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MACROINVERTEBRATE COMMUNITY RESPONSE TO
EXPERIMENTAL MANIPULATION OF LIGHT AND
NUTRIENTS IN SECOND ORDER STREAMS OF

' NORTHERN CALIFORNIA

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OSVALDO HERNANDEZ

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MACRONVERTEBRATE COMMUNITY RESPONSE TO EXPERIMENTAL
MANIPULATION OF LIGHT AND NUTRIENTS IN SECOND-ORDER STREAMS
OF NORTHERN CALIFORNIA.

By

Osvaldo Hernandez

A DISSERTATION

Submitted to
Michigan State University
in partial fulﬁllment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY
Entomology

2009

ABSTRACT
MACROINVERTEBRATE COMMUNITY RESPONSE TO EXPERIMENTAL
MANIPULATION OF LIGHT AND NUTRIENTS IN SECOND-ORDER STREAMS
OF NORTHERN CALIFORNIA.
By
Osvaldo Hernandez
Benthic macroinvertebrate community response to increased light and

nutrient augmentation, via riparian canopy removal and salmon carcass addition,
was evaluated in second order coastal streams of northern California. The study
followed a split—plot experimental design in a total of six streams within the
Klamath and Smith River catchments. Each stream consisted of two study
reaches (100m) separated by a 250m buffer. Three randomly selected streams
received carcass addition (whole-plot factor) and within each stream a study
reach randomly received riparian hardwood removal (subplot factor), resulting in
four replicated treatments (0- control, N- nutrient augmented, L- light augmented,
L+N- light and nutrient augmented). Stream reaches were sampled once pre-
manipulation and four times post-manipulation. Main effect results show that
while carcass addition had no effect on macroinvertebrate biomass, canopy
removal led to moderately greater biomass (F(1,4)=5.33 p=0.0819), and all post-
manipulation biomass were greater than pre-manipulation levels (F(4, 149)=5.14

=0.0007). Pairwise comparisons of macroinvertebrate biomass treatment
means, pre and post-manipulation, Show no difference within C and N
treatments, and greater macroinvertebrate biomass in L (p=0.09) and L+N

augmented treatments (p=0.0025). Additionally, red alder (Alnus rubra) leaf pack

mass loss was evaluated among experimental treatments in both spring and fall
2002. During both seasons, 1) all experimental treatments (N, L, L+N) had
greater mass loss relative to controls, and 2) control leaf packs had slowest
decay coefﬁcients (k), while light (L) treatments had fastest decay coefﬁcients.
Decay coefﬁcients in the spring were signiﬁcantly faster in streams with carcass
additions than in streams with out carcass additions. Decay coefﬁcients in the fall
were signiﬁcantly faster in the light (L) treatment than in the controls (C).

Results of this study revealed no signiﬁcant main effect of carcass addition
on benthic macroinvertebrate community structure and function in these low
order coastal California streams. An ongoing study on nutrient (P) spiraling in the
study streams (Harvey and Hill in prep) has shown streams are very non-
retentive, i.e. they leak phosphorus rapidly. So the physical, chemical, and
biological conditions to effectively take up phosphorus are absent with or without
carcasses. Nonetheless, pairwise comparisons pre and post-treatment did
reveal a signiﬁcant effect of augmenting both light and nutrients on mean
invertebrate biomass. These results suggest the importance of considering light
availability as a component of studies investigating management options that aim

to increase salmonid ﬁsh production through cascading food web pathways.

I am indebted to a countless number of people who have helped me along the
way. I am especially grateful to Dr. Richard W. Merritt, who despite receiving
letters and applications from more qualiﬁed and apt prospective students, was
both willing to give me the opportunity and persistent enough to see me through

it all. Thank you, Rico.

iv

ACKNOWLEDGEMENTS

The Green Diamond Resource Company provided access to their property
and logistical support. California Cooperative Fish Research Unit at Humboldt
State University provided additional logistical support. The Ofﬁce of Diversity and
Pluralism at Michigan State University provided some ﬁnancial support for ﬁeld
equipment. Moral support provided by fellow friends and students at both

Humboldt State University and Michigan State University.

