatment
Bacillariophyta
Chlorophyta
Cyanobacteria
“Other”
40.450 (<0.001)
1.760 (0.173)
2.099 (0.078)
4.961 (0.013)
0.617 (0.608)
0.251 (0.956)
12.738 (<0.001)
2.111 (0.116)
0.085 (0.997)
0.366 (0.696)
0.572 (0.637)
1.489 (0.211)
38.646 (<0.001)
1.025 (0.393)
1.443 (0.226)
15.167 (<0.001)
1.008 (0.401)
0.603 (0.726)
13.548 (<0.001)
2.310 (0.093)
0.094 (0.997)
2.288 (0.116)
1.206 (0.322)
1.876 (0.113)
Table 16. Two-way ANOVA results for macroinvertebrate guild biomass (mg DW/m2)
and density (number/m2) showing F values with p-values in parentheses.
Factor
Biomass
Stream
Nutrient Treatment
Stream *Nutrient
Treatment
Density
Stream
Nutrient Treatment
Stream*Nutrient
Treatment
CollectorGatherer
CollectorFilterer
Predator
Scraper
“Other”
31.458
(<0.001)
2.040
(0.126)
1.132
(0.364)
16.990
(<0.001)
0.230
(0.875)
1.089
(0.388)
17.286
(<0.001)
1.630
(0.200)
0.735
(0.625)
1.060
(0.357)
0.542
(0.647)
0.217
(0.969)
29.737
(<0.001)
0.261
(0.853)
2.239
(0.062)
43.049
(<0.001)
2.504
(0.075)
1.310
(0.279)
25.971
(<0.001)
1.654
(0.195)
4.465
(0.002)
22.920
(<0.001)
2.301
(0.094)
0.590
(0.736)
7.202
(0.002)
0.732
(0.540)
0.659
(0.683)
33.025
(<0.001)
0.039
(0.989)
1.517
(0.201)
36
Percent Biomass (mg DW/m2)
A
100%
90%
80%
70%
60%
Other
50%
Cyanobacteria
40%
Chlorophyta
30%
Bacillariophyta
20%
10%
0%
Control
Percent Biomass (mg DW/m2)
B
N
P
N+P
100%
90%
80%
70%
Other
60%
50%
Cyanobacteria
40%
Chlorophyta
30%
Bacillariophyta
20%
10%
0%
C
Percent Biomass (mg DW/m2)
Control
N
P
N+P
100%
90%
80%
70%
60%
50%
40%
30%
20%
10%
0%
Other
Cyanobacteria
Chlorophyta
Bacillariophyta
Control
N
P
N+P
Figure 6. Biomass distribution (mg DW/m2) of algal groups by stream and treatment
with Bacillariophyta at the bottom and “Other” at the top; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments. A) Mesotrophic stream B)
Eutrophic stream C) Hypereutrophic stream.
37
Percent Biomass (mg DW/m2)
A
100%
90%
80%
70%
Other
60%
Scraper
50%
Predator
40%
Collector-filterer
30%
Collector-gatherer
20%
10%
0%
Control
B
N
P
N+P
Percent Biomass (mg DW/m2)
100%
90%
80%
70%
Other
60%
Scraper
50%
Predator
40%
Collector-filterer
30%
Collector-gatherer
20%
10%
0%
Control
C
N
P
N+P
Percent Biomass (mg DW/m2)
100%
90%
80%
70%
Other
60%
Scraper
50%
Predator
40%
Collector-filterer
30%
Collector-gatherer
20%
10%
0%
Control
N
P
N+P
Figure 7. Biomass distribution (mg DW/m2) of macroinvertebrate guilds by stream and
treatment with Collector-gatherer at the bottom and “Other” at the top; N=nitrogen
treatments, P=phosphorus treatments, N+P=dual nutrient treatments. A) Mesotrophic
stream B) Eutrophic stream C) Hypereutrophic stream.
38
The density of both algal (Table 17) and macroinvertebrate groups (Table 18)
varied by stream only (MANOVA: Wilks’ Lambda=0.256, F8,64=7.824, p<0.001; Wilks’
Lambda=0.025, F10,62=32.796, p<0.001, respectively). Two-way ANOVA indicated
significant differences among all algal groups but those listed as “other” (Table 15;
Figure 8). Tukey post-hoc tests showed the mesotrophic stream had the highest
Bacillariophyta, Chlorophyta, and Cyanobacteria densities (p≤0.001 in all comparisons).
Moreover, two-way ANOVA indicated differences by stream in all five
macroinvertebrate groups by density (Table 16; Figure 9). Tukey post-hoc tests indicated
lowest collector-gather densities were in the mesotrophic stream and highest in the
hypereutrophic stream (mesotrophic to both eutrophic and hypereutrophic: p<0.001;
eutrophic to hypereutrophic: p=0.004). Collector-filterer density was lowest in the
eutrophic stream (p<0.001 in both comparisons) while predator density was highest in the
eutrophic stream (p<0.001 for both comparisons). Scraper density was higher in the
hypereutrophic stream than in the eutrophic stream (p=0.002). “Other” density was
lowest in the mesotrophic stream (p<0.001 for both comparisons). A significant
difference was seen by treatment in predators, where density on N treatments was less
than P treatments (p=0.039, respectively).
39
Table 17. Algal density (cells/cm2) for each group by stream and treatment. “Other”
includes Rhodophyta, Euglenophyta, and Chrysophyta. Numbers represent means ±1
standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient
treatments.
Bacillariophyta Chlorophyta Cyanobacteria
Mesotrophic
Control
N
P
N+P
Stream
Average
Eutrophic
Control
N
P
N+P
Stream
Average
Hypereutrophic
Control
N
P
N+P
Stream
Average
“Other”
1.36x106
(3.45x105)
1.45x106
(3.68x105)
1.62x106
(3.83x105)
1.51x106
(3.11x105)
1.49x106
(3.30x105)
1.76x105
(3.42x105)
7.53x103
(6.27x103)
2.58x104
(3.74x104)
1.57x104
(1.78x104)
5.64x104
(1.70x105)
1.15x106
(6.66x105)
1.87x106
(9.02x105)
9.64x105
(3.87x105)
1.48x106
(8.91x105)
1.37x106
(7.54x105)
7.40x101
(1.08x102)
1.29x101
(1.50x101)
6.86x100
(1.37x101)
6.08x100
(1.22x101)
2.50x101
(5.76x101)
6.90x105
(2.85x105)
7.08x105
(1.85x105)
3.99x105
(8.88x104)
8.37x105
(2.55x105)
6.58x105
(2.55x105)
3.35x103
(2.64x103)
5.84x102
(2.57x102)
1.04x103
(6.00x102)
3.56x103
(5.89x103)
2.13x103
(3.21x103)
5.48x105
(4.93x105)
5.21x105
(9.92x104)
3.14x105
(2.30x105)
6.29x105
(3.76x105)
5.03x105
(3.22x105)
9.00x100
(1.06x101)
2.88x101
(1.41x101)
1.47x101
(1.11x101)
8.86x101
(5.54x101)
3.53x101
(4.20x101)
5.75x105
(1.77x105)
5.97x105
(1.57x105)
6.55x105
(3.21x104)
6.84x105
(3.34x105)
6.26x105
(2.30x105)
1.51x103
(1.74x103)
1.58x103
(6.16x102)
1.93x103
(7.25x102)
2.55x103
(2.68x103)
1.89x103
(1.59x103)
3.67x105
(1.39x105)
5.15x105
(1.12x105)
4.51x105
(4.01x105)
5.61x105
(2.32x105)
4.75x105
(2.18x105)
1.70x101
(1.17x101)
6.23x101
(7.99x101)
2.70x101
(4.68x101)
2.41x101
(3.61x101)
3.30x101
(4.83x101)
40
Table 18. Macroinvertebrate density (number/m2) for each group by stream and
treatment. “Other” includes parasites, pupae, adults, scavengers, and shredders. Numbers
represent means ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments,
N+P=dual nutrient treatments.
Mesotrophic
Control
N
P
N+P
Stream Average
Eutrophic
Control
N
P
N+P
Stream Average
Hypereutrophic
Control
N
P
N+P
Stream Average
Collector-gatherer
Collector-filterer
Scraper
Predator
“Other”
477.05
(189.05)
325.51
(113.73)
426.53
(135.94)
516.33
(255.96)
436.36
(178.84)
207.66
(102.67)
145.92
(38.88)
89.80
(40.99)
185.21
(107.47)
157.14
(84.79)
39.29
(21.49)
50.51
(46.28)
33.67
(38.88)
84.19
(64.48)
51.91
(45.43)
594.90
(183.76)
409.70
(242.05)
667.86
(204.42)
578.07
(318.21)
562.63
(237.93)
129.08
(74.17)
140.31
(49.78)
95.41
(21.49)
162.76
(69.50)
131.89
(57.31)
796.95
(359.42)
796.95
(418.58)
1694.92
(372.28)
886.74
(379.43)
1043.89
(519.16)
5.61
(11.23)
72.96
(49.78)
61.74
(46.28)
61.74
(94.14)
50.51
(58.83)
22.45
(31.75)
28.06
(11.22)
16.84
(11.22)
28.06
(21.49)
23.85
(19.17)
1240.32
(626.00)
1161.75
(405.90)
2570.43
(707.66)
1273.99
(278.21)
1561.62
(768.49)
280.62
(53.44)
482.66
(277.08)
370.41
(158.21)
415.31
(299.23)
387.25
(211.15)
1745.43
(357.02)
1543.38
(658.18)
1990.50
(810.77)
1341.34
(508.42)
1632.81
(571.87)
291.84
(115.93)
202.04
(133.44)
172.11
(64.81)
162.76
(95.90)
209.53
(110.20)
61.74
(21.49)
84.19
(61.82)
97.28
(34.29)
117.86
(59.04)
89.80
(48.00)
550.01
(241.44)
544.39
(299.72)
636.06
(342.92)
578.07
(362.62)
573.20
(279.22)
544.39
(90.50)
331.13
(131.38)
493.88
(214.15)
308.68
(71.87)
414.56
(156.43)
41
Percent density (cells/cm2)
A
100%
90%
80%
70%
60%
Other
50%
Cyanobacteria
40%
Chlorophyta
30%
Bacillariophyta
20%
10%
0%
Control
B
N
P
N+P
100%
Percent density (cells/cm2)
90%
80%
70%
60%
Other
50%
Cyanobacteria
40%
Chlorophyta
30%
Bacillariophyta
20%
10%
0%
Control
C
N
P
N+P
Percent density (cells/cm2)
100%
90%
80%
70%
Other
60%
50%
Cyanobacteria
40%
Chlorophyta
30%
Bacillariophyta
20%
10%
0%
Control
N
P
2
N+P
Figure 8. Density distribution (cells/cm ) for algal groups by stream and treatment with
Bacillariophyta at the bottom and “Other” at the top; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments. A) Mesotrophic stream B)
Eutrophic stream C) Hypereutrophic stream.
42
A
Percent Density (number/m2)
100%
90%
80%
70%
Other
60%
Scraper
50%
Predator
40%
Collector-filterer
30%
Collector-gatherer
20%
10%
0%
Control
B
N
P
N+P
Percent Density (number/m2)
100%
90%
80%
70%
Other
60%
Scraper
50%
Predator
40%
Collector-filterer
30%
Collector-gatherer
20%
10%
0%
Control
C
N
P
N+P
Percent Density (number/m2)
100%
90%
80%
70%
Other
60%
Scraper
50%
Predator
40%
Collector-filterer
30%
Collector-gatherer
20%
10%
0%
Control
N
P
2
N+P
Figure 9. Density distribution (number/m ) for macroinvertebrate guilds by stream and
treatment with Collector-gatherer at the bottom and “Other” at the top; N=nitrogen
treatments, P=phosphorus treatments, N+P=dual nutrient treatments. A) Mesotrophic
stream B) Eutrophic stream C) Hypereutrophic stream.
43
Trophic Relationships
Algal biomass (as chlorophyll-a) and macroinvertebrate biomass followed
opposite patterns, with algal biomass decreasing as productivity increased, while
macroinvertebrate biomass increased as productivity increased. Broken down by guild,
grazers (collector-gatherers and scrapers) are seen to drive this pattern, while predators
show a peak at the eutrophic stream (Figure 10). Doubling time of the algal dry weight
was significantly correlated to macroinvertebrate biomass and density (r=0.411, p=0.004,
and r=0.538, p<0.001, respectively, n=47), with the direction indicating that doubling
time increases with increasing macroinvertebrate biomass and density. Additionally, the
doubling time of the macroinvertebrate biomass was significantly shorter than the
doubling time of the algal dry weight (paired T-test: T=9.31, p<0.001, n=47), while the
doubling time of the macroinvertebrate population density was significantly longer than
that of the algal dry weight (paired T-test: T= -10.47, p<0.001, n=47). Likewise, the algal
biomass doubling time was significantly correlated to the doubling time of
macroinvertebrate biomass and density (r= -0.464, p=0.001, n=47; r= -0.564, p<0.001,
n=47, respectively). Moderate correlations were found between algal dry weight and
macroinvertebrate density (r= -0.545, p<0.001, n=47) and biomass (r= -0.385, p=0.008,
n=47). Macroinvertebrate density was also found to influence the species richness of
algae (r= -0.335, p=0.021, n=47). For both macroinvertebrate and algal biomass, species
richness and biomass of the respective trophic level showed positive correlations
(r=0.381, p=0.003, n=47; r=0.410, p=0.004, n=47, respectively) in that as biomass of that
trophic level increased, the species richness increased as well.
44
A
400
Biomass (mg DW/m2)
350
300
250
Control
200
N
150
P
100
N+P
50
0
Mesotrophic
B
Eutrophic
Hypereutrophic
Biomass (mg DW/m2)
250
200
Control
150
N
100
P
N+P
50
0
Mesotrophic
C
Eutrophic
Hypereutrophic
90
Biomass (mg Chl-a/m2)
80
70
60
Control
50
N
40
P
30
N+P
20
10
0
Mesotrophic
Eutrophic
Hypereutrophic
Figure 10. Mean biomass of guilds by treatment and stream; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments. A) Dry weight of predators B)
Dry weight of grazers (collector-gatherers and scrapers) C) Biomass of algae as
chlorophyll-a.
45
Discussion
Algal and macroinvertebrate biomass did not significantly respond to
experimental nutrient enrichment across the trophic gradient but did vary among the
streams. The responses at the primary producer (algae) and primary consumer
(herbivorous macroinvertebrates) levels showed significant negative correlation to each
other, indicating tight coupling between the trophic levels rather than direct linear
increases at both levels due to nutrient enrichment. Additionally, diversity did not show a
typical unimodal response (e.g., Leibold 1999); rather, lowest diversity was found for
both algae and macroinvertebrates in the eutrophic stream. Nutrient enrichment within
the streams appeared to have some influence upon diversity measures, especially
evenness. These results seem to indicate that varying complexity in median trophic levels
(especially macroinvertebrates) across the trophic gradient served to mediate responses
between primary producers and higher trophic levels.
Biomass Responses to Nutrient Enrichment
In the mesotrophic stream, a trend (though non-significant) of increasing algal
biomass (chlorophyll-a) with nutrient addition was observed: algal biomass increased in
both the N and P treatments over the controls, and the dual N+P nutrient treatment
showed the highest biomass. This response is typical of bio-limiting nutrient additions;
synergistic influence of dual nutrient enrichment on primary productivity is well
documented in the literature (e.g., Elser et al. 2007; Fairchild et al. 1985; Francoeur
2001). The lowest numbers of macroinvertebrates were present in the mesotrophic
stream, decreasing grazing pressure (e.g., Lamberti and Resh 1983; Rosemond et al.
46
1993) and possibly allowing the algae to respond to nutrient addition (Hillebrand 2002),
although top-down force was still strong enough to prevent a significant response. The
highest macroinvertebrate biomass in the mesotrophic stream was on the N+P treatments,
a pattern similar to the algae and dissimilar to the other streams where differences
occurred between N and P treatments. This seems to suggest that bottom-up effects (seen
through nutrient enrichment) on algal biomass were strongest in the mesotrophic stream
and were conserved at the next trophic level, perhaps intimating a connection between
top-down and bottom-up influences whereby both are influencing the food chain but
one’s effects are stronger at any given time (Gruner et al. 2008; McIntosh et al. 2005;
Rosemond et al. 1993).
The eutrophic and hypereutrophic streams showed little to no response to
nutrients, indicating they may not have been limited by nutrients at this time. There is
ample evidence that just as often as nutrient enrichment increases algal biomass, it fails to
produce a response (Francoeur 2001). These streams typically support high nutrient
concentrations and corresponding benthic chlorophyll-a concentrations (Carrick et al.
2009), and while the nutrient loads provided in this experiment were higher than ambient
conditions, another factor may be more strongly limiting than nutrients. Limitation by
macroinvertebrate grazing is a likely factor (Corkum 1996; Rosemond et al. 1993), given
that in the eutrophic and hypereutrophic streams macroinvertebrate biomass comprised a
greater portion of the total biomass, and therefore they may have exerted more influence
on the algal biomass than those in the mesotrophic stream. Macroinvertebrates can
consume large amounts of algae (Holomuzki et al. 2010); in fact, Chironomidae, the
dominant family in all three streams (63.6-95.8% of macroinvertebrate density and 27.1-
47
97.7% of macroinvertebrate biomass), have been found to consume up to 125% of algal
standing crop at their highest densities when examined as a group (Goldfinch and
Carman 2000). Other studies have similarly found that grazing by macroinvertebrates can
be a considerable factor governing algal biomass (e.g., Lamberti and Resh 1983;
Lamberti et al. 1987; Wallace and Webster 1996). The highest densities of primary
consumers, which included mostly herbivorous Chironomidae such as Micropsectra and
Cricotopus (herbivorous Chironomidae range: 23.5-84.3% of Chironomidae density; 5.759.2% of Chironomidae biomass across all three streams), were found in the eutrophic
and hypereutrophic streams. Thus, it follows that higher algal consumption rates (i.e.,
grazing) occurred in these streams relative to the mesotrophic stream, and if grazing rates
were even moderate, then grazing rates would likely have outperformed algal growth.
Accordingly, the lower algal biomass observed in the eutrophic and hypereutrophic
streams seems reasonable, particularly given the high macroinvertebrate densities that
were inversely related to algae. Additionally, grazers have been known to drive increased
algal turnover times (Lamberti and Resh 1983), which in turn can support high secondary
production, even up to 10-20 times primary producer biomass (Steinman et al. 2006). For
this to occur, tight coupling between trophic levels is necessary and has often been seen
between stream macroinvertebrates and algae (Wallace and Webster 1996). This
experiment supported this idea in that the eutrophic and hypereutrophic streams, where
more macroinvertebrates were present, algal biomass was lower and biomass doubling
times were longer than the mesotrophic stream. Interestingly, algal doubling time was
positively correlated to macroinvertebrate biomass/density, again suggesting tight
coupling between the two trophic levels (Lamberti and Resh 1983). While the specific
48
food habits of a macroinvertebrate group may vary between sites (Cummins 1973), the
evidence in this study nonetheless indicates a strong relationship between algae and
macroinvertebrates, likely driven by grazing.
Diversity Response to Nutrient Enrichment
The mesotrophic stream exhibited a more balanced (greater Brillouin’s Evenness)
and diverse algal and macroinvertebrate community compared with the other two
streams. The algal assemblage here was generally comprised of smaller Cyanobacteria
and Chlorophyta cells relative to the eutrophic or hypereutrophic stream. For instance,
spherical Cyanobacteria such as Merismopedia and Microcystis were common as were
the chlorophytes Pediastrum and Scenedesmus. Various species in these genera have
been associated with both oligotrophic and eutrophic waters (Garg and Garg 2002;
Wrigley et al. 1991); they were therefore already likely present in the mesotrophic stream
and persisted with nutrient enrichment, rather than being replaced by other taxa
associated with only eutrophic waters. The small-celled filamentous chlorophyte
Stigeoclonium and filamentous diatom Melosira were also commonly found in this
stream. Two factors may be contributing to the dominance of smaller cells and these
filamentous organisms. The lower grazing pressure (fewer macroinvertebrates) present
here may have led to the establishment of Stigeoclonium and Melosira as similarly
observed by Rosemond et al. (1993) and DeNicola et al. (1990) in ungrazed treatments.
Additionally, lower grazing pressure may have led to dominance by less easily consumed
smaller cells due either to algal prostrate growth morphology and/or lack of appropriate
mouthparts for grazing a particular cell shape/size (DeNicola et al. 1990). The
49
macroinvertebrate community in the mesotrophic stream showed the least number of
Chironomidae, and also had various Trichoptera rarely encountered in the other streams,
especially the genera Neureclipsis, Polycentropus, Psychomyia, and Helicopsyche as nonpredatory macroinvertebrates. Similar to the eutrophic stream, there were also many
Ceratopsyche net-spinning caddisflies. These organisms are likely to be important in the
grazing and collecting of algal material in this stream.
Although the hypereutrophic stream contained a greater percentage of larger,
filamentous algal taxa such as Schizomeris, Oedogonium, Cylindrocapsa, and
Cladophora (often considered an indicator of eutrophic conditions; Kelly and Whitton
1988) than the other streams, they were still in low numbers with diatoms being the
primary organisms. Diatom dominance is common in strongly grazed algal communities,
where the algal community is kept in an early successional stage by preferential feeding
on filamentous organisms (e.g., DeNicola et al. 1990; Lamberti and Resh 1983). Indeed,
the most collector-gatherers were found in the hypereutrophic stream, comprised mainly
of the chironomid tribe Tanytarsini (e.g., Micropsectra, Paratanytarsus), with fewer
Orthocladiinae (e.g., Cricotopus) and Chironomini (e.g., Dictrotendipes, Polypedilum), as
were the most scrapers, consisting mostly of various Gastropoda and the amphipod genus
Gammarus. These organisms could have driven the algal community composition to
diatom dominance over the filamentous organisms and also the low algal biomass,
suggesting top-down control through grazing.
The algal taxa in the eutrophic stream appeared to comprise an intermediate
assemblage taxonomically between the other two streams, with many of the smaller taxa
found in the mesotrophic stream and also the filamentous taxa found in the
50
hypereutrophic stream. Collector-filterers were least common in this stream, with a noted
decrease in the trichopteran genus Ceratopsyche which was common in the other streams.
While it is possible that they were emerging and therefore missed, no pupae were found
as they were for Hydroptilidae; hydropsychids such as Ceratopsyche are known to be
preyed upon by sculpins, the numerically dominant fishes in the eutrophic stream, which
may have reduced their numbers (Fairchild and Holomuzki 2005). Since Ceratopsyche is
not a predatory macroinvertebrate, this would provide evidence that lower trophic level
fish were not just preying upon predatory macroinvertebrates, but all trophic levels.
Contrarily, predatory chironomids in the subfamily Tanypodinae were common in this
stream compared with the macroinvertebrates present in the mesotrophic and
hypereutrophic streams. The Tanypodinae were variable in size (although lengths were
significantly greater here than the mesotrophic stream), and therefore may have been
feeding on both algal matter and other chironomids (Baker and McLachlan 1979)
dependent on size/instar, influencing the macroinvertebrate and algal communities
simultaneously and further complicating the trophic dynamics via omnivory. The
probable presence of high omnivory in this stream through sculpin feeding upon all levels
of macroinvertebrates and Tanypodinae feeding on other small macroinvertebrates and
algae would lead to a higher degree of complexity within the median trophic levels in this
stream. This may then prevent direct simple responses from being observable at all
levels, as seems to be the case here in the macroinvertebrates.
A significant difference was observed in the density of predators concurrent with
a slight but likely biologically significant difference in collector-gatherer density between
N and P treatments, which were both comprised primarily of Chironomidae. In addition
51
to being less dense, the assemblage occurring on the N treatments had greater evenness
values, indicating communities that were closer to maximum Brillouin’s diversity, while
the P treatments had the lowest diversity and evenness. These differences may reflect a
dense community on the P treatments dominated by relatively few species. Previous work
has found positive responses in larval Chironomidae to gradients of P (Ramírez and
Pringle 2006), matching results here of highest densities and quickest doubling times.
The macroinvertebrate community as a whole showed a negative response to N addition
in density and doubling time. Armitage and Fong (2004) found nitrogen additions caused
negative responses in Gastropoda due to an increase in Cyanobacteria; Cyanobacteria
were most abundant on the nitrogen treatments in this experiment as well, and may
therefore be a factor in the decrease in density and growth of collector-gatherers and
predators on N treatments in this experiment. A more significant (and therefore likely
stronger) negative response was noticed in predators, possibly indicating an N
bioaccumulation effect in higher trophic levels.
The highest macroinvertebrate diversity was observed on the N+P treatments,
which further supports the contention of differential effects of N and P. For instance,
research has shown that dual additions of N and P can induce a synergistic effect on
primary production (e.g., Elser et al. 2007); these nutrient synergisms appear here to be
conserved or maintained through higher trophic levels such that diversity is increased.
Despite this trend, measures like biomass and density still appear negatively affected by
the presence of N, even when co-enriched with P. In contrast to macroinvertebrate
taxonomic nutrient response, algal control treatments supported the highest evenness,
suggesting that an increase in biomass does not necessarily create a more balanced
52
community. Instead, it may decrease diversity responses; this decrease has previously
been observed in eutrophic systems (e.g., Hautier et al. 2009; Leibold 1999).
Diversity across this trophic gradient did not show a typical unimodal distribution
(Tilman 1982), nor did it follow the monotonic changes observed here in other algal and
macroinvertebrate measures (i.e., biomass and density). Instead, the opposite was seen for
both algae and macroinvertebrates in that the lowest diversity was found in the eutrophic
stream. While the highest diversity in the mesotrophic stream follows the idea of
moderately productive systems harboring high diversity (e.g., Abrams 1995), the fact that
the hypereutrophic stream also had higher diversity than the eutrophic stream is contrary
to both unimodal and increasing monotonic models discussed in Abrams (1995).
Interestingly, though diversity is generally expected to increase monotonically if not
responding unimodally (Abrams 1995; Pringle 1990), it has also been documented to
decrease monotonically with increasing productivity (Goldberg and Miller 1990). Though
some suggest that with a monotonic response only parts of the unimodal curve are being
perceived across the productivity measured, Abrams (1995) discusses that true monotonic
responses are just as likely. This experiment seems to suggest that diversity responses to
productivity gradients may be trophic level specific, such that greater complexity
between and within median trophic levels are shifting typical response curves. For
instance, in the eutrophic stream the prevalence of Tanypodinae was striking (averaging
56.6% of eutrophic biomass and 49.6% of eutrophic density), and was a likely factor in
the high unevenness in the macroinvertebrate community which may have then led to
lower diversity. As previously stated, they were likely behaving as omnivores, yet to
higher trophic levels in the stream were simply a prey choice regardless of trophic
53
position. The unidirectional connection between the Tanypodinae and higher trophic
levels suggests they may be functionally redundant with other macroinvertebrates to
those higher levels. Yet, to lower levels there is a multidirectional connection, and so
though they may be thought to be structurally redundant with other macroinvertebrates
(and may often be treated as such by examination at the family level, e.g., Corkum 1996;
Goldfinch and Carman 2000; Ramírez and Pringle 2006), it is likely they are not
functionally redundant but instead play an integral role in mediating responses between
trophic levels.
Importance of Median Trophic Levels
As discussed above, it is possible that organisms situated at median levels of the
food chain constrain diversity measures and influence biomass and taxonomic
composition. For instance, macroinvertebrates constrained algal species richness,
suggested by the moderate negative relationship and weak bottom-up influences; this was
also observed by Opsahl et al. (2003) via exclusion of macroinvertebrates. Additionally,
the correlations of algal biomass and doubling time with macroinvertebrate biomass and
doubling time suggest tight coupling in the system. The macroinvertebrates were seen to
constitute nearly 30% of biomass in these streams, indicating their importance in energy
transfer in the streams. Their median position would allow them to pass energy from the
primary producers to higher levels present. In all of these streams, higher trophic levels
were present as various fish species, indicating energy to be passing along the chain
through the macroinvertebrates to at least some of the fish, especially as each stream has
a “top predator” that may only be present with ample resource availability (Post 2002).
54
Internal complexity within median levels may be preventing simple top-down or
bottom-up effects from being seen in these streams (Paine 1966). An assessment of algal
and macroinvertebrate biomass and their responses to nutrient enrichment in this
experiment suggests that the primary factor governing biomass levels at this point in time
was not nutrient limitation. It seems likely that consumers played a large role relative to
nutrients, as the mesotrophic stream’s response appears to be that of a three-step food
chain while the eutrophic and hypereutrophic stream appear to respond as four-step food
chains. The fish composition in each of these streams, however, complicates these ideas,
as they do not directly seem to support the idea of strictly three- and four-step food
chains. In the mesotrophic stream, there was an abundance of lower trophic level fish,
especially blacknose dace and other minnows. Yet, many American eels were also
present, holding the top predator role. The number and type of fish present indicates
something other than a three-step chain is functioning in the system, yet the response in
algae and macroinvertebrates still mirrors three-step responses (e.g., Hairston et al. 1960).
The eutrophic stream also shows many lower trophic level fish, primarily sculpins, with
brown trout occupying more of a top predator role. Again, this in addition to the presence
of many predatory Tanypodinae seems to suggest something other than a four-step chain,
yet the response remains that of a four-step in algal response (e.g., Carpenter et al. 1985).
The hypereutrophic stream has less fish overall, but is still occupied by similar lower
trophic level fish and many brown trout, yet algal and macroinvertebrate response seems
to be that of a four-step chain. This cursory look at higher trophic levels in these streams
indicates that there is something more complicated happening than the simple responses
suggested by algal and macroinvertebrate factors in these streams in order for primary
55
production to be supporting these higher trophic levels without a simple response evident
in the fish.
The presence of top predatory fish in each food chain may have influenced the
lower levels through direct or indirect means such as drift feeding (Dahl and Greenberg
1996; Nakano et al. 1999), competition (Holomuzki et al. 2010), growth rates of other
species (Ruetz et al. 2003), and other behavioral influences (Werner and Peacor 2003).
For example, limited direct interaction may be seen between brown trout and lower
trophic levels in summer due to selective feeding on terrestrial insects falling into the
stream (Dahl 1998; Nakano et al. 1999) or emerging aquatic insects (Elliott 1967;
Wagner et al. 2012), but the interaction changes seasonally when trout are more reliant on
autochthonous sources supported by high algal productivity (Bridcut and Giller 1995;
Elliott 1967). Fish predators are longer lived than macroinvertebrates and algae; as such,
rather than the quick turnover times seen in algal populations (Lamberti and Resh 1983;
Steinman et al. 2006), fish biomass generally undergoes slow increases that are
conserved. These situations can be manifested as inverted biomass pyramids (Sandin et
al. 2008) readily observed in streams (Power 2001) and common in offshore ocean
systems (e.g., Gasol et al. 1997). These pyramids are only observable in open systems
(Gasol et al. 1997) or when tight coupling is occurring (Sandin et al. 2008). The
relationships between algae and macroinvertebrates here suggest that there is strong
coupling at the lower levels of these streams which is likely passed on to the higher levels
through high primary production and efficient trophic energy transfers (Sandin et al.
2008); when algal standing crop is low due to high grazing, algal populations must
56
therefore by necessity experience quick turnover mediated by median trophic levels to
pass energy to the top of a food chain.
Conclusions
The experimental responses measured in this study are reasonable as the
experimental design used here appeared to be effective in reflecting patterns seen
spatially and temporally in temperate streams. First, terracotta pot saucers have
previously been utilized as an inert growth surface and nutrient point-source for benthic
communities (e.g., Fairchild et al. 1985; Godwin et al. 2009) which enables a high degree
of replication, controls nutrient release, and provides a uniform surface for growth which
adequately supported an algal and macroinvertebrate community in this experiment. The
algal growth was within ranges previously observed in all three streams (unpublished
data). The macroinvertebrate community was similar to rocks in the surrounding stream
in terms of density, diversity, and species richness: hierarchical cluster analysis did not
separate rocks from NDS (unpublished data), and colonization by Trichoptera occurred in
all three streams with this design. Additionally, all life stages were collected, including
adults, indicating the macroinvertebrates were utilizing the NDS as they would natural
surfaces. The nutrient concentrations used in this experiment have been commonly used
in similar studies (Fairchild et al. 1985; Corkum 1996; Carrick and Price 2011) and lead
to elevated, conserved, and balanced N:P loading rates similar to Fairchild et al. (1985)
based upon surface area to volume ratios. While the groupings (both algal and
macroinvertebrate) used in this experiment assumed that structural redundancy mirrors
functional redundancy at these guild levels, the functional trophic position of a given
57
taxon may vary based upon stream conditions (Cummins 1973). Nevertheless, the use of
functional guilds is common in lotic research, especially for macroinvertebrates (e.g.
Vannote et al. 1980) as food habit studies for a given taxon would need to be conducted
across many of their natural habitats. Since more specific data is not typically available,
the use of functional feeding guilds is justified in experiments such as these where the
greater functioning of food chains across systems is the goal, and not the role of a
specific taxon (Corkum 1992).
Although nutrients did not appear to be strongly limiting growth during this
experiment and responses appeared more consumer driven, experimentation at more time
points in all seasons would be necessary to determine whether top-down consumer or
bottom-up nutrient factors were consistently the dominant controlling factor (Batzer
1998; Boyer et al. 2003; Francoeur 2001; Gasol et al. 1995). Regardless, strong patterns
throughout the functional food chain were not evident here. None of the streams across
the trophic gradient observed here responded in a simple linear fashion; the interactions
were complicated, and while assessed as functional food chains (e.g., Post 2002), these
lotic systems clearly function as webs in reality. This complexity is not uncommon in
biological systems which are characterized by links that change in response to both
external and internal environmental factors (Biggs et al. 2009). Although these streams
were comparable in biogeochemistry and other abiotic features, the stream communities
as a whole showed variation in trophic relationships and factors like diversity and
biomass that may have been missed without such an inclusive ecosystem level
experimental design (Fretwell 1987). The abundance of fishes in higher trophic levels in
these streams suggests tight coupling between trophic levels must be present in order to
58
support such communities and must be facilitated through median trophic levels. Indeed,
lower trophic level responses may be dependent upon food web configuration as
mediated by complexity within median trophic levels, whereby some systems (i.e.,
functional four-step chains) exhibit greater predation constraints and others (i.e.,
functional three-step chains) respond more freely to bottom-up influences.
59
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Appendix A. Presence/absence of algal taxa in each of the three streams. An “x”
indicates the taxon was present in that stream.