TABLE OF CONTENTS

LIST OF TABLES ................................................................................................ vii
LIST OF FIGURES ............................................................................................. viii
CHAPTER 1

INTRODUCTION

Introduction ........................................................................................................... 1
Literature Cited ..................................................................................................... 5
CHAPTER 2

BENTHIC MACROINVERTEBRATE RESPONSE TO EXPERIMENTAL
MANIPULATION OF LIGHT AND NUTRIENTS IN SECOND ORDER STREAMS
OF NORTHERN CALIFORNIA

Introduction ........................................................................................................... 8
Methods .............................................................................................................. 1 0
Results ................................................................................................................ 14
Discussion ........................................................................................................... 19
Conclusion .......................................................................................................... 24
Literature Cited ................................................................................................... 45
CHAPTER 3

RED ALDER (Alnus rubra) LEAF PACK MASS LOSS AND
MACROINVERTEBRATE RESPONSE TO EXPERIMENTAL MANIPULATION
OF LIGHT AND NUTRIENTS IN SECOND ORDER STREAMS OF NORTHERN
CALIFORNIA

Introduction ......................................................................................................... 50
Methods .............................................................................................................. 51
Results ................................................................................................................ 55
Discussion ........................................................................................................... 59
Conclusions ........................................................................................................ 60
Literature Cited ................................................................................................... 77
APPENDIX .......................................................................................................... 81

vi

LIST OF TABLES

CHAPTER 2
Table 2.1. Study stream characteristics. * Denotes streams with carcass
addition ................................................................................................................ 26

Table 2.2. Checklist of taxa collected from cobble habitats within study reaches
of the Smith and Klamath River basins of Northern California. (a- premaipulation,
b- postmanipulation, c- open canopy, d- closed canopy, e- control, f- nutrient, 9—
light, h- Iight+nutrient, Sc- scraper, Gc- gathering collector, Fc— ﬁltering collector,
Sh- shredder, Pi- piercer, Pr— predator, Pa- parasite, A- Accidental Drifter, B-
Behavioral drifter) ............................................................................................... 27

Table 2.3. Mean P/R ratio analog, calculated from macroinvertebrate biomass by
sampling event and treatment. Ratios (>075) are autotrophic, (<0.75) are

heterotrophic. No statistically signiﬁcant differences ........................................... 33
Table 2.4. Percent of total biomass of top ﬁve taxa per sampling event .............. 34
CHAPTER 3

Table 3.1. Study stream characteristics. * Denotes streams with carcass

addition ................................................................................................................ 62

Table 3.2. Daily decay rates (k) of red alder leaf packs from experimental study
reaches of the Smith and Klamath River basins, northern California. (C= controls,
N= augmented nutrients, L= augmented light, L+N= augmented light and
nutrients). k values with same letters are not signiﬁcantly different from each
other, Fishers LSD (p =0.05) ................................................................................ 63

Table 3.3. Biomass (mg) and percent dominance of the most abundant taxa:
nutrient treatment (carcass, no carcass) by time (Day 15, Day 23, Day 30), from
Spring experimental red alder (Alnus rubra) leaf packs ........................ . .............. 64

Table 3.4. Biomass (mg) and percent dominance of the most abundant taxa in
(C) and (L) treatments by time (Day 15, Day 23, Day 30) from fall experimental
red alder (Alnus rubra) leaf packs ........................................................................ 65

Table 3.5. Checklist of taxa collected from experimental red alder (Alnus rubra)
leaf-packs from study reaches of the Smith and Klamath River basins of Northern
California. (a- Spring, b- Fall, Sc- Scraper, Gc— gathering collector, Fc- ﬁltering
collector, Sh- shredder, Pi- piercer, Pr- predator) ................................................ 66

vii

LIST OF FIGURES

CHAPTER 2
Figure 2.1. Study reaches within the lower Smith and Klamath River basins of

northern California. Fish symbol refer to streams that received carcass additions
(after WIlzbach et al. 2005) .................................................................................. 35

Figure 2.2. Mean macroinvertebrate biomass (: 1 SE.) by (a) sampling event.
Bars with same letters are not signiﬁcantly different from each other, Fishers LSD
(p = 0.05). And (b) sampling event by carcass interaction, March 2002 t = 2.19,
df=9, p = 0.05 ....................................................................................................... 36

Figure 2.3. Pairwise comparison of pre and post-treatment macroinvertebrate
biomass treatment means (1 1 SE). L+N signiﬁcant t = 3.08, df=150, p =0.0025

............................................................................................................................. 37

Figure 2.4. Mean percent Chironomidae by (a) the main effects of canopy type,
and sampling event, and by (b) interaction between canopy and sampling event
treatments. Bars with same letters are not signiﬁcantly different from each other,
Fishers LSD (p = 0.05) ......................................................................................... 38

Figure 2.5. (a) Mean Richness and (0) Mean Shannon-Weiner Diversity pre and
post-treatment (:I: 1 SE). Bars with same letters are not signiﬁcantly different
from each other (p < 0.05) ................................................................................... 39

Figure 2.6. Functional feeding group composition pre (October 2001) and pos-
treatment (October 2003) across all treatments. * Indicates signiﬁcantly different
pre and post-treatment (ANOVA, p < 0.05) .......................................................... 40

Figure 2.7. Interaction effect on percent Shredder (a) pre (October 2001) and
post-treatment (October 2003) within closed-canopy reaches, and (b) post-
treatment open versus closed-canopy reaches (1 1 SE). Bars with same letters

are not signiﬁcantly different from each other, Fishers LSD (p = 0.05) ................ 41

Figure 2.8. Mean CPOM / FPOM ratio (:l: 1 SE.) calculated from
macroinvertebrate biomass by (a) sampling event, (b) sampling event by carcass
streams, and (c) sampling event by no carcass streams. Bars with same letters
are not signiﬁcantly different from each other, Fishers LSD (p = 0.05) ................ 42

Figure 2.9. Sampling event effect on mean behavioral / accidental drift ratio (:1: 1

SE). Bars with same letters are not signiﬁcantly different from each other,
Fishers LSD (p = 0.05) ......................................................................................... 43

viii

Figure 2.10. Interaction effect (sampling event by carcass treatment) on mean
behavioral / accidental drift ratios (1 1 SE.) in (a) no carcass streams, and (b) in
March 2002 between carcass and no carcass streams. Bars with same letters
are not signiﬁcantly different from each other, Fishers LSD (p = 0.05) ................ 44

CHAPTER 3

Figure 3.1. Study reaches within the lower Smith and Klamath River basins of
northern California. Fish symbol refer to streams that received carcass additions
(after WIlzbach et al. 2005) .................................................................................. 71

Figure 3.2. Percent AFDM experimental red alder leaf pack remaining (+/- 1 SE.)
at day (2, 15, 23, and 30) in study reaches of the Smith and Klamath River
basins during a) spring and b) fall ........................................................................ 72

Figure 3.3. Spring a) dry mass and b) Shannon-Weiner diversity (H') (+/- 1 SE.)
of macroinvertebrates on experimental red alder leaf packs at days (15, 23 and
30) from streams of the Smith and Klamath River basins .................................... 73

Figure 3.4. Fall dry mass (+/- 1 SE.) of invertebrates on experimental red alder
leaf packs at days (15, 23 and 30) from a) streams and b) study reaches of the
Smith and Klamath River basins .......................................................................... 74

Figure 3.5. Fall Shannon-Weiner diversity (H') (+/- 1 SE.) of macroinvertebrates
on experimental red alder leaf packs at days (15, 23 and 30) from a) streams and
b) treatment study reaches of the Smith and Klamath River basins .................... 75

Figure 6. Fall Shredder biomass (+/- 1 SE.) on experimental red alder leaf packs
among study treatments in the Smith and Klamath River basins ........................ 76

ix

CHAPTER 1

INTRODUCTION

Northern California’s economy is dependent on its natural resources.
Much of the landscape has been designated for timber harvest. Del Norte, the
most northwestern county has almost equal acreage designated for national
forest (157,000) than it does for the forest industry (135,000), although timber
production has steadily been decreasing from 60,105,000 board feet in 1996 to
22,691,000 board feet in 2001. Humboldt County located just south of Del Norte,
has 262,000 acres designated as national forest and 608,000 acres in the forest
industry. Similarly, timber production has been steadily declining in Humboldt
County from 517,524,000 board feet in 1996 to 358,225,000 board feet in 2001
(California Department of Forestry Website).