Algal Division
Bacillariophyta
Genus
Achnanthes-like
Mesotrophic Eutrophic Hypereutrophic
X
X
X
Bacillariophyta
Amphora
X
X
X
Bacillariophyta
Campylodiscus
X
X
X
Bacillariophyta
Cocconeis
X
X
X
Bacillariophyta
Cyclotella
X
X
X
Bacillariophyta
Cymatopleura
X
X
X
Bacillariophyta
Cymbelloid
X
X
X
Bacillariophyta
Diatoma
X
X
X
Bacillariophyta
Fragillaria
X
X
X
Bacillariophyta
Gomphonema
X
X
X
Bacillariophyta
Gyrosigma
X
X
X
Bacillariophyta
Melosira
X
X
X
Bacillariophyta
Meridion
X
Bacillariophyta
Naviculoid (small)
X
X
X
Bacillariophyta
Naviculoid (large)
X
X
X
Bacillariophyta
Nitzschia
X
X
X
Bacillariophyta
Rhoicosphenia
X
X
X
Bacillariophyta
Surrirella
X
X
X
Bacillariophyta
Synedra
X
X
X
Chlorophyta
Cladophora glomerata
X
X
X
Chlorophyta
Closterium (small)
X
X
X
Chlorophyta
Closterium (medium)
X
X
X
Chlorophyta
Closterium (large)
X
X
X
Chlorophyta
Cosmarium
X
X
X
Chlorophyta
Cylindrocapsa
X
X
Chlorophyta
Gloeocystis
X
Chlorophyta
Green balls
X
Chlorophyta
Mougeotia
Chlorophyta
Oedogonium (small)
Chlorophyta
Oedogonium (medium)
Chlorophyta
Pediastrum
Chlorophyta
Rhizoclonium
X
X
X
X
X
X
X
X
X
X
70
Appendix A. (continued) Presence/absence of algal taxa in each of the three streams. An
“x” indicates the taxon was present in that stream.
Algal Division
Chlorophyta
Genus
Scenedesmus
Chlorophyta
Schizomeris
Chlorophyta
Stigeoclonium
Chlorophyta
Mesotrophic Eutrophic Hypereutrophic
X
X
X
X
X
X
X
X
Ulothrix
X
X
X
Cyanobacteria
Aphanocapsa
X
X
X
Cyanobacteria
Blue-green balls
X
X
X
Cyanobacteria
Chroococcus
X
Cyanobacteria
Homeothrix
X
Cyanobacteria
Lyngbya
X
X
X
Cyanobacteria
Lyngbya (thin)
X
X
X
Cyanobacteria
Merismopedia
X
X
X
Cyanobacteria
Microcystis
X
Cyanobacteria
Oscillatoria
X
Cyanobacteria
Phormidium
X
X
X
Cyanobacteria
Spirulina
X
X
X
Cyanobacteria
Stauromatonema
X
Euglenophyta
Euglena
X
X
X
Chrysophyta
Synura
X
Rhodophyta
Batrachospermum
X
X
X
X
71
Appendix B. Algal biomass (mg DW/m2) in mesotrophic stream by taxa for treatment.
Values represent mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus
treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Bacillariophyta Achnanthes-like
Bacillariophyta Amphora
Bacillariophyta Campylodiscus
Bacillariophyta Cocconeis
Bacillariophyta Cyclotella
Bacillariophyta Cymatopleura
Bacillariophyta Cymbelloid
Bacillariophyta Diatoma
Bacillariophyta Fragillaria
Bacillariophyta Gomphonema
Bacillariophyta Gyrosigma
Bacillariophyta Melosira
Bacillariophyta Meridion
Bacillariophyta Naviculoid (small)
Bacillariophyta Naviculoid (large)
Bacillariophyta Nitzschia
Bacillariophyta Rhoicosphenia
Bacillariophyta Surrirella
Bacillariophyta Synedra
Control
N
P
N+P
887.09
(295.39)
4.45
(2.83)
4.70
(9.40)
1498.72
(317.47)
0.00
(0.00)
0.51
(0.60)
119.59
(67.24)
91.68
(25.35)
0.02
(0.04)
636.66
(230.87)
34.46
(21.79)
146.17
(45.17)
0.00
(0.00)
84.92
(28.92)
1513.82
(131.40)
1128.42
(610.76)
188.51
(80.29)
1.24
(2.48)
234.05
(120.23)
864.64
(249.20)
4.93
(3.11)
9.10
(10.53)
1620.26
(637.63)
10.49
(12.14)
0.27
(0.54)
168.96
(168.74)
166.89
(109.34)
0.02
(0.03)
787.71
(230.77)
32.17
(19.45)
119.42
(36.97)
0.00
(0.00)
152.50
(118.66)
1977.10
(559.65)
712.91
(490.40)
142.90
(104.91)
0.48
(0.96)
238.25
(138.64)
1134.37
(281.69)
6.73
(4.37)
14.93
(19.87)
1962.92
(308.47)
8.34
(16.68)
1.18
(1.46)
62.57
(44.36)
141.85
(62.34)
0.00
(0.00)
724.09
(90.56)
40.61
(16.24)
156.65
(57.52)
0.00
(0.00)
131.12
(75.90)
1498.96
(293.15)
991.39
(300.22)
150.65
(36.29)
56.97
(113.94)
340.75
(125.22)
1004.02
(227.06)
6.83
(2.74)
3.93
(7.87)
1323.43
(438.21)
22.17
(44.34)
0.81
(0.57)
65.47
(78.38)
139.18
(137.32)
1.80
(3.60)
671.42
(224.28)
41.25
(15.35)
172.30
(119.24)
0.32
(0.63)
120.73
(13.15)
1754.79
(374.62)
1258.05
(456.67)
102.01
(55.84)
0.00
(0.00)
417.47
(40.77)
Stream
Average
972.53
(261.44)
5.73
(3.17)
8.17
(12.31)
1601.33
(466.12)
10.25
(23.35)
0.69
(0.87)
104.15
(101.32)
134.90
(88.64)
0.46
(1.80)
704.97
(190.99)
37.12
(16.93)
148.63
(67.69)
0.08
(0.32)
122.32
(69.33)
1686.17
(390.44)
1022.69
(475.44)
146.02
(73.37)
14.67
(56.87)
307.63
(128.22)
72
Appendix B. (continued) Algal biomass (mg DW/m2) in mesotrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Control
N
P
N+P
Chlorophyta
Cladophora glomerata
Chlorophyta
Closterium (small)
Chlorophyta
Closterium (medium)
Chlorophyta
Closterium (large)
Chlorophyta
Cosmarium
Chlorophyta
Cylindrocapsa
Chlorophyta
Gloeocystis
Chlorophyta
Green balls
Chlorophyta
Mougeotia
Chlorophyta
Oedogonium (small)
Chlorophyta
Oedogonium (medium)
Chlorophyta
Pediastrum
Chlorophyta
Rhizoclonium
Chlorophyta
Scenedesmus
Chlorophyta
Schizomeris
Chlorophyta
Stigeoclonium
Chlorophyta
Ulothrix
Cyanobacteria
Aphanocapsa
Cyanobacteria
Blue-green balls
Cyanobacteria
Chroococcus
0.00
(0.00)
2.19
(4.39)
0.00
(0.00)
4.74
(3.66)
0.00
(0.00)
0.24
(0.10)
0.00
(0.00)
0.38
(0.76)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
7.37
(14.23)
0.00
(0.00)
168.01
(328.13)
0.00
(0.00)
25.44
(18.24)
0.00
(0.00)
15.48
(3.04)
94.86
(39.28)
0.00
(0.00)
0.50
(0.99)
3.71
(7.43)
51.59
(103.19)
3.66
(2.53)
0.00
(0.00)
0.38
(0.25)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.70
(0.30)
0.00
(0.00)
2.31
(1.60)
0.00
(0.00)
27.68
(32.43)
6.08
(12.15)
8.36
(11.67)
89.37
(31.93)
1.07
(2.14)
0.00
(0.00)
0.00
(0.00)
0.19
(0.39)
4.74
(1.76)
6.91
(13.82)
0.54
(0.38)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
19.33
(36.96)
0.00
(0.00)
5.30
(0.95)
0.00
(0.00)
28.77
(21.45)
0.00
(0.00)
4.78
(4.25)
97.16
(26.59)
0.00
(0.00)
0.00
(0.00)
9.99
(19.98)
0.24
(0.49)
8.39
(7.21)
12.96
(25.91)
0.90
(0.49)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
7.74
(15.48)
0.00
(0.00)
0.22
(0.32)
0.00
(0.00)
6.34
(2.12)
0.00
(0.00)
253.73
(465.71)
3.54
(7.09)
7.68
(9.17)
102.67
(34.45)
0.00
(0.00)
Stream
Average
0.12
(0.50)
3.97
(10.46)
13.01
(51.56)
5.38
(4.29)
4.97
(14.27)
0.51
(0.40)
0.00
(0.00)
0.10
(0.38)
0.00
(0.00)
1.94
(7.74)
0.00
(0.00)
6.91
(19.42)
0.00
(0.00)
45.49
(163.94)
0.00
(0.00)
83.90
(232.39)
2.40
(6.83)
9.08
(8.13)
96.02
(30.26)
0.27
(1.07)
73
Appendix B. (continued) Algal biomass (mg DW/m2) in mesotrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Cyanobacteria
Homeothrix
Cyanobacteria
Lyngbya
Cyanobacteria
Lyngbya (thin)
Cyanobacteria
Merismopedia
Cyanobacteria
Microcystis
Cyanobacteria
Oscillatoria
Cyanobacteria
Phormidium
Cyanobacteria
Spirulina
Cyanobacteria
Stauromatonema
Euglenophyta
Euglena
Chrysophyta
Synura
Rhodophyta
Batrachospermum
Control
N
P
N+P
0.51
(1.02)
0.00
(0.00)
43.06
(38.36)
0.32
(0.25)
55.49
(83.40)
0.00
(0.00)
39.09
(78.19)
0.00
(0.00)
0.23
(0.47)
0.10
(0.14)
0.61
(1.22)
0.00
(0.00)
0.98
(1.31)
131.10
(154.38)
19.97
(24.46)
0.61
(0.60)
70.72
(112.93)
0.00
(0.00)
0.00
(0.00)
0.00
(0.01)
1.32
(2.63)
0.09
(0.10)
0.00
(0.00)
0.00
(0.00)
1.04
(1.45)
0.04
(0.08)
26.25
(21.80)
0.31
(0.20)
34.96
(69.59)
0.00
(0.00)
5.00
(10.00)
0.00
(0.00)
0.00
(0.00)
0.05
(0.09)
0.00
(0.00)
0.00
(0.00)
0.97
(1.95)
0.00
(0.00)
39.17
(18.91)
0.22
(0.11)
48.27
(60.44)
9.44
(18.88)
311.17
(622.33)
0.00
(0.01)
0.00
(0.00)
0.04
(0.08)
0.00
(0.00)
0.00
(0.00)
Stream
Average
0.87
(1.33)
32.78
(90.57)
32.12
(25.97)
0.37
(0.34)
52.36
(76.28)
2.36
(9.44)
88.81
(310.68)
0.00
(0.01)
0.39
(1.32)
0.07
(0.10)
0.15
(0.61)
0.00
(0.00)
74
Appendix C. Algal biomass (mg DW/m2) in eutrophic stream by taxa for treatment.
Values represent mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus
treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Bacillariophyta Achnanthes-like
Bacillariophyta Amphora
Bacillariophyta Campylodiscus
Bacillariophyta Cocconeis
Bacillariophyta Cyclotella
Bacillariophyta Cymatopleura
Bacillariophyta Cymbelloid
Bacillariophyta Diatoma
Bacillariophyta Fragillaria
Bacillariophyta Gomphonema
Bacillariophyta Gyrosigma
Bacillariophyta Melosira
Bacillariophyta Meridion
Bacillariophyta Naviculoid (small)
Bacillariophyta Naviculoid (large)
Bacillariophyta Nitzschia
Bacillariophyta Rhoicosphenia
Bacillariophyta Surrirella
Bacillariophyta Synedra
Control
N
P
N+P
301.90
(122.03)
7.06
(3.25)
2.77
(5.54)
668.58
(208.13)
46.62
(26.25)
3.23
(3.25)
534.91
(174.70)
179.33
(75.35)
15.50
(29.94)
90.76
(74.72)
31.16
(9.06)
37.06
(30.44)
0.00
(0.00)
80.24
(50.63)
1008.23
(228.71)
1052.48
(857.63)
8.41
(10.33)
294.66
(254.02)
59.80
(53.08)
456.70
(165.03)
3.50
(1.69)
3.64
(7.28)
778.29
(208.98)
5.49
(10.99)
1.92
(0.78)
532.14
(164.23)
151.53
(126.90)
3.33
(3.89)
120.55
(60.29)
19.54
(8.59)
11.52
(7.75)
0.00
(0.00)
65.98
(22.91)
783.56
(209.39)
686.37
(355.99)
14.44
(12.64)
180.40
(250.73)
30.63
(12.72)
264.32
(60.30)
3.07
(1.20)
2.18
(4.37)
491.45
(96.19)
10.29
(12.20)
0.68
(0.57)
436.00
(165.37)
148.89
(50.70)
0.04
(0.06)
46.13
(32.85)
16.64
(11.08)
5.82
(6.36)
0.00
(0.00)
36.85
(24.32)
538.39
(257.00)
195.66
(221.56)
0.00
(0.00)
97.30
(111.37)
17.54
(19.07)
443.20
(308.40)
6.30
(4.06)
0.00
(0.00)
680.47
(340.14)
19.57
(14.41)
2.39
(0.76)
644.69
(51.70)
101.36
(96.81)
1.52
(2.97)
143.81
(29.98)
30.40
(8.88)
42.74
(33.40)
0.00
(0.00)
80.38
(33.22)
1291.39
(915.47)
1097.74
(745.03)
14.78
(22.23)
440.32
(586.65)
59.06
(16.88)
Stream
Average
366.53
(189.24)
4.98
(3.07)
2.15
(4.74)
654.70
(231.94)
20.49
(22.44)
2.05
(1.82)
536.93
(152.72)
145.28
(87.08)
5.10
(14.97)
100.31
(60.50)
24.43
(10.76)
24.28
(26.40)
0.00
(0.00)
65.86
(35.95)
905.39
(531.49)
758.06
(657.94)
9.41
(13.80)
253.17
(338.35)
41.76
(32.89)
75
Appendix C. (continued) Algal biomass (mg DW/m2) in eutrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Chlorophyta
Genus
Chlorophyta
Cladophora
glomerata
Closterium (small)
Chlorophyta
Closterium (medium)
Chlorophyta
Closterium (large)
Chlorophyta
Cosmarium
Chlorophyta
Cylindrocapsa
Chlorophyta
Gloeocystis
Chlorophyta
Green balls
Chlorophyta
Mougeotia
Chlorophyta
Oedogonium (small)
Chlorophyta
Chlorophyta
Oedogonium
(medium)
Pediastrum
Chlorophyta
Rhizoclonium
Chlorophyta
Scenedesmus
Chlorophyta
Schizomeris
Chlorophyta
Stigeoclonium
Chlorophyta
Ulothrix
Cyanobacteria
Aphanocapsa
Cyanobacteria
Blue-green balls
Control
N
P
N+P
0.58
(1.15)
13.67
(27.34)
32.68
(65.35)
1.71
(2.55)
6.56
(13.13)
0.22
(0.39)
0.00
(0.00)
0.00
(0.00)
1.53
(3.06)
0.00
(0.00)
5.77
(11.54)
0.00
(0.00)
0.00
(0.00)
0.60
(0.93)
10.05
(20.09)
0.00
(0.00)
0.00
(0.00)
0.23
(0.45)
59.76
(52.78)
0.00
(0.00)
0.00
(0.00)
0.13
(0.26)
1.56
(2.05)
8.64
(17.27)
0.12
(0.08)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.12
(3.53)
0.32
(0.22)
0.00
(0.00)
1.22
(2.43)
0.00
(0.00)
0.61
(0.95)
63.26
(7.53)
0.21
(0.42)
3.04
(6.08)
0.04
(0.09)
0.29
(0.58)
0.00
(0.00)
0.06
(0.10)
9.33
(18.67)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.68
(0.89)
0.07
(0.10)
0.00
(0.00)
10.76
(20.32)
0.00
(0.00)
0.90
(1.59)
38.52
(26.28)
0.00
(0.00)
0.00
(0.00)
62.47
(124.60)
1.46
(1.97)
7.51
(15.03)
0.12
(0.08)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.36
(0.30)
0.00
(0.00)
0.00
(0.00)
3.46
(6.93)
0.50
(0.59)
67.34
(43.50)
Stream
Average
0.20
(0.60)
4.18
(13.80)
23.83
(68.40)
1.26
(1.82)
5.68
(12.30)
0.13
(0.19)
2.33
(9.33)
0.00
(0.00)
0.38
(1.53)
0.00
(0.00)
1.44
(5.77)
0.00
(0.00)
0.70
(1.86)
0.34
(0.49)
2.51
(10.05)
2.99
(10.27)
0.87
(3.46)
0.56
(0.93)
57.22
(34.89)
76
Appendix C. (continued) Algal biomass (mg DW/m2) in eutrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Cyanobacteria
Chroococcus
Cyanobacteria
Homeothrix
Cyanobacteria
Lyngbya
Cyanobacteria
Lyngbya (thin)
Cyanobacteria
Merismopedia
Cyanobacteria
Microcystis
Cyanobacteria
Oscillatoria
Cyanobacteria
Phormidium
Cyanobacteria
Spirulina
Cyanobacteria
Stauromatonema
Euglenophyta
Euglena
Chrysophyta
Synura
Rhodophyta
Batrachospermum
Control
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.15
(0.21)
12.90
(17.32)
0.47
(0.81)
0.00
(0.00)
0.00
(0.00)
22.86
(45.71)
0.01
(0.01)
0.00
(0.00)
0.06
(0.07)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.27
(0.23)
9.20
(12.95)
0.09
(0.07)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.28
(0.57)
0.00
(0.00)
0.19
(0.09)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.09
(0.12)
4.99
(6.21)
0.14
(0.15)
0.00
(0.00)
0.00
(0.00)
0.29
(0.58)
0.00
(0.00)
0.00
(0.00)
0.10
(0.07)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.66
(0.45)
10.49
(13.22)
0.09
(0.07)
0.00
(0.00)
0.00
(0.00)
66.16
(132.32)
0.00
(0.01)
0.00
(0.00)
0.39
(0.09)
0.00
(0.00)
1.88
(3.77)
Stream
Average
0.00
(0.00)
0.00
(0.00)
0.29
(0.34)
9.39
(12.04)
0.20
(0.41)
0.00
(0.00)
0.00
(0.00)
22.33
(68.52)
0.07
(0.28)
0.00
(0.00)
0.19
(0.15)
0.00
(0.00)
0.47
(1.88)
77
Appendix D. Algal biomass (mg DW/m2) in hypereutrophic stream by taxa for treatment.