As a result, previously harvested areas are now characterized by a dense
red alder (Alnus rubra) riparian canopy. Red alder is an early successional
species that quickly colonizes disturbed areas with exposed mineral soils. Vlﬁth
rapid growth rates of one meter or more in the ﬁrst year and maximum annual
growth of more than three meters a year by two to ﬁve year old seedlings
(Harrington and Curtis 1986), it quickly dominates riparian margins. However, red
alder is a relatively short-lived species maturing at 60 to 70 years with a
maximum age of about 100 years and has low commercial value. However, its
ecological value to streams has been the focus of studies in the Paciﬁc

Northwest. Red alder is capable of ﬁxing atmospheric nitrogen (N2) that can

result in increases in nitrogen availability both in the soil and in leaf litter that
could potentially beneﬁt the quality of allochthonous organic matter to adjacent
streams. For example, in an effort to determine how the distribution of
macroinvertebrates is affected by interstitial detritus quality and quantity, Culp
and Davies (1985) found that total macroinvertebrate densities and biomass
were greater in alder detritus than in no detritus and low hemlock detritus in the
main channel of Carnation Creek, Vancouver Island, British Columbia. In
addition, research on the effects of canopy type on benthic macroinvertebrate
and detritus export from headwater streams in southeast Alaska, has shown
streams with a red alder dominated young growth canopy exported signiﬁcantly
more macroinvertebrates than did streams with a conifer-dominated young
growth canopy (Piccolo and VIﬁpﬂi 2002).

In addition to the economic importance of timber in northern California,
there are numerous streams that are economically important for salmonid ﬁsh
production. The commercial value of samonids in California in 2001 was
$4,692,093 (Department of Fish and Game website) primarily from Chinook
salmon (Oncorhynchus tshawytscha) and steelhead (Oncorhynchus mykiss).
Coho salmon (Oncorhynchus kisutch) present in some watersheds in California
are currently listed as threatened, which reﬂects the decline in salmonid
populations in the Paciﬁc Northwest (Gresh et al. 2000). The importance of
salmonid ﬁsh to California’s economy has resulted a need for management

options that will likely lead to their increased production.

One current management option being considered to stimulate ﬁsh
productiOn is removal of riparian red alder. Removal of riparian canopy has led to
increases in periphyton production (Hansmann and Phinney 1973; Duncan and
Brusven 1985a; Feminella et al. 1989; Hetrick et al. 1998a) that enhances
benthic invertebrate production (Burns 1972; Newbold et al. 1980; Hawkins et al.
1982; Duncan and Brusven 1985b; Hetrick et al. 1998b). This enhanced I
invertebrate production may partly be due to the generally higher protein content
and digestibility of algae and algal-based detritus than most incoming terrestrial
plant matter (T riska et al. 1975). The change in the relative importance from
allochthonous organic matter sources on which small order streams depend
upon (Vannote et al. 1980) to autochthonous organic matter will result in changes
to the invertebrate community composition from primarily shredders and
detritivores to scrapers and gathering—collectors (Gregory et al. 1987). These
changes in community composition and greater production of benthic
invertebrates will result in greater food availability for ﬁsh by way of greater
invertebrate abundances or possibly through behavioral differences such as
invertebrate drift, ultimately leading to greater production of ﬁsh.