Values represent mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus
treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Bacillariophyta Achnanthes-like
Bacillariophyta Amphora
Bacillariophyta Campylodiscus
Bacillariophyta Cocconeis
Bacillariophyta Cyclotella
Bacillariophyta Cymatopleura
Bacillariophyta Cymbelloid
Bacillariophyta Diatoma
Bacillariophyta Fragillaria
Bacillariophyta Gomphonema
Bacillariophyta Gyrosigma
Bacillariophyta Melosira
Bacillariophyta Meridion
Bacillariophyta Naviculoid (small)
Bacillariophyta Naviculoid (large)
Bacillariophyta Nitzschia
Bacillariophyta Rhoicosphenia
Bacillariophyta Surrirella
Bacillariophyta Synedra
Control
N
P
N+P
412.82
(130.97)
1.55
(0.76)
0.00
(0.00)
360.38
(174.40)
3.50
(6.99)
0.22
(0.44)
350.13
(258.91)
177.38
(64.48)
4.61
(6.35)
279.03
(193.52)
23.90
(12.09)
42.89
(21.32)
0.00
(0.00)
52.14
(23.31)
543.75
(197.13)
263.12
(321.98)
30.05
(8.75)
101.18
(119.42)
14.35
(12.30)
397.88
(104.52)
0.86
(0.54)
21.16
(21.18)
375.92
(173.30)
22.82
(20.10)
0.75
(0.87)
291.72
(153.10)
133.84
(72.32)
0.00
(0.00)
346.28
(158.13)
20.25
(3.59)
47.96
(32.05)
0.00
(0.00)
50.89
(6.97)
514.47
(98.71)
508.54
(342.33)
10.81
(8.57)
29.67
(59.34)
19.71
(9.31)
470.26
(265.56)
1.52
(0.33)
0.00
(0.00)
524.58
(204.03)
3.00
(5.19)
1.50
(1.51)
367.84
(134.86)
167.33
(12.33)
3.09
(5.35)
407.02
(311.72)
28.80
(12.65)
36.76
(10.54)
4.00
(6.93)
44.95
(1.87)
548.75
(129.90)
365.27
(367.81)
10.48
(9.62)
0.00
(0.00)
19.25
(1.75)
456.62
(250.56)
1.81
(1.33)
37.33
(31.24)
518.94
(222.15)
15.38
(10.43)
1.96
(0.77)
466.49
(347.67)
116.71
(46.76)
0.00
(0.00)
440.75
(100.52)
33.13
(8.12)
59.66
(30.79)
0.20
(0.40)
55.63
(39.66)
408.51
(344.58)
610.15
(287.04)
15.85
(13.14)
67.06
(134.11)
25.32
(3.82)
Stream
Average
432.01
(174.55)
1.43
(0.85)
15.60
(23.89)
439.65
(189.14)
11.72
(14.10)
1.08
(1.07)
369.13
(228.93)
147.58
(56.26)
1.85
(4.14)
365.69
(183.46)
26.37
(9.89)
47.49
(24.69)
0.85
(3.09)
51.30
(21.88)
500.88
(204.39)
441.54
(322.58)
17.22
(12.30)
52.77
(95.61)
19.68
(8.47)
78
Appendix D. (continued) Algal biomass (mg DW/m2) in hypereutrophic stream by taxa
for treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Chlorophyta
Chlorophyta
Chlorophyta
Genus
Control
N
P
Cladophora
glomerata
Closterium (small)
0.00
(0.00)
4.73
(9.45)
0.11
(0.23)
0.00
(0.00)
0.00
(0.00)
0.09
(0.18)
31.85
(63.71)
0.00
(0.00)
0.00
(0.00)
1.61
(3.23)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.07
(0.13)
93.90
(108.74)
7.85
(6.79)
0.00
(0.00)
0.00
(0.00)
37.74
(18.11)
0.00
(0.00)
10.43
(14.89)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
9.07
(18.15)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
6.90
(13.80)
0.00
(0.00)
0.00
(0.00)
0.37
(0.69)
10.31
(20.62)
9.34
(15.36)
15.24
(20.64)
0.07
(0.14)
46.89
(12.69)
0.00
(0.00)
24.48
(42.40)
0.00
(0.00)
35.72
(61.87)
0.00
(0.00)
0.00
(0.00)
17.01
(29.46)
0.00
(0.00)
0.00
(0.00)
11.72
(10.84)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.14
(0.25)
50.33
(87.18)
7.78
(13.47)
1.62
(2.81)
0.00
(0.00)
35.71
(46.79)
Chlorophyta
Closterium
(medium)
Closterium (large)
Chlorophyta
Cosmarium
Chlorophyta
Cylindrocapsa
Chlorophyta
Gloeocystis
Chlorophyta
Green balls
Chlorophyta
Mougeotia
Chlorophyta
Chlorophyta
Oedogonium
(small)
Oedogonium
(medium)
Pediastrum
Chlorophyta
Rhizoclonium
Chlorophyta
Scenedesmus
Chlorophyta
Schizomeris
Chlorophyta
Stigeoclonium
Chlorophyta
Ulothrix
Cyanobacteria
Aphanocapsa
Cyanobacteria
Blue-green balls
Chlorophyta
Stream
Average
0.00
0.00
(0.00)
(0.00)
3.04
9.75
(6.08)
(19.95)
0.00
0.03
(0.00)
(0.12)
1.05
7.42
(1.21)
(27.60)
8.77
2.34
(10.25)
(6.22)
0.02
0.03
(0.04)
(0.09)
19.78
19.59
(39.57) (38.42)
0.00
0.00
(0.00)
(0.00)
0.00
0.00
(0.00)
(0.00)
0.00
2.77
(0.00)
(6.40)
0.00
1.84
(0.00)
(7.12)
0.00
0.00
(0.00)
(0.00)
0.67
0.18
(1.34)
(0.69)
0.00
0.14
(0.00)
(0.37)
249.43
104.37
(498.86) (257.36)
18.03
10.95
(21.68) (14.39)
11.06
7.34
(22.12) (15.54)
0.00
0.02
(0.00)
(0.07)
60.12
45.74
(20.58) (24.65)
N+P
79
Appendix D. (continued) Algal biomass (mg DW/m2) in hypereutrophic stream by taxa
for treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Control
N
P
N+P
Cyanobacteria
Chroococcus
Cyanobacteria
Homeothrix
Cyanobacteria
Lyngbya
Cyanobacteria
Lyngbya (thin)
Cyanobacteria
Merismopedia
Cyanobacteria
Microcystis
Cyanobacteria
Oscillatoria
Cyanobacteria
Phormidium
Cyanobacteria
Spirulina
Cyanobacteria
Stauromatonema
Euglenophyta
Euglena
Chrysophyta
Synura
Rhodophyta
Batrachospermum
0.00
(0.00)
0.00
(0.00)
6.95
(13.51)
3.50
(4.15)
0.05
(0.10)
0.00
(0.00)
0.00
(0.00)
21.80
(43.61)
0.00
(0.00)
0.00
(0.00)
0.11
(0.08)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.63
(1.26)
6.60
(12.66)
17.85
(13.95)
0.44
(0.79)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.14
(0.08)
0.00
(0.00)
2.55
(5.09)
0.00
(0.00)
0.00
(0.00)
17.39
(30.13)
9.99
(17.30)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.01)
0.00
(0.00)
0.18
(0.31)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.46
(0.43)
8.11
(9.42)
0.13
(0.19)
0.02
(0.05)
0.00
(0.00)
67.96
(135.93)
0.00
(0.00)
0.00
(0.00)
0.16
(0.24)
0.00
(0.00)
0.00
(0.00)
Stream
Average
0.00
(0.00)
0.17
(0.65)
7.22
(15.44)
9.85
(11.74)
0.17
(0.42)
0.01
(0.02)
0.00
(0.00)
23.94
(72.17)
0.00
(0.00)
0.00
(0.00)
0.15
(0.17)
0.00
(0.00)
0.68
(2.63)
80
Appendix E. Algal density (cells/cm2) in mesotrophic stream by taxa for treatment.
Values represent mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus
treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Bacillariophyta Achnanthes-like
Bacillariophyta Amphora
Bacillariophyta Campylodiscus
Bacillariophyta Cocconeis
Bacillariophyta Cyclotella
Bacillariophyta Cymatopleura
Bacillariophyta Cymbelloid
Bacillariophyta Diatoma
Bacillariophyta Fragillaria
Bacillariophyta Gomphonema
Bacillariophyta Gyrosigma
Bacillariophyta Melosira
Bacillariophyta Meridion
Bacillariophyta Naviculoid (small)
Bacillariophyta Naviculoid (large)
Bacillariophyta Nitzschia
Bacillariophyta Rhoicosphenia
Bacillariophyta Surrirella
Bacillariophyta Synedra
Control
N
P
N+P
6.33x105
(2.11x105)
4.21x102
(2.68x102)
7.26x100
(1.45x101)
1.05x105
(2.23x104)
0.00x100
(0.00x100)
1.28x101
(1.51x101)
6.30x103
(3.54x103)
5.73x103
(1.58x103)
7.31x100
(1.46x101)
1.30x105
(4.72x104)
4.96x102
(3.14x102)
5.09x103
(1.57x103)
0.00x100
(0.00x100)
9.36x104
(3.19x104)
1.79x105
(1.56x104)
1.48x105
(8.03x104)
3.77x104
(1.61x104)
1.45x101
(2.90x101)
1.16x104
(5.96x103)
6.17x105
(1.78x105)
4.67x102
(2.95x102)
1.41x101
(1.63x101)
1.14x105
(4.48x104)
1.46x103
(1.69x103)
6.75x100
(1.35x101)
8.90x103
(8.89x103)
1.04x104
(6.83x103)
6.75x100
(1.35x101)
1.61x105
(4.72x104)
4.64x102
(2.80x102)
4.16x103
(1.29x103)
0.00x100
(0.00x100)
1.68x105
(1.31x105)
2.34x105
(6.63x104)
9.38x104
(6.45x104)
2.86x104
(2.10x104)
5.63x100
(1.13x101)
1.18x104
(6.87x103)
8.09x105
(2.01x105)
6.38x102
(4.14x102)
2.31x101
(3.07x101)
1.38x105
(2.17x104)
1.16x103
(2.32x103)
2.96x101
(3.66x101)
3.30x103
(2.34x103)
8.87x103
(3.90x103)
0.00x100
(0.00x100)
1.48x105
(1.85x104)
5.85x102
(2.34x102)
5.45x103
(2.00x103)
0.00x100
(0.00x100)
1.45x105
(8.37x104)
1.78x105
(3.47x104)
1.30x105
(3.95x104)
3.01x104
(7.26x103)
6.68x102
(1.34x103)
1.69x104
(6.21x103)
7.16x105
(1.62x105)
6.47x102
(2.59x102)
6.08x100
(1.22x101)
9.30x104
(3.08x104)
3.09x103
(6.17x103)
2.03x101
(1.42x101)
3.45x103
(4.13x103)
8.70x103
(8.58x103)
7.30x102
(1.46x103)
1.37x105
(4.59x104)
5.94x102
(2.21x102)
6.00x103
(4.15x103)
5.00x101
(9.99x101)
1.33x105
(1.45x104)
2.08x105
(4.44x104)
1.65x105
(6.01x104)
2.04x104
(1.12x104)
0.00x100
(0.00x100)
2.07x104
(2.02x103)
Stream
Average
6.94x105
(1.86x105)
5.43x102
(3.01x102)
1.26x101
(1.90x101)
1.13x105
(3.27x104)
1.43x103
(3.25x103)
1.74x101
(2.16x101)
5.49x103
(5.34x103)
8.43x103
(5.54x103)
1.86x102
(7.29x102)
1.44x105
(3.91x104)
5.35x102
(2.44x102)
5.17x103
(2.36x103)
1.25x101
(5.00x101)
1.35x105
(7.64x104)
2.00x105
(4.62x104)
1.35x105
6.25x104)
2.92x104
(1.47x104)
1.72x102
(6.66x102)
1.52x104
(6.36x103)
81
Appendix E. (continued) Algal density (cells/cm2) in mesotrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Chlorophyta
Chlorophyta
Chlorophyta
Genus
Control
N
P
N+P
Cladophora
glomerata
Closterium (small)
7.31x100
(1.46x101)
0.00x100
(0.00x100)
5.54x101
(4.27x101)
0.00x100
(0.00x100)
4.47x101
(1.90x101)
0.00x100
(0.00x100)
1.32x102
(2.63x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
7.37x103
(1.42x104)
0.00x100
(0.00x100)
1.68x105
(3.28x105)
0.00x100
(0.00x100)
8.48x102
(6.08x102)
0.00x100
(0.00x100)
8.50x103
(1.67x103)
6.78x105
(2.81x105)
1.24x101
(2.48x101)
2.50x103
(5.01x103)
4.28x101
(2.95x101)
0.00x100
(0.00x100)
7.03x101
(4.59x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
7.65x102
(1.53x103)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
7.01x102
(2.99x102)
0.00x100
(0.00x100)
2.31x103
(1.60x103)
0.00x100
(0.00x100)
9.23x102
(1.08x103)
2.03x102
(4.05x102)
4.59x103
(6.41x103)
6.38x105
(2.28x105)
0.00x100
(0.00x100)
9.39x100
(1.88x101)
5.54x101
(2.05x101)
9.00x100
(1.80x101)
1.01x102
(7.12x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.93x104
(3.70x104)
0.00x100
(0.00x100)
5.30x103
(9.54x102)
0.00x100
(0.00x100)
9.59x102
(7.15x102)
0.00x100
(0.00x100)
2.62x103
(2.33x103)
6.94x105
(1.90x105)
3.33x101
(6.66x101)
1.18x101
(2.36x101)
9.80x101
(8.42x101)
1.69x101
(3.38x101)
1.68x102
(9.21x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.58x102
(5.16x102)
0.00x100
(0.00x100)
2.15x102
(3.22x102)
0.00x100
(0.00x100)
6.34x103
(2.12x103)
0.00x100
(0.00x100)
8.46x103
(1.55x104)
1.18x102
(2.36x102)
4.22x103
(5.03x103)
7.33x105
(2.46x105)
Chlorophyta
Closterium
(medium)
Closterium (large)
Chlorophyta
Cosmarium
Chlorophyta
Cylindrocapsa
Chlorophyta
Gloeocystis
Chlorophyta
Green balls
Chlorophyta
Mougeotia
Chlorophyta
Chlorophyta
Oedogonium
(small)
Oedogonium
(medium)
Pediastrum
Chlorophyta
Rhizoclonium
Chlorophyta
Scenedesmus
Chlorophyta
Schizomeris
Chlorophyta
Stigeoclonium
Chlorophyta
Ulothrix
Cyanobacteria
Aphanocapsa
Cyanobacteria
Blue-green balls
Chlorophyta
Stream
Average
1.32x101
(3.49x101)
6.31x102
(2.50x103)
6.29x101
(5.01x101)
6.47x100
(1.86x101)
9.59x101
(7.39x101)
0.00x100
(0.00x100)
3.29x101
(1.32x102)
1.91x102
(7.65x102)
0.00x100
(0.00x100)
6.45x101
(2.58x102)
0.00x100
(0.00x100)
6.91x103
(1.94x104)
0.00x100
(0.00x100)
4.55x104
(1.64x105)
0.00x100
(0.00x100)
2.80x103
(7.75x103)
8.02x101
(2.28x102)
4.98x103
(4.46x103)
6.86x105
(2.16x105)
82
Appendix E. (continued) Algal density (cells/cm2) in mesotrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Cyanobacteria
Chroococcus
Cyanobacteria
Homeothrix
Cyanobacteria
Lyngbya
Cyanobacteria
Lyngbya (thin)
Cyanobacteria
Merismopedia
Cyanobacteria
Microcystis
Cyanobacteria
Oscillatoria
Cyanobacteria
Phormidium
Cyanobacteria
Spirulina
Cyanobacteria
Stauromatonema
Euglenophyta
Euglena
Chrysophyta
Synura
Rhodophyta
Batrachospermum
Control
N
P
N+P
0.00x100
(0.00x100)
5.08x102
(1.02x103)
0.00x100
(0.00x100)
3.08x105
(2.74x105)
3.22x103
(2.46x103)
1.11x105
(1.67x105)
0.00x100
(0.00x100)
3.98x104
(7.96x104)
0.00x100
(0.00x100)
2.34x102
(4.68x102)
1.55x101
(2.09x101)
5.85x101
(1.17x102)
0.00x100
(0.00x100)
1.07x103
(2.14x103)
9.78x102
(1.31x103)
9.36x105
(1.10x106)
1.43x105
(1.75x105)
6.14x103
(5.98x103)
1.41x105
(2.26x105)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.13x101
(2.25x101)
1.32x103
(2.63x103)
1.29x101
(1.50x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.04x103
(1.45x103)
2.96x102
(5.91x102)
1.88x105
(1.56x105)
3.08x103
(1.97x103)
6.99x104
(1.39x105)
0.00x100
(0.00x100)
5.09x103
(1.02x104)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
6.86x100
(1.37x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
9.73x102
(1.95x103)
0.00x100
(0.00x100)
2.80x105
(1.35x105)
2.22x103
(1.12x103)
9.65x104
(1.21x105)
4.50x104
(8.99x104)
3.17x105
(6.34x105)
8.33x100
(1.67x101)
0.00x100
(0.00x100)
6.08x100
(1.22x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
Stream
Average
2.67x102
(1.07x103)
8.75x102
(1.33x103)
2.34x105
(6.47x105)
2.29x105
(1.85x105)
3.67x103
(3.42x103)
1.05x105
(1.53x105)
1.12x104
(4.50x104)
9.04x104
(3.16x105)
4.89x100
(1.35x101)
3.88x102
(1.32x103)
1.04x101
(1.47x101)
1.46x101
(5.85x101)
0.00x100
(0.00x100)
83
Appendix F. Algal density (cells/cm2) in eutrophic stream by taxa for treatment. Values
represent mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments,
N+P=dual nutrient treatments.
Algal Division
Genus
Bacillariophyta Achnanthes-like
Bacillariophyta Amphora
Bacillariophyta Campylodiscus
Bacillariophyta Cocconeis
Bacillariophyta Cyclotella
Bacillariophyta Cymatopleura
Bacillariophyta Cymbelloid
Bacillariophyta Diatoma
Bacillariophyta Fragillaria
Bacillariophyta Gomphonema
Bacillariophyta Gyrosigma
Bacillariophyta Melosira
Bacillariophyta Meridion
Bacillariophyta Naviculoid (small)
Bacillariophyta Naviculoid (large)
Bacillariophyta Nitzschia
Bacillariophyta Rhoicosphenia
Bacillariophyta Surrirella
Bacillariophyta Synedra
Control
N
P
N+P
2.15x105
(8.70x104)
6.68x102
(3.08x102)
4.28x100
(8.55x100)
4.70x104
(1.46x104)
6.49x103
(3.66x103)
8.07x101
(8.11x101)
2.82x104
(9.20x103)
1.12x104
(4.71x103)
6.28x103
(1.21x104)
1.86x104
(1.53x104)
4.49x102
(1.31x102)
1.29x103
(1.06x103)
0.00x100
(0.00x100)
8.85x104
(5.58x104)
1.19x105
(2.71x104)
1.38x105
(1.13x105)
1.68x103
(2.07x103)
3.45x103
(2.98x103)
2.96x103
(2.63x103)
3.26x105
(1.18x105)
3.32x102
(1.60x102)
5.63x100
(1.13x101)
5.47x104
(1.47x104)
7.65x102
(1.53x103)
4.80x101
(1.94x101)
2.80x104
(8.65x103)
9.47x103
(7.93x103)
1.35x103
(1.58x103)
2.47x104
(1.23x104)
2.81x102
(1.24x102)
4.01x102
(2.70x102)
0.00x100
(0.00x100)
7.27x104
(2.53x104)
9.28x104
(2.48x104)
9.03x104
(4.68x104)
2.89x103
(2.53x103)
2.11x103
(2.94x103)
1.52x103
(6.31x102)
1.89x105
(4.30x104)
2.91x102
(1.14x102)
3.38x100
(6.75x100)
3.45x104
(6.76x103)
1.43x103
(1.70x103)
1.69x101
(1.44x101)
2.30x104
(8.71x103)
9.31x103
(3.17x103)
1.78x101
(2.45x101)
9.44x103
(6.72x103)
2.40x102
(1.60x102)
2.03x102
(2.21x102)
0.00x100
(0.00x100)
4.06x104
(2.68x104)
6.38x104
(3.04x104)
2.57x104
(2.91x104)
0.00x100
(0.00x100)
1.14x103
(1.31x103)
8.70x102
(9.45x102)
3.16x105
(2.20x105)
5.97x102
(3.84x102)
0.00x100
(0.00x100)
4.78x104
(2.39x104)
2.73x103
(2.01x103)
5.97x101
(1.91x101)
3.40x104
(2.72x103)
6.33x103
(6.05x103)
6.14x102
(1.20x103)
2.94x104
(6.14x103)
4.38x102
(1.28x102)
1.49x103
(1.16x103)
0.00x100
(0.00x100)
8.86x104
(3.66x104)
1.53x105
(1.08x105)
1.44x105
(9.80x104)
2.96x103
(4.45x103)
5.16x103
(6.87x103)
2.93x103
(8.37x102)
Stream
Average
2.61x105
(1.35x105)
4.72x102
(2.91x102)
3.32x100
(7.32x100)
4.60x104
(1.63x104)
2.85x103
(3.12x103)
5.13x101
(4.55x101)
2.83x104
(8.05x103)
9.08x103
(5.44x103)
2.07x103
(6.07x103)
2.05x104
(1.24x104)
3.52x102
(1.55x102)
8.45x102
(9.19x102)
0.00x100
(0.00x100)
7.26x104
(3.96x104)
1.07x105
(6.30x104)
9.97x104
(8.65x104)
1.88x103
(2.76x103)
2.97x103
(3.96x103)
2.07x103
(1.63x103)
84
Appendix F. (continued) Algal density (cells/cm2) in eutrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Chlorophyta
Cladophora
glomerata
Closterium (small)
Chlorophyta
Chlorophyta
Chlorophyta
Closterium
(medium)
Closterium (large)
Chlorophyta
Cosmarium
Chlorophyta
Cylindrocapsa
Chlorophyta
Gloeocystis
Chlorophyta
Green balls
Chlorophyta
Mougeotia
Chlorophyta
Chlorophyta
Oedogonium
(small)
Oedogonium
(medium)
Pediastrum
Chlorophyta
Rhizoclonium
Chlorophyta
Scenedesmus
Chlorophyta
Schizomeris
Chlorophyta
Stigeoclonium
Chlorophyta
Ulothrix
Cyanobacteria
Aphanocapsa
Cyanobacteria
Blue-green balls
Chlorophyta
Control
N
P
N+P
4.56x101
(9.11x101)
1.59x103
(3.17x103)
2.00x101
(2.98x101)
8.55x100
(1.71x101)
4.11x101
(7.20x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
8.85x102
(1.77x103)
2.36x101
(4.73x101)
0.00x100
(0.00x100)
9.62x101
(1.92x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
6.03x102
(9.26x102)
4.28x101
(8.55x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.24x102
(2.48x102)
4.27x105
(3.77x105)
0.00x100
(0.00x100)
6.19x100
(1.24x101)
1.82x101
(2.39x101)
1.13x101
(2.25x101)
2.15x101
(1.52x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.69x102
(2.81x102)
3.18x102
(2.18x102)
0.00x100
(0.00x100)
4.05x101
(8.10x101)
0.00x100
(0.00x100)
3.36x102
(5.21x102)
4.52x105
(5.38x104)
1.01x101
(2.03x101)
2.17x100
(4.33x100)
3.38x100
(6.75x100)
0.00x100
(0.00x100)
1.18x101
(1.84x101)
2.03x102
(4.05x102)
0.00x100
(0.00x100)
3.21x102
(6.42x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
5.45x101
(7.07x101)
7.05x101
(1.00x102)
0.00x100
(0.00x100)
3.59x102
(6.77x102)
0.00x100
(0.00x100)
4.95x102
(8.74x102)
2.75x105
(1.88x105)
0.00x100
(0.00x100)
3.03x103
(6.05x103)
1.70x101
(2.30x101)
9.79x100
(1.96x101)
2.18x101
(1.58x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
3.60x102
(3.03x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.15x102
(2.31x102)
2.73x102
(3.25x102)
4.81x105
(3.11x105)
Stream
Average
1.39x101
(4.60x101)
1.16x103
(3.32x103)
1.47x101
(2.13x101)
7.40x100
(1.60x101)
2.40x101
(3.63x101)
5.06x101
(2.03x102)
0.00x100
(0.00x100)
3.01x102
(9.21x102)
5.91x100
(2.36x101)
0.00x100
(0.00x100)
2.40x101
(9.62x101)
0.00x100
(0.00x100)
5.58x101
(1.48x102)
3.38x102
(4.89x102)
1.07x101
(4.28x101)
9.98x101
(3.42x102)
2.89x101
(1.15x102)
3.07x102
(5.09x102)
4.09x105
(2.49x105)
85
Appendix F. (continued) Algal density (cells/cm2) in eutrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Cyanobacteria
Chroococcus
Cyanobacteria
Homeothrix
Cyanobacteria
Lyngbya
Cyanobacteria
Lyngbya (thin)
Cyanobacteria
Merismopedia
Cyanobacteria
Microcystis
Cyanobacteria
Oscillatoria
Cyanobacteria
Phormidium
Cyanobacteria
Spirulina
Cyanobacteria
Stauromatonema
Euglenophyta
Euglena
Chrysophyta
Synura
Rhodophyta
Batrachospermum
Control
N
P
N+P
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.07x103
(1.53x103)
9.21x104
(1.24x105)
4.73x103
(8.07x103)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.33x104
(4.65x104)
1.6x101
(2.33x101)
0.00x100
(0.00x100)
9.00x100
(1.06x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.94x103
(1.63x103)
6.57x104
(9.25x104)
8.96x102
(7.16x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
7.65x102
(1.53x103)
0.00x100
(0.00x100)
2.88x101
(1.41x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
6.65x102
(8.49x102)
3.57x104
(4.44x104)
1.36x103
(1.50x103)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.97x102
(5.95x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.47x101
(1.11x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
4.70x103
(3.18x103)
7.49x104
(9.44x104)
8.96x102
(7.03x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
6.74x104
(1.35x105)
9.51x100
(1.90x101)
0.00x100
(0.00x100)
5.82x101
(1.27x101)
0.00x100
(0.00x100)
3.04x101
(6.08x101)
Stream
Average
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.09x103
(2.41x103)
6.71x104
(8.60x104)
1.97x103
(4.05x103)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.27x104
(6.98x104)
1.98x102
(7.63x102)
0.00x100
(0.00x100)
2.77x101
(2.25x101)
0.00x100
(0.00x100)
7.59x100
(3.04x101)
86
Appendix G. Algal density (cells/cm2) in hypereutrophic stream by taxa for treatment.
Values represent mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus
treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Bacillariophyta Achnantheslike
Bacillariophyta Amphora
Bacillariophyta Campylodiscus
Bacillariophyta Cocconeis
Bacillariophyta Cyclotella
Bacillariophyta Cymatopleura
Bacillariophyta Cymbelloid
Bacillariophyta Diatoma
Bacillariophyta Fragillaria
Bacillariophyta Gomphonema
Bacillariophyta Gyrosigma
Bacillariophyta Melosira
Bacillariophyta Meridion
Bacillariophyta Naviculoid
(small)
Bacillariophyta Naviculoid
(large)
Bacillariophyta Nitzschia
Bacillariophyta Rhoicosphenia
Bacillariophyta Surrirella
Bacillariophyta Synedra
Control
N
P
N+P
2.94x105
(9.34x104)
1.47x102
(7.21x101)
0.00x100
(0.00x100)
2.53x104
(1.23x104)
4.87x102
(9.74x102)
5.51x100
(1.10x101)
1.84x104
(1.36x104)
1.11x104
(4.03x103)
1.87x103
(2.57x103)
5.71x104
(3.96x104)
3.44x102
(1.74x102)
1.49x103
(7.42x102)
0.00x100
(0.00x100)
5.75x104
(2.57x104)
6.44x104
(2.34x104)
3.46x104
(4.23x104)
6.01x103
(1.75x103)
1.19x103
(1.40x103)
7.11x102
(6.10x102)
2.84x105
(7.46x104)
8.16x101
(5.09x101)
3.27x101
(3.27x101)
2.64x104
(1.22x104)
3.18x103
(2.80x103)
1.87x101
(2.16x101)
1.54x104
(8.07x103)
8.37x103
(4.52x103)
0.00x100
(0.00x100)
7.09x104
(3.24x104)
2.92x102
(5.18x101)
1.67x103
(1.12x103)
0.00x100
(0.00x100)
5.61x104
(7.68x103)
6.09x104
(1.17x104)
6.69x104
(4.50x104)
2.16x103
(1.71x103)
3.48x102
(6.95x102)
9.77x102
(4.61x102)
3.35x105
(1.89x105)
1.44x102
(3.17x101)
0.00x100
(0.00x100)
3.69x104
(1.43x104)
4.17x102
(7.23x102)
3.75x101
(3.77x101)
1.94x104
(7.10x103)
1.05x104
(7.71x102)
1.25x103
(2.17x103)
8.33x104
(6.38x104)
4.15x102
(1.82x102)
1.28x103
(3.67x102)
6.30x102
(1.09x103)
4.96x104
(2.06x103)
6.50x104
(1.54x104)
4.80x104
(4.84x104)
2.10x103
(1.92x103)
0.00x100
(0.00x100)
9.54x102
(8.68x101)
3.26x105
(1.79x105)
1.71x102
(1.26x102)
5.77x101
(4.83x101)
3.65x104
(1.56x104)
2.14x103
(1.45x103)
4.89x101
(1.94x101)
2.46x104
(1.83x104)
7.29x103
(2.92x103)
0.00x100
(0.00x100)
9.02x104
(2.06x104)
4.77x102
(1.17x102)
2.08x103
(1.07x103)
3.18x101
(6.36x101)
6.13x104
(4.37x104)
4.84x104
(4.08x104)
8.03x104
(3.78x104)
3.17x103
(2.63x103)
7.86x102
(1.57x103)
1.26x103
(1.89x102)
Stream
Average
3.08x105
(1.25x105)
1.35x102
(8.02x101)
2.41x101
(3.69x101)
3.09x104
(1.33x104)
1.63x103
(1.96x103)
2.70x101
(2.69x101)
1.94x104
(1.21x104)
9.22x103
(3.52x103)
7.48x102
(1.68x103)
7.48x104
(3.75x104)
3.80x102
(1.42x102)
1.65x103
(8.59x102)
1.35x102
(4.87x102)
5.66x104
(2.41x104)
5.93x104
(2.42x104)
5.81x104
(4.24x104)
3.44x103
(2.46x103)
6.18x102
(1.12x103)
9.76x102
(4.20x102)
87
Appendix G. (continued) Algal density (cells/cm2) in hypereutrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Chlorophyta
Genus
Chlorophyta
Cladophora
glomerata
Closterium
(small)
Closterium
(medium)
Closterium
(large)
Cosmarium
Chlorophyta
Cylindrocapsa
Chlorophyta
Gloeocystis
Chlorophyta
Green balls
Chlorophyta
Mougeotia
Chlorophyta
Chlorophyta
Oedogonium
(small)
Oedogonium
(medium)
Pediastrum
Chlorophyta
Rhizoclonium
Chlorophyta
Scenedesmus
Chlorophyta
Schizomeris
Chlorophyta
Stigeoclonium
Chlorophyta
Ulothrix
Chlorophyta
Chlorophyta
Chlorophyta
Chlorophyta
Cyanobacteria Aphanocapsa
Cyanobacteria Blue-green
balls
Control
N
P
N+P
1.58x101
(3.15x101)
5.51x100
(1.10x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.65x101
(3.31x101)
6.91x102
(1.38x103)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
5.38x101
(1.08x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
6.62x101
(1.32x102)
4.00x102
(4.63x102)
2.62x102
(2.26x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.70x105
(1.29x105)
3.48x101
(4.96x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.97x102
(3.94x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.15x102
(2.30x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
3.69x102
(6.92x102)
4.39x101
(8.78x101)
3.11x102
(5.12x102)
5.08x102
(6.88x102)
3.95x101
(7.90x101)
3.35x105
(9.06x104)
8.16x101
(1.41x102)
0.00x100
(0.00x100)
4.17x102
(7.23x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
3.69x102
(6.39x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
3.91x102
(3.61x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.44x102
(2.49x102)
2.14x102
(3.71x102)
2.59x102
(4.49x102)
5.40x101
(9.35x101)
0.00x100
(0.00x100)
2.55x105
(3.34x105)
1.01x101
(2.03x101)
0.00x100
(0.00x100)
1.23x101
(1.42x101)
1.14x101
(1.34x101)
3.94x100
(7.88x100)
4.29x102
(8.58x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
5.32x101
(1.06x102)
0.00x100
(0.00x100)
1.06x103
(2.12x103)
6.01x102
(7.23x102)
3.69x102
(7.37x102)
0.00x100
(0.00x100)
4.29x105
(1.47x105)
Stream
Average
3.25x101
(6.65x101)
1.47x100
(5.69x100)
8.67x101
(3.22x102)
3.05x100
(8.10x100)
5.46x100
(1.73x101)
4.25x102
(8.33x102)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
9.25x101
(2.13x102)
3.07x101
(1.19x102)
0.00x100
(0.00x100)
1.42x101
(5.49x101)
1.45x102
(3.71x102)
4.44x102
(1.10x103)
3.65x102
(4.80x102)
2.45x102
(5.18x102)
1.05x101
(4.08x101)
3.27x105
(1.76x105)
88
Appendix G. (continued) Algal density (cells/cm2) in hypereutrophic stream by taxa for
treatment. Values represent mean ±1 standard deviation; N=nitrogen treatments,
P=phosphorus treatments, N+P=dual nutrient treatments.
Algal Division
Genus
Cyanobacteria Chroococcus
Cyanobacteria Homeothrix
Cyanobacteria Lyngbya
Cyanobacteria Lyngbya (thin)
Cyanobacteria Merismopedia
Cyanobacteria Microcystis
Cyanobacteria Oscillatoria
Cyanobacteria Phormidium
Cyanobacteria Spirulina
Cyanobacteria Stauromatonema
Euglenophyta
Euglena
Chrysophyta
Synura
Rhodophyta
Batrachospermum
Control
N
P
N+P
0.00x100
(0.00x100)
0.00x100
(0.00x100)
4.97x104
(9.65x104)
2.50x104
(2.96x104)
5.20x102
(1.04x103)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.22x104
(4.44x104)
3.94x100
(7.88x100)
0.00x100
(0.00x100)
1.70x101
(1.17x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
6.29x102
(1.26x103)
4.71x104
(9.04x104)
1.27x105
(9.96x104)
4.43x103
(7.89x103)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
9.06x100
(1.05x101)
0.00x100
(0.00x100)
2.12x101
(1.26x101)
0.00x100
(0.00x100)
4.11x101
(8.21x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
1.24x105
(2.15x105)
7.13x104
(1.24x105)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
9.00x100
(1.56x101)
0.00x100
(0.00x100)
2.70x101
(4.68x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
3.30x103
(3.06x103)
5.80x104
(6.73x104)
1.30x103
(1.92x103)
4.73x101
(9.45x101)
0.00x100
(0.00x100)
6.92x104
(1.38x105)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
2.41x101
(3.60x101)
0.00x100
(0.00x100)
0.00x100
(0.00x100)
Stream
Average
0.00x100
(0.00x100)
1.68x102
(6.49x102)
5.15x104
(1.10x105)
7.04x104
(8.39x104)
1.67x103
(4.19x103)
1.26x101
(4.88x101)
0.00x100
(0.00x100)
2.44x104
(7.35x104)
5.27x100
(9.35x100)
0.00x100
(0.00x100)
2.20x101
(2.58x101)
0.00x100
(0.00x100)
1.10x101
(4.24x101)
89
Appendix H. Presence/absence of macroinvertebrate taxa in each of the three streams.
An “x” indicates the taxon was present in that stream. A * indicates the organism was
terrestrial and excluded from further analysis. Under family, (L)=larva, (P)=pupa,
(A)=represents adult. Under genus, S.F.=subfamily, T.=tribe, MT=mesotrophic stream,
ET=eutrophic stream, HT=hypereutrophic stream.
Order
Family
Acariformes
Hydrachnidia
Amphipoda
Gammaridae
Genus
Gammarus
Guild
MT
ET
HT
Parasite
X
X
X
Scraper
X
*Araneae
X
X
Coleoptera
Elmidae (A)
Coleoptera
Elmidae (L)
Stenelmis
Coleoptera (L)
Scraper
X
Collector-gatherer
X
Collector-gatherer
*Coleoptera (A)
terr.
Collembola
Poduridae
X
X
X
X
X
Podura aquatica
Cyclopoida
Collector-gatherer
X
Collector-filterer
Other
X
X
Diptera
Chironomidae (A)
Diptera
Chironomidae (L)
Clinotanypus
Predator
Diptera
Chironomidae (L)
Nilotanypus
Predator
Diptera
Chironomidae (L)
Psectrotanypus
Predator
Diptera
Chironomidae (L)
Brillia
Collector-gatherer
Diptera
Chironomidae (L)
Cricotopus
Collector-gatherer
Diptera
Chironomidae (L)
Parametriocnemus
Collector-gatherer
Diptera
Chironomidae (L)
Dicrotendipes
Collector-gatherer
Diptera
Chironomidae (L)
Glyptotendipes
Collector-gatherer
X
Diptera
Chironomidae (L)
Polypedilum
Collector-gatherer
X
Diptera
Chironomidae (L)
Microsectra
Collector-gatherer
X
X
X
Diptera
Chironomidae (L)
Paratanytarsus
Collector-gatherer
X
X
X
Diptera
Chironomidae (L)
Rheotanytarsus
Collector-filterer
X
X
Diptera
Chironomidae (L)
Tanytarsus
Collector-gatherer
X
X
Diptera
Chironomidae (P)
Non-feeding
X
X
X
Diptera
Empididae
Hemerodromia
Predator
X
X
X
Diptera
Empididae
Metachela/Chelifera Predator
Diptera
Simuliidae (A)
Diptera
Simuliidae (L)
Diptera
Simuliidae (L)
Collector-filterer
Diptera
Simuliidae (P)
Non-feeding
Other
Prosimulium
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Collector-filterer
X
X
X
X
90
Appendix H. (continued) Presence/absence of macroinvertebrate taxa in each of the
three streams. An “x” indicates the taxon was present in that stream. A * indicates the
organism was terrestrial and excluded from further analysis. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe,
MT=mesotrophic stream, ET=eutrophic stream, HT=hypereutrophic stream.
Order
Family
Genus
Guild
MT
ET
HT
Diptera
Tipulidae
Antocha
Collector-gatherer
X
X
X
X
X
X
X
X
X
Diptera (A)
Other
Ephemeroptera
Baetidae
Collector-gatherer
Ephemeroptera
Baetidae
Baetis
Collector-gatherer
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
X
X
Ephemeroptera
Heptageniidae
Scraper
X
X
Ephemeroptera
Isonychiidae
Isonychia
Collector-filterer
X
Ephemeroptera
Leptohypidae
Tricorythodes
Collector-gatherer
Ephemeroptera
Collector-gatherer
X
X
X
X
X
X
Gastropoda
Ancylidae
Ferrissia
Scraper
X
Gastropoda
Ancylidae
Laevapex fuscus
Scraper
X
Gastropoda
Bithyniidae
Bithynia tentaculata
Scraper
X
Gastropoda
Hydrobiidae
Amnicola
Scraper
X
Gastropoda
Physidae
Physa
Scraper
Gastropoda
Physidae
Physella gyrina
Scraper
Gastropoda
Physidae
Gastropoda
Planorbidae
X
Scraper
Menetus dilatatus
Gastropoda
*Hemiptera
X
X
Scraper
Scraper
Aphidoidea
*Hemiptera (A)
X
X
X
X
X
X
*Hymenoptera
*Hymenoptera
(A)
Diapriidae (A)
*Insecta (A)
terr.
Isopoda
Asellidae
Lepidoptera
Crambidae
X
X
X
Caecidotea
Scavenger
X
Shredder
X
Nematoda
Collector-gatherer
X
Oligochaeta
Collector-gatherer
X
Predator
X
Plecoptera
Perlidae
Plecoptera
Perlodidae
Podocopa
Agnetina
Predator
Collector-filterer
X
X
X
X
X
91
Appendix H. (continued) Presence/absence of macroinvertebrate taxa in each of the
three streams. An “x” indicates the taxon was present in that stream. A * indicates the
organism was terrestrial and excluded from further analysis. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe,
MT=mesotrophic stream, ET=eutrophic stream, HT=hypereutrophic stream.
Order
Family
Genus
Guild
MT
Trichoptera
Apataniidae
Apatania
Scraper
X
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
X
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Cheumatopsyche
Trichoptera
Hydropsychidae
Potamyia flava
Trichoptera
Hydropsychidae
Trichoptera
Hydroptilidae
Trichoptera
Hydroptilidae (L)
Piercer
X
Trichoptera
Hydroptilidae (P)
Non-feeding
X
Trichoptera
Leptoceridae
Collector-gatherer
Trichoptera
Polycentropodidae Neureclipsis
Collector-filterer
X
Trichoptera
Polycentropodidae Nyctiophylax
Predator
X
Trichoptera
Polycentropodidae Polycentropus
Predator
X
Trichoptera
Polycentropodidae
Collector-filterer
X
Trichoptera
Psychomyiidae
Psychomyia
Collector-gatherer
X
Trichoptera
Rhyacophilidae
Rhyacophila
Predator
X
X
Trichoptera (L)
Collector-filterer
X
X
Trichoptera (P)
Non-feeding
Ochrotrichia
ET
HT
X
X
X
Collector-filterer
X
X
Collector-filterer
X
X
X
Collector-filterer
X
Piercer
X
X
X
X
X
Appendix I. Macroinvertebrate biomass (mg DW/m2) in mesotrophic stream by taxa for treatment. Values represent mean ±1
standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Acariformes
Hydrachnidia
Amphipoda
Gammaridae
Gammarus
Scraper
Coleoptera
Elmidae (A)
Stenelmis
Scraper
Coleoptera
Elmidae (L)
Parasite
Collector-gatherer
Coleoptera (L)
Collembola
Guild
Collector-gatherer
Poduridae
Podura aquatic
Cyclopoida
Collector-gatherer
Collector-filterer
Diptera
Chironomidae (A)
Other
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
Diptera
Chironomidae (P)
Non-feeding
N
P
N+P
1.15
(2.30)
0.00
(0.00)
2.25
(4.49)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.90
(4.53)
64.42
(33.93)
1.98
(1.23)
3.55
(1.58)
6.03
(3.44)
28.35
(14.88)
0.00
(0.00)
88.31
(113.96)
0.00
(0.00)
0.65
(1.30)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
30.42
(19.43)
2.41
(2.73)
1.56
(1.81)
5.06
(5.42)
32.21
(11.43)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
4.44
(5.15)
62.17
(32.80)
3.12
(0.49)
2.09
(2.93)
6.94
(4.25)
21.91
(4.93)
0.00
(0.00)
0.00
(0.00)
2.25
(4.49)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
41.54
(30.94)
4.81
(3.18)
5.92
(8.63)
6.35
(1.37)
37.37
(15.96)
Stream
Average
0.29
(1.15)
22.08
(64.48)
1.12
(3.07)
0.16
(0.65)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.58
(4.10)
49.64
(30.47)
3.08
(2.26)
3.28
(4.56)
6.09
(3.57)
29.96
(12.66)
92
Control
Appendix I. (continued) Macroinvertebrate biomass (mg DW/m2) in mesotrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Diptera
Empididae
Hemerodromia
Predator
Diptera
Empididae
Metachela/Chelifera
Predator
Diptera
Simuliidae (A)
Diptera
Simuliidae (L)
Diptera
Simuliidae (L)
Collector-filterer
Diptera
Simuliidae (P)
Non-feeding
Diptera
Tipulidae
Other
Prosimulium
Antocha
Diptera (A)
Collector-filterer
Collector-gatherer
Other
Ephemeroptera
Baetidae
Collector-gatherer
Ephemeroptera
Baetidae
Baetis
Collector-gatherer
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
Ephemeroptera
Heptageniidae
Ephemeroptera
Isonychiidae
Scraper
Isonychia
Collector-filterer
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.18
(0.20)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
14.40
(28.80)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.18
(0.21)
0.00
(0.00)
0.03
(0.06)
0.00
(0.00)
0.00
(0.00)
0.15
(0.31)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.36
(0.52)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
9.64
(19.27)
0.00
(0.00)
0.00
(0.00)
0.25
(0.31)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.12
(0.15)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
26.99
(31.55)
0.00
(0.00)
0.00
(0.00)
Stream
Average
0.06
(0.18)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.21
(0.29)
0.00
(0.00)
0.01
(0.03)
0.00
(0.00)
9.16
(20.07)
0.04
(0.15)
3.60
(14.40)
93
Control
Appendix I. (continued) Macroinvertebrate biomass (mg DW/m2) in mesotrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Ephemeroptera
Family
Leptohyphidae
Genus
Tricorythodes
Ephemeroptera
Control
N
P
N+P
Collector-gatherer
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
8.82
(17.63)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.07
(0.13)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Collector-gatherer
Gastropoda
Ancylidae
Ferrissia
Scraper
Gastropoda
Ancylidae
Laevapex fuscus
Scraper
Gastropoda
Bithyniidae
Bithynia tentaculata
Scraper
Gastropoda
Hydrobiidae
Amnicola
Scraper
Gastropoda
Physidae
Physa
Scraper
Gastropoda
Physidae
Physella gyrina
Scraper
Gastropoda
Physidae
Gastropoda
Planorbidae
Scraper
Menetus dilatatus
Gastropoda
Scraper
Scraper
Isopoda
Asellidae
Lepidoptera
Crambidae
Caecidotea
Scavenger
Shredder
Stream
Average
0.00
(0.00)
0.02
(0.07)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.20
(8.82)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
94
Guild
Appendix I. (continued) Macroinvertebrate biomass (mg DW/m2) in mesotrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Control
N
P
N+P
Nematoda
Collector-gatherer
Oligochaeta
Collector-gatherer
0.00
(0.00)
0.00
(0.00)
1.55
(3.10)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
9.68
(18.24)
37.41
(14.58)
0.00
(0.00)
0.85
(1.69)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
43.77
(29.81)
0.00
(0.00)
0.47
(0.94)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.65
(1.29)
1.39
(2.78)
10.80
(12.48)
1.30
(2.59)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.09
(0.19)
0.00
(0.00)
78.12
(156.24)
0.00
(0.00)
0.00
(0.00)
0.36
(0.42)
5.44
(6.88)
31.67
(28.50)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Perlidae
Plecoptera
Perlodidae
Agnetina
Predator
Predator
Podocopa
Collector-filterer
Trichoptera
Apataniidae
Apatania
Scraper
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Cheumatopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Potamyia flava
Collector-filterer
Trichoptera
Hydropsychidae
Trichoptera
Hydroptilidae
Trichoptera
Hydroptilidae (L)
Collector-filterer
Ochrotrichia
Piercer
Piercer
Stream
Average
0.02
(0.09)
0.00
(0.00)
19.92
(78.03)
0.00
(0.00)
0.00
(0.00)
0.25
(0.67)
4.13
(9.63)
30.91
(24.02)
0.32
(1.30)
0.33
(0.94)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
95
Guild
Plecoptera
Genus
Appendix I. (continued) Macroinvertebrate biomass (mg DW/m2) in mesotrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Control
N
P
N+P
0.00
(0.00)
0.00
(0.00)
9.98
(15.37)
0.84
(1.69)
6.46
(7.53)
0.00
(0.00)
0.00
(0.00)
0.35
(0.70)
0.19
(0.39)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
17.63
(9.76)
0.78
(1.56)
3.51
(7.01)
0.00
(0.00)
1.76
(3.51)
0.78
(1.56)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
6.90
(4.78)
1.45
(2.90)
28.17
(28.23)
0.00
(0.00)
4.66
(5.58)
0.00
(0.00)
0.28
(0.57)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
11.28
(8.25)
0.63
(0.74)
14.51
(17.31)
0.00
(0.00)
2.36
(4.72)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Trichoptera
Hydroptilidae (P)
Non-feeding
Trichoptera
Leptoceridae
Collector-gatherer
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
Trichoptera
Polycentropodidae
Nyctiophylax
Predator
Trichoptera
Polycentropodidae
Polycentropus
Predator
Trichoptera
Polycentropodidae
Trichoptera
Psychomyiidae
Psychomyia
Collector-gatherer
Trichoptera
Rhyacophilidae
Rhyacophila
Predator
Collector-filterer
Trichoptera (L)
Collector-filterer
Trichoptera (P)
Non-feeding
Stream
Average
0.00
(0.00)
0.00
(0.00)
11.45
(10.04)
0.93
(1.72)
13.16
(18.38)
0.00
(0.00)
2.19
(4.01)
0.28
(0.83)
0.12
(0.33)
0.00
(0.00)
96
Appendix J. Macroinvertebrate biomass (mg DW/m2) in eutrophic stream by taxa for treatment. Values represent mean ±1 standard
deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva, (P)=pupa,
(A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Acariformes
Hydrachnidia
Amphipoda
Gammaridae
Coleoptera
Elmidae (A)
Coleoptera
Elmidae (L)
Genus
Parasite
Gammarus
Stenelmis
Scraper
Scraper
Collector-gatherer
Coleoptera (L)
Collembola
Guild
Collector-gatherer
Poduridae
Podura aquatic
Cyclopoida
Collector-gatherer
Collector-filterer
Diptera
Chironomidae (A)
Other
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
Diptera
Chironomidae (P)
Non-feeding
Control
N
P
N+P
1.90
1.38
1.86
3.02
(0.51)
(1.07)
(1.49)
(2.13)
0.00
0.00
0.00
(0.00)
(0.00)
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.73
(0.85)
0.00
(0.00)
3.73
(4.32)
190.81
(155.12)
2.98
(2.45)
4.23
(2.96)
18.08
(15.18)
43.81
(6.65)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.38
(2.05)
0.00
(0.00)
0.00
(0.00)
155.15
(45.71)
6.89
(5.90)
4.15
(3.91)
12.34
(6.07)
68.29
(26.41)
0.00
(0.00)
0.18
(0.35)
0.06
(0.11)
0.56
(1.11)
0.00
(0.00)
4.73
(9.46)
342.50
(155.77)
7.70
(3.70)
15.49
(4.27)
27.78
(11.59)
77.32
(40.59)
0.00
(0.00)
0.00
(0.00)
0.12
(0.23)
0.56
(0.70)
0.00
(0.00)
0.00
(0.00)
222.12
(25.53)
3.95
(1.68)
5.51
(8.12)
14.89
(5.27)
50.25
(29.27)
97
0.00
(0.00)
Stream
Average
2.04
(1.42)
0.00
(0.00)
0.00
(0.00)
0.04
(0.18)
0.04
(0.12)
1.05
(1.40)
0.00
(0.00)
2.12
(5.15)
227.64
(124.51)
5.38
(3.95)
7.35
(6.75)
18.27
(11.07)
59.92
(29.03)
Appendix J. (continued) Macroinvertebrate biomass (mg DW/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Diptera
Empididae
Hemerodromia
Predator
Diptera
Empididae
Metachela/Chelifera
Predator
Diptera
Simuliidae (A)
Diptera
Simuliidae (L)
Diptera
Simuliidae (L)
Collector-filterer
Diptera
Simuliidae (P)
Non-feeding
Diptera
Tipulidae
Other
Prosimulium
Antocha
Diptera (A)
Collector-filterer
Collector-gatherer
Other
Ephemeroptera
Baetidae
Collector-gatherer
Ephemeroptera
Baetidae
Baetis
Collector-gatherer
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
Ephemeroptera
Heptageniidae
Ephemeroptera
Isonychiidae
Scraper
Isonychia
Collector-filterer
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
56.63
(108.61)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.08
(2.15)
1.91
(2.45)
0.00
(0.00)
28.42
(25.57)
0.00
(0.00)
0.00
(0.00)
0.23
(0.45)
0.00
(0.00)
1.54
(3.07)
0.00
(0.00)
0.18
(0.36)
0.00
(0.00)
1.40
(0.97)
0.00
(0.00)
1.09
(2.18)
0.00
(0.00)
37.32
(74.64)
0.12
(0.25)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.54
(0.83)
0.00
(0.00)
1.76
(3.52)
9.80
(19.60)
45.75
(58.75)
0.37
(0.74)
0.00
(0.00)
Stream
Average
0.06
(0.23)
0.00
(0.00)
0.38
(1.54)
0.00
(0.00)
0.04
(0.18)
0.00
(0.00)
0.49
(0.82)
0.27
(1.08)
1.19
(2.29)
2.45
(9.80)
42.03
(66.41)
0.12
(0.38)
0.00
(0.00)
98
Control
Appendix J. (continued) Macroinvertebrate biomass (mg DW/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Ephemeroptera
Family
Leptohyphidae
Genus
Tricorythodes
Ephemeroptera
Control
N
P
N+P
Collector-gatherer
3.39
(6.77)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.05
(0.10)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
3.11
(6.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
6.62
(13.25)
0.00
(0.00)
0.05
(0.10)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.31
(4.62)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
12.80
(25.59)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Collector-gatherer
Gastropoda
Ancylidae
Ferrissia
Scraper
Gastropoda
Ancylidae
Laevapex fuscus
Scraper
Gastropoda
Bithyniidae
Bithynia tentaculata
Scraper
Gastropoda
Hydrobiidae
Amnicola
Scraper
Gastropoda
Physidae
Physa
Scraper
Gastropoda
Physidae
Physella gyrina
Scraper
Gastropoda
Physidae
Gastropoda
Planorbidae
Scraper
Menetus dilatatus
Gastropoda
Scraper
Scraper
Isopoda
Asellidae
Lepidoptera
Crambidae
Caecidotea
Scavenger
Shredder
Stream
Average
0.85
(3.39)
0.03
(0.07)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.35
(3.75)
0.00
(0.00)
3.20
(12.80)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.66
(6.62)
99
Guild
Appendix J. (continued) Macroinvertebrate biomass (mg DW/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Control
N
P
N+P
Nematoda
Collector-gatherer
Oligochaeta
Collector-gatherer
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
7.67
(15.34)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.47
(0.95)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
19.52
(16.71)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
17.62
(20.89)
0.54
(1.08)
0.94
(1.89)
0.00
(0.00)
0.00
(0.00)
0.75
(1.51)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.74
(1.47)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
38.73
(69.93)
0.00
(0.00)
3.94
(7.88)
0.00
(0.00)
0.00
(0.00)
0.22
(0.45)
Perlidae
Plecoptera
Perlodidae
Agnetina
Predator
Predator
Podocopa
Collector-filterer
Trichoptera
Apataniidae
Apatania
Scraper
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Cheumatopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Potamyia flava
Collector-filterer
Trichoptera
Hydropsychidae
Trichoptera
Hydroptilidae
Trichoptera
Hydroptilidae (L)
Collector-filterer
Ochrotrichia
Piercer
Piercer
Stream
Average
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.18
(0.74)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
20.89
(36.10)
0.14
(0.54)
1.22
(3.99)
0.00
(0.00)
0.00
(0.00)
0.36
(0.87)
100
Guild
Plecoptera
Genus
Appendix J. (continued) Macroinvertebrate biomass (mg DW/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Trichoptera
Hydroptilidae (P)
Non-feeding
Trichoptera
Leptoceridae
Collector-gatherer
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
Trichoptera
Polycentropodidae
Nyctiophylax
Predator
Trichoptera
Polycentropodidae
Polycentropus
Predator
Trichoptera
Polycentropodidae
Trichoptera
Psychomyiidae
Psychomyia
Collector-gatherer
Trichoptera
Rhyacophilidae
Rhyacophila
Predator
Collector-filterer
Trichoptera (L)
Collector-filterer
Trichoptera (P)
Non-feeding
Control
N
P
N+P
2.01
(2.69)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.32
(4.63)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.31
(4.62)
0.12
(0.24)
0.00
(0.00)
0.79
(1.01)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.24
(0.89)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Stream
Average
1.01
(1.54)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.16
(3.16)
0.03
(0.12)
0.00
(0.00)
101
Appendix K. Macroinvertebrate biomass (mg DW/m2) in hypereutrophic stream by taxa for treatment. Values represent mean ±1
standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Acariformes
Hydrachnidia
Amphipoda
Gammaridae
Gammarus
Scraper
Coleoptera
Elmidae (A)
Stenelmis
Scraper
Coleoptera
Elmidae (L)
Parasite
Collector-gatherer
Coleoptera (L)
Collembola
Guild
Collector-gatherer
Poduridae
Podura aquatic
Cyclopoida
Collector-gatherer
Collector-filterer
Diptera
Chironomidae (A)
Other
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
Diptera
Chironomidae (P)
Non-feeding
N
P
N+P
0.41
(0.82)
46.83
(60.06)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.64
(0.74)
0.00
(0.00)
2.08
(4.15)
73.47
(38.60)
10.53
(7.62)
3.65
(0.68)
43.95
(9.00)
113.40
(17.86)
1.14
(1.33)
74.88
(86.53)
2.25
(4.49)
0.11
(0.22)
0.00
(0.00)
2.00
(3.14)
0.00
(0.00)
1.80
(3.60)
63.67
(34.60)
7.25
(4.77)
3.97
(1.34)
29.21
(17.18)
67.01
(26.95)
1.60
(1.42)
144.14
(204.02)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.46
(0.80)
0.16
(0.28)
0.00
(0.00)
76.94
(61.22)
6.96
(2.90)
4.50
(4.34)
48.88
(16.77)
106.52
(43.84)
1.39
(0.97)
38.08
(45.08)
0.00
(0.00)
0.11
(0.22)
0.00
(0.00)
1.02
(0.69)
0.00
(0.00)
2.22
(4.44)
60.79
(37.60)
4.24
(0.54)
2.98
(1.53)
22.99
(10.52)
63.14
(14.80)
Stream
Average
1.10
(1.11)
71.44
(101.91)
0.60
(2.32)
0.06
(0.15)
0.00
(0.00)
1.07
(1.67)
0.03
(0.13)
1.62
(3.38)
68.17
(38.21)
7.26
(4.93)
3.73
(2.00)
35.41
(16.10)
86.25
(33.20)
102
Control
Appendix K. (continued) Macroinvertebrate biomass (mg DW/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Diptera
Empididae
Hemerodromia
Predator
Diptera
Empididae
Metachela/Chelifera
Predator
Diptera
Simuliidae (A)
Diptera
Simuliidae (L)
Diptera
Simuliidae (L)
Collector-filterer
Diptera
Simuliidae (P)
Non-feeding
Diptera
Tipulidae
Other
Prosimulium
Antocha
Diptera (A)
Collector-filterer
Collector-gatherer
Other
Ephemeroptera
Baetidae
Collector-gatherer
Ephemeroptera
Baetidae
Baetis
Collector-gatherer
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
Ephemeroptera
Heptageniidae
Ephemeroptera
Isonychiidae
Scraper
Isonychia
Collector-filterer
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.94
(3.88)
0.00
(0.00)
8.30
(10.59)
8.43
(5.93)
0.41
(0.81)
18.74
(28.66)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.44
(1.94)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.77
(5.53)
6.40
(6.26)
0.00
(0.00)
0.00
(0.00)
1.89
(3.77)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.30
(0.52)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
3.69
(6.39)
7.78
(3.27)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
15.91
(12.82)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Stream
Average
0.65
(1.43)
0.06
(0.23)
0.00
(0.00)
0.52
(2.00)
0.00
(0.00)
3.69
(6.83)
9.75
(8.25)
0.11
(0.42)
5.00
(15.80)
0.50
(1.95)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
103
Control
Appendix K. (continued) Macroinvertebrate biomass (mg DW/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Ephemeroptera
Family
Leptohyphidae
Genus
Tricorythodes
Ephemeroptera
Control
N
P
N+P
Collector-gatherer
0.00
(0.00)
0.00
(0.00)
0.36
(0.24)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.75
(0.59)
0.00
(0.00)
9.56
(14.86)
0.00
(0.00)
0.00
(0.00)
0.38
(0.75)
0.08
(0.17)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.80
(1.40)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.36
(2.36)
0.00
(0.00)
0.11
(0.19)
0.00
(0.00)
0.05
(0.08)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.92
(0.87)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
6.45
(8.42)
0.38
(0.75)
0.00
(0.00)
1.01
(2.02)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.91
(0.34)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Collector-gatherer
Gastropoda
Ancylidae
Ferrissia
Scraper
Gastropoda
Ancylidae
Laevapex fuscus
Scraper
Gastropoda
Bithyniidae
Bithynia tentaculata
Scraper
Gastropoda
Hydrobiidae
Amnicola
Scraper
Gastropoda
Physidae
Physa
Scraper
Gastropoda
Physidae
Physella gyrina
Scraper
Gastropoda
Physidae
Gastropoda
Planorbidae
Scraper
Menetus dilatatus
Gastropoda
Scraper
Scraper
Isopoda
Asellidae
Lepidoptera
Crambidae
Caecidotea
Scavenger
Shredder
Stream
Average
0.27
(1.06)
0.10
(0.39)
1.86
(4.84)
0.10
(0.39)
0.01
(0.04)
0.27
(1.04)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.57
(0.95)
0.00
(0.00)
2.55
(8.15)
0.00
(0.00)
104
Guild
Appendix K. (continued) Macroinvertebrate biomass (mg DW/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Control
N
P
N+P
Nematoda
Collector-gatherer
Oligochaeta
Collector-gatherer
0.11
(0.21)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.16
(0.31)
0.00
(0.00)
0.00
(0.00)
170.94
(79.62)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.04
(0.08)
0.10
(0.20)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
77.84
(38.34)
0.00
(0.00)
0.00
(0.00)
0.51
(0.51)
0.71
(1.42)
0.00
(0.00)
0.00
(0.00)
0.24
(0.42)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
110.08
(85.53)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.24
(2.15)
0.00
(0.00)
0.06
(0.13)
0.05
(0.09)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
133.50
(141.82)
0.00
(0.00)
1.05
(2.10)
0.24
(0.49)
0.00
(0.00)
0.00
(0.00)
Perlidae
Plecoptera
Perlodidae
Agnetina
Predator
Predator
Podocopa
Collector-filterer
Trichoptera
Apataniidae
Apatania
Scraper
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Cheumatopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Potamyia flava
Collector-filterer
Trichoptera
Hydropsychidae
Trichoptera
Hydroptilidae
Trichoptera
Hydroptilidae (L)
Collector-filterer
Ochrotrichia
Piercer
Piercer
Stream
Average
0.06
(0.13)
0.09
(0.21)
0.00
(0.00)
0.00
(0.00)
0.04
(0.16)
0.00
(0.00)
0.00
(0.00)
123.96
(91.29)
0.00
(0.00)
0.28
(1.08)
0.20
(0.39)
0.44
(1.17)
0.00
(0.00)
105
Guild
Plecoptera
Genus
Appendix K. (continued) Macroinvertebrate biomass (mg DW/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Trichoptera
Hydroptilidae (P)
Non-feeding
Trichoptera
Leptoceridae
Collector-gatherer
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
Trichoptera
Polycentropodidae
Nyctiophylax
Predator
Trichoptera
Polycentropodidae
Polycentropus
Predator
Trichoptera
Polycentropodidae
Trichoptera
Psychomyiidae
Psychomyia
Collector-gatherer
Trichoptera
Rhyacophilidae
Rhyacophila
Predator
Collector-filterer
Trichoptera (L)
Collector-filterer
Trichoptera (P)
Non-feeding
Control
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.13
(0.25)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.12
(0.23)
0.00
(0.00)
0.00
(0.00)
0.04
(0.08)
0.00
(0.00)
Stream
Average
0.00
(0.00)
0.03
(0.13)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.03
(0.12)
0.00
(0.00)
0.00
(0.00)
0.01
(0.04)
0.00
(0.00)
106
Appendix L. Macroinvertebrate density (number/m2) in mesotrophic stream by taxa for treatment. Values represent mean ±1 standard
deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva, (P)=pupa,
(A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Acariformes
Hydrachnidia
Amphipoda
Gammaridae
Gammarus
Scraper
Coleoptera
Elmidae (A)
Stenelmis
Scraper
Coleoptera
Elmidae (L)
Parasite
Collector-gatherer
Coleoptera (L)
Collembola
Guild
Collector-gatherer
Poduridae
Podura aquatic
Cyclopoida
Collector-gatherer
Collector-filterer
Diptera
Chironomidae (A)
Other
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
Diptera
Chironomidae (P)
Non-feeding
N
P
N+P
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
22.45
(0.00)
544.39
(202.77)
84.18
(33.67)
61.74
(11.22)
303.06
(144.91)
123.47
(64.81)
0.00
(0.00)
22.45
(31.75)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
381.64
(260.51)
56.12
(28.98)
28.06
(21.49)
202.04
(111.50)
140.31
(49.78)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
11.22
(12.96)
600.52
(218.71)
95.41
(49.78)
50.51
(46.28)
246.94
(79.90)
95.41
(21.49)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
505.11
(309.17)
117.86
(78.57)
95.41
(99.34)
263.78
(53.05)
162.76
(69.50)
Stream
Average
1.40
(5.61)
5.61
(17.39)
2.81
(7.67)
1.40
(5.61)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
8.42
(11.22)
507.91
(239.53)
88.39
(51.50)
58.93
(56.12)
253.96
(99.63)
130.49
(55.12)
107
Control
Appendix L. (continued) Macroinvertebrate density (number/m2) in mesotrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Diptera
Empididae
Hemerodromia
Predator
Diptera
Empididae
Metachela/Chelifera
Predator
Diptera
Simuliidae (A)
Diptera
Simuliidae (L)
Diptera
Simuliidae (L)
Collector-filterer
Diptera
Simuliidae (P)
Non-feeding
Diptera
Tipulidae
Other
Prosimulium
Antocha
Diptera (A)
Collector-filterer
Collector-gatherer
Other
Ephemeroptera
Baetidae
Collector-gatherer
Ephemeroptera
Baetidae
Baetis
Collector-gatherer
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
Ephemeroptera
Heptageniidae
Ephemeroptera
Isonychiidae
Scraper
Isonychia
Collector-filterer
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
22.45
(18.33)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
16.84
(21.49)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
16.84
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
11.22
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
16.84
(21.49)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
16.84
(21.49)
5.61
(11.22)
0.00
(0.00)
Stream
Average
2.81
(7.67)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
18.24
(16.84)
0.00
(0.00)
2.81
(7.67)
0.00
(0.00)
7.02
(13.52)
4.21
(9.05)
1.40
(5.61)
108
Control
Appendix L. (continued) Macroinvertebrate density (number/m2) in mesotrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Ephemeroptera
Family
Leptohyphidae
Genus
Tricorythodes
Ephemeroptera
Control
N
P
N+P
Collector-gatherer
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
22.45
(44.90)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
11.22
(22.45)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Collector-gatherer
Gastropoda
Ancylidae
Ferrissia
Scraper
Gastropoda
Ancylidae
Laevapex fuscus
Scraper
Gastropoda
Bithyniidae
Bithynia tentaculata
Scraper
Gastropoda
Hydrobiidae
Amnicola
Scraper
Gastropoda
Physidae
Physa
Scraper
Gastropoda
Physidae
Physella gyrina
Scraper
Gastropoda
Physidae
Gastropoda
Planorbidae
Scraper
Menetus dilatatus
Gastropoda
Scraper
Scraper
Isopoda
Asellidae
Lepidoptera
Crambidae
Caecidotea
Scavenger
Shredder
Stream
Average
0.00
(0.00)
1.40
(5.61)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.40
(5.61)
0.00
(0.00)
0.00
(0.00)
8.42
(24.42)
0.00
(0.00)
0.00
(0.00)
109
Guild
Appendix L. (continued) Macroinvertebrate density (number/m2) in mesotrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Control
N
P
N+P
Nematoda
Collector-gatherer
Oligochaeta
Collector-gatherer
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
22.45
(31.75)
123.47
(38.88)
0.00
(0.00)
11.22
(22.45)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
95.41
(49.78)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
28.06
(28.25)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
11.22
(12.96)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
11.22
(12.96)
44.90
(63.50)
117.86
(84.74)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Perlidae
Plecoptera
Perlodidae
Agnetina
Predator
Predator
Podocopa
Collector-filterer
Trichoptera
Apataniidae
Apatania
Scraper
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Cheumatopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Potamyia flava
Collector-filterer
Trichoptera
Hydropsychidae
Trichoptera
Hydroptilidae
Trichoptera
Hydroptilidae (L)
Collector-filterer
Ochrotrichia
Piercer
Piercer
Stream
Average
2.81
(7.67)
1.40
(5.61)
2.81
(7.67)
0.00
(0.00)
0.00
(0.00)
4.21
(9.05)
18.24
(36.86)
91.20
(62.68)
1.40
(5.61)
4.21
(12.21)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
110
Guild
Plecoptera
Genus
Appendix L. (continued) Macroinvertebrate density (number/m2) in mesotrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Control
N
P
N+P
0.00
(0.00)
0.00
(0.00)
61.74
(61.82)
5.61
(11.22)
33.67
(53.44)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
39.29
(21.49)
5.61
(11.22)
16.84
(33.67)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
39.29
(38.34)
5.61
(11.22)
61.74
(33.67)
5.61
(11.22)
16.84
(21.49)
0.00
(0.00)
11.22
(22.45)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
61.74
(56.12)
11.22
(12.96)
44.90
(51.84)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Trichoptera
Hydroptilidae (P)
Non-feeding
Trichoptera
Leptoceridae
Collector-gatherer
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
Trichoptera
Polycentropodidae
Nyctiophylax
Predator
Trichoptera
Polycentropodidae
Polycentropus
Predator
Trichoptera
Polycentropodidae
Trichoptera
Psychomyiidae
Psychomyia
Collector-gatherer
Trichoptera
Rhyacophilidae
Rhyacophila
Predator
Collector-filterer
Trichoptera (L)
Collector-filterer
Trichoptera (P)
Non-feeding
Stream
Average
0.00
(0.00)
0.00
(0.00)
50.51
(43.76)
7.02
(10.75)
39.29
(42.99)
2.81
(7.67)
7.02
(13.52)
2.81
(7.67)
5.61
(12.96)
0.00
(0.00)
111
Appendix M. Macroinvertebrate density (number/m2) in eutrophic stream by taxa for treatment. Values represent mean ±1 standard
deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva, (P)=pupa,
(A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Acariformes
Hydrachnidia
Amphipoda
Gammaridae
Gammarus
Scraper
Coleoptera
Elmidae (A)
Stenelmis
Scraper
Coleoptera
Elmidae (L)
Parasite
Collector-gatherer
Coleoptera (L)
Collembola
Guild
Collector-gatherer
Poduridae
Podura aquatic
Cyclopoida
Collector-gatherer
Collector-filterer
Diptera
Chironomidae (A)
Other
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
Diptera
Chironomidae (P)
Non-feeding
N
P
N+P
72.96
(46.28)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
16.84
(11.22)
0.00
(0.00)
11.22
(12.96)
1229.09
(615.17)
112.25
(95.24)
95.41
(33.67)
555.62
(329.11)
190.82
(28.98)
179.59
(314.82)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
44.90
(40.99)
0.00
(0.00)
5.61
(11.22)
1144.91
(397.80)
145.92
(97.85)
196.43
(162.27)
387.25
(202.77)
297.45
(115.02)
22.45
(18.33)
0.00
(0.00)
0.00
(0.00)
16.84
(33.67)
5.61
(11.22)
11.22
(22.45)
0.00
(0.00)
11.22
(22.45)
2553.60
(695.90)
218.88
(107.47)
286.23
(104.29)
1105.62
(317.68)
336.74
(176.77)
179.59
(170.97)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
16.84
(21.49)
0.00
(0.00)
16.84
(33.67)
1262.77
(282.41)
101.02
(38.88)
157.14
(180.53)
538.78
(201.63)
218.88
(127.49)
Stream
Average
113.65
(176.47)
0.00
(0.00)
0.00
(0.00)
4.21
(16.84)
2.81
(7.67)
22.45
(27.19)
0.00
(0.00)
11.22
(20.08)
1547.59
(762.85)
144.52
(92.73)
183.80
(138.92)
646.82
(370.99)
260.97
(126.43)
112
Control
Appendix M. (continued) Macroinvertebrate density (number/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Diptera
Empididae
Hemerodromia
Predator
Diptera
Empididae
Metachela/Chelifera
Predator
Diptera
Simuliidae (A)
Diptera
Simuliidae (L)
Diptera
Simuliidae (L)
Collector-filterer
Diptera
Simuliidae (P)
Non-feeding
Diptera
Tipulidae
Other
Prosimulium
Antocha
Diptera (A)
Collector-filterer
Collector-gatherer
Other
Ephemeroptera
Baetidae
Collector-gatherer
Ephemeroptera
Baetidae
Baetis
Collector-gatherer
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
Ephemeroptera
Heptageniidae
Ephemeroptera
Isonychiidae
Scraper
Isonychia
Collector-filterer
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
22.45
(31.75)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
16.84
(11.22)
0.00
(0.00)
22.45
(18.33)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
44.90
(25.92)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
22.45
(31.75)
0.00
(0.00)
11.22
(22.45)
33.67
(67.35)
11.22
(12.96)
11.22
(22.45)
0.00
(0.00)
Stream
Average
1.40
(5.61)
0.00
(0.00)
1.40
(5.61)
0.00
(0.00)
1.40
(5.61)
0.00
(0.00)
18.24
(26.20)
1.40
(5.61)
8.42
(13.90)
8.42
(33.67)
15.43
(19.60)
4.21
(12.21)
0.00
(0.00)
113
Control
Appendix M. (continued) Macroinvertebrate density (number/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Ephemeroptera
Family
Leptohyphidae
Genus
Tricorythodes
Ephemeroptera
Control
N
P
N+P
Collector-gatherer
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Collector-gatherer
Gastropoda
Ancylidae
Ferrissia
Scraper
Gastropoda
Ancylidae
Laevapex fuscus
Scraper
Gastropoda
Bithyniidae
Bithynia tentaculata
Scraper
Gastropoda
Hydrobiidae
Amnicola
Scraper
Gastropoda
Physidae
Physa
Scraper
Gastropoda
Physidae
Physella gyrina
Scraper
Gastropoda
Physidae
Gastropoda
Planorbidae
Scraper
Menetus dilatatus
Gastropoda
Scraper
Scraper
Isopoda
Asellidae
Lepidoptera
Crambidae
Caecidotea
Scavenger
Shredder
Stream
Average
1.40
(5.61)
4.21
(9.05)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
2.81
(7.67)
0.00
(0.00)
1.40
(5.61)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
1.40
(5.61)
114
Guild
Appendix M. (continued) Macroinvertebrate density (number/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Control
N
P
N+P
Nematoda
Collector-gatherer
Oligochaeta
Collector-gatherer
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
67.35
(57.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
33.67
(42.99)
5.61
(11.22)
11.22
(22.45)
0.00
(0.00)
0.00
(0.00)
11.22
(22.45)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
44.90
(60.79)
0.00
(0.00)
16.84
(33.67)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
Perlidae
Plecoptera
Perlodidae
Agnetina
Predator
Predator
Podocopa
Collector-filterer
Trichoptera
Apataniidae
Apatania
Scraper
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Cheumatopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Potamyia flava
Collector-filterer
Trichoptera
Hydropsychidae
Trichoptera
Hydroptilidae
Trichoptera
Hydroptilidae (L)
Collector-filterer
Ochrotrichia
Piercer
Piercer
Stream
Average
0.00
(0.00)
1.40
(5.61)
0.00
(0.00)
1.40
(5.61)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
37.88
(48.30)
1.40
(5.61)
7.02
(19.60)
0.00
(0.00)
0.00
(0.00)
5.61
(12.96)
115
Guild
Plecoptera
Genus
Appendix M. (continued) Macroinvertebrate density (number/m2) in eutrophic stream by taxa for treatment. Values represent mean
±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Control
N
P
N+P
16.84
(21.49)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
16.84
(21.49)
5.61
(11.22)
0.00
(0.00)
11.22
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
16.84
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Trichoptera
Hydroptilidae (P)
Non-feeding
Trichoptera
Leptoceridae
Collector-gatherer
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
Trichoptera
Polycentropodidae
Nyctiophylax
Predator
Trichoptera
Polycentropodidae
Polycentropus
Predator
Trichoptera
Polycentropodidae
Trichoptera
Psychomyiidae
Psychomyia
Collector-gatherer
Trichoptera
Rhyacophilidae
Rhyacophila
Predator
Collector-filterer
Trichoptera (L)
Collector-filterer
Trichoptera (P)
Non-feeding
Stream
Average
11.22
(14.20)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(12.96)
2.81
(7.67)
0.00
(0.00)
116
Appendix N. Macroinvertebrate density (number/m2) in hypereutrophic stream by taxa for treatment. Values represent mean ±1
standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family, (L)=larva,
(P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Acariformes
Hydrachnidia
Amphipoda
Gammaridae
Gammarus
Scraper
Coleoptera
Elmidae (A)
Stenelmis
Scraper
Coleoptera
Elmidae (L)
Parasite
Collector-gatherer
Coleoptera (L)
Collembola
Guild
Collector-gatherer
Poduridae
Podura aquatic
Cyclopoida
Collector-gatherer
Collector-filterer
Diptera
Chironomidae (A)
Other
Diptera
Chironomidae (L)
S.F. Tanypodinae
Predator
Diptera
Chironomidae (L)
S.F. Orthocladiinae
Collector-gatherer
Diptera
Chironomidae (L)
T. Chironomini
Collector-gatherer
Diptera
Chironomidae (L)
T. Tanytarsini
Collector-gatherer
Diptera
Chironomidae (P)
Non-feeding
N
P
N+P
5.61
(11.22)
16.84
(21.49)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
11.22
(12.96)
0.00
(0.00)
5.61
(11.22)
550.01
(241.44)
134.70
(66.09)
72.96
(33.67)
1296.44
(407.14)
493.88
(77.77)
33.67
(42.99)
22.45
(25.92)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
28.06
(42.50)
0.00
(0.00)
5.61
(11.22)
510.72
(287.71)
145.92
(76.68)
78.57
(28.98)
1038.28
(481.61)
291.84
(117.37)
22.45
(22.45)
37.42
(46.73)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
7.48
(12.96)
7.48
(12.96)
0.00
(0.00)
621.09
(340.71)
89.80
(59.40)
97.28
(12.96)
1504.10
(669.35)
463.95
(190.93)
28.06
(21.49)
11.22
(12.96)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
16.84
(11.22)
0.00
(0.00)
5.61
(11.22)
578.07
(362.62)
101.02
(38.88)
95.41
(102.67)
886.74
(265.31)
275.00
(64.48)
Stream
Average
22.45
(26.83)
20.95
(26.11)
1.50
(5.80)
2.99
(7.90)
0.00
(0.00)
16.46
(23.19)
1.50
(5.80)
4.49
(9.29)
561.23
(276.77)
119.73
(59.80)
85.31
(53.12)
1159.88
(469.65)
375.65
(144.61)
117
Control
Appendix N. (continued) Macroinvertebrate density (number/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Diptera
Empididae
Hemerodromia
Predator
Diptera
Empididae
Metachela/Chelifera
Predator
Diptera
Simuliidae (A)
Diptera
Simuliidae (L)
Diptera
Simuliidae (L)
Collector-filterer
Diptera
Simuliidae (P)
Non-feeding
Diptera
Tipulidae
Other
Prosimulium
Antocha
Diptera (A)
Collector-filterer
Collector-gatherer
Other
Ephemeroptera
Baetidae
Collector-gatherer
Ephemeroptera
Baetidae
Baetis
Collector-gatherer
Ephemeroptera
Heptageniidae
Maccaffertium
Scraper
Ephemeroptera
Heptageniidae
Ephemeroptera
Isonychiidae
Scraper
Isonychia
Collector-filterer
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
16.84
(21.49)
196.43
(97.64)
5.61
(11.22)
22.45
(18.33)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
28.06
(21.49)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
202.04
(135.94)
0.00
(0.00)
11.22
(22.45)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
7.48
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
7.48
(12.96)
269.39
(124.99)
0.00
(0.00)
7.48
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
213.27
(138.99)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Stream
Average
7.48
(16.25)
1.50
(5.80)
0.00
(0.00)
1.50
(5.80)
1.50
(5.80)
7.48
(13.86)
217.01
(114.68)
1.50
(5.80)
11.97
(16.68)
1.50
(5.80)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
118
Control
Appendix N. (continued) Macroinvertebrate density (number/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Ephemeroptera
Family
Leptohyphidae
Genus
Tricorythodes
Ephemeroptera
Control
N
P
N+P
Collector-gatherer
0.00
(0.00)
5.61
(11.22)
16.84
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
22.45
(18.33)
5.61
(11.22)
28.06
(33.67)
0.00
(0.00)
0.00
(0.00)
16.84
(33.67)
11.22
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
33.67
(28.98)
11.22
(22.45)
0.00
(0.00)
0.00
(0.00)
7.48
(12.96)
0.00
(0.00)
7.48
(12.96)
0.00
(0.00)
7.48
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
44.90
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
39.29
(46.28)
5.61
(11.22)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
56.12
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
Collector-gatherer
Gastropoda
Ancylidae
Ferrissia
Scraper
Gastropoda
Ancylidae
Laevapex fuscus
Scraper
Gastropoda
Bithyniidae
Bithynia tentaculata
Scraper
Gastropoda
Hydrobiidae
Amnicola
Scraper
Gastropoda
Physidae
Physa
Scraper
Gastropoda
Physidae
Physella gyrina
Scraper
Gastropoda
Physidae
Gastropoda
Planorbidae
Scraper
Menetus dilatatus
Gastropoda
Scraper
Scraper
Isopoda
Asellidae
Lepidoptera
Crambidae
Caecidotea
Scavenger
Shredder
Stream
Average
1.50
(5.80)
5.99
(17.93)
19.46
(26.65)
1.50
(5.80)
1.50
(5.80)
1.50
(5.80)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
38.91
(21.58)
4.49
(12.59)
7.48
(20.20)
0.00
(0.00)
119
Guild
Appendix N. (continued) Macroinvertebrate density (number/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Control
N
P
N+P
Nematoda
Collector-gatherer
Oligochaeta
Collector-gatherer
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
280.62
(110.74)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
179.59
(118.79)
0.00
(0.00)
0.00
(0.00)
22.45
(18.33)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
7.48
(12.96)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
172.11
(64.81)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
7.48
(12.96)
0.00
(0.00)
5.61
(11.22)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
112.25
(60.79)
0.00
(0.00)
11.22
(22.45)
11.22
(22.45)
0.00
(0.00)
0.00
(0.00)
Perlidae
Plecoptera
Perlodidae
Agnetina
Predator
Predator
Podocopa
Collector-filterer
Trichoptera
Apataniidae
Apatania
Scraper
Trichoptera
Helicopsychidae
Helicopsyche
Scraper
Trichoptera
Hydropsychidae
Ceratopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Cheumatopsyche
Collector-filterer
Trichoptera
Hydropsychidae
Potamyia flava
Collector-filterer
Trichoptera
Hydropsychidae
Trichoptera
Hydroptilidae
Trichoptera
Hydroptilidae (L)
Collector-filterer
Ochrotrichia
Piercer
Piercer
Stream
Average
4.49
(9.29)
4.49
(9.29)
0.00
(0.00)
0.00
(0.00)
1.50
(5.80)
0.00
(0.00)
0.00
(0.00)
187.08
(105.86)
0.00
(0.00)
2.99
(11.59)
8.98
(16.54)
2.99
(7.90)
0.00
(0.00)
120
Guild
Plecoptera
Genus
Appendix N. (continued) Macroinvertebrate density (number/m2) in hypereutrophic stream by taxa for treatment. Values represent
mean ±1 standard deviation; N=nitrogen treatments, P=phosphorus treatments, N+P=dual nutrient treatments. Under family,
(L)=larva, (P)=pupa, (A)=represents adult. Under genus, S.F.=subfamily, T.=tribe.
Order
Family
Genus
Guild
Trichoptera
Hydroptilidae (P)
Non-feeding
Trichoptera
Leptoceridae
Collector-gatherer
Trichoptera
Polycentropodidae
Neureclipsis
Collector-filterer
Trichoptera
Polycentropodidae
Nyctiophylax
Predator
Trichoptera
Polycentropodidae
Polycentropus
Predator
Trichoptera
Polycentropodidae
Trichoptera
Psychomyiidae
Psychomyia
Collector-gatherer
Trichoptera
Rhyacophilidae
Rhyacophila
Predator
Collector-filterer
Trichoptera (L)
Collector-filterer
Trichoptera (P)
Non-feeding
Control
N
P
N+P
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
5.61
(11.22)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
16.84
(33.67)
0.00
(0.00)
0.00
(0.00)
11.22
(12.96)
5.61
(11.22)
Stream
Average
0.00
(0.00)
1.50
(5.80)
0.00
(0.00)
0.00
(0.00)
0.00
(0.00)
4.49
(17.39)
0.00
(0.00)
0.00
(0.00)
2.99
(7.90)
1.50
(5.80)
121