Another management option for increased ﬁsh production is fertilization of
streams. Harvey et al. (1998) showed that experimental additions of phosphorus
and nitrogen to an arctic tundra stream stimulated production at all trophic levels
relative to an unfertilized reach. Declining nutrient supplies historically derived
from carcasses of spawned salmon, known as marine derived nutrients, has

likely led to increased nutrient limitation in streams of the Paciﬁc Northwest

(Gresh et al. 2000). Fertilization of streams is believed to result in a positive
feedback mechanism whereby nutrients from salmon carcasses are incorporated
by stream autotrophs. The increased production of autotrophs would then result
in greater food availability for organisms in higher trophic levels, up to top
predators (salmonid ﬁsh). Greater production of ﬁsh would result in greater
spawner returns and therefore a greater nutrient subsidy of marine derived
nutrients.

In an effort to assess the efﬁcacy of these management options we
propose to experimentally manipulate light and nutrient levels in previously
harvested second order streams with a red alder riparian canopy in northern
California. Experimental manipulation is a logical approach to determining the
relative effects of increased light, nutrients, or both by contrasting manipulated
and controlled areas in natural streams. The objective of this study is to
document the effects of increased light and nutrients on coastal cutthroat (Salmo

clarkr) ﬁsh production by the food web pathways that support them.

LITERATURE CITED

Literature Cited

Burns,J.W. 1972. Some effects of logging and associated road construction on
northern California streams. Transactions of the American Fisheries
Society 101: 1—17.

Culp,J.M. and Davies,R.W. 1985. Responses of benthic macroinvertebrate
species to manipulation of interstitial detritus in Carnation Creek, British
Columbia. Canadian Journal of Fisheries and Aquatic Sciences 42: 139-
146.

Duncan,W.F.A. and Brusven,M.A. 1985a. Energy dynamics of three low-order
southeast Alaskan streams: Autochthonous production. Journal of
Freshwater Ecology. La Crosse 3: 155-166.

Feminella,J.W., M.E.Power, and V.H.Resh 1989. Periphyton responses to
invertebrate grazing and riparian canopy in three California coastal
streams. Freshwater Biology 22: 445-457.

Gregory S.V., G.A.Lamberti, D.C.Erman, K.V.Koski, M.L.Murphy, and J.R.Sedell.
1987. Inﬂuence of forest practices on aquatic production. In Streamside
management: forestry and ﬁshery interactions. Edited by E.O.Salo and
T.W.Cundy. Institute of Forest Resources, Contribution 57, University of
Washington AR-10, Seattle, WA pp. 233-255.

Gresh,T., Lichatowich,J., and Schoonmaker,P. 2000. An Estimation of Historic
and Current Levels of Salmon Production in the Northeast Paciﬁc
Ecosystem: Evidence of a Nutrient Deﬁcit in the Freshwater Systems of
the Paciﬁc Northwest. Fisheries 25: 15-21.

Hansmann,E.W. and Phinney,H.K. 1973. Effects of logging on periphyton in
coastal streams of Oregon. Ecology 54: 194499.

Harrington,C.A. and R.O.Curtis 1986. Height growth and site index curves for red
alder. USDA For Serv Res Pap PNW US Pac Northwest For Range Exp
Stn.

Harvey,C.J., Peterson,B.J., Bowden,W.B., Hershey,A.E., Miller,M.C.,
Deegan,L.A., and Finlay,J.C. 1998. Biological responses to fertilization of

Oksrukuyik Creek, a tundra stream. Journal of the North American
Benthological Society 17: 190-209.

Hawkins,C.P., Murphy,M.L., and Anderson,N.H. 1982. Effect of canopy,
substrate composition, and gradient on the structure of macroinvertebrate
communities in cascade range streams of Oregon. Ecology 63: 1840-
1856.

Hetrick,N.J., Brusven,M.A., Meehan,W.R., and Bjornn,T.C. 1998a. Changes in
solar input, water temperature, periphyton accumulation, and
allochthonous input and storage after canopy removal along two small
salmon streams in southeast Alaska. Transactions of the American
Fisheries Society 127: 859-875.

Hetrick,N.J., Brusven,M.A., Bjornn,T.C., Keith,R.M., and Meehan,W.R. 1998b.
Effects of canopy removal on invertebrates and diet of juvenile Coho
salmon in a small stream in southeast Alaska. Transactions of the
American Fisheries Society 127: 876-888.

Newbold,J.D., Erman,D.C., and Roby,K.B. 1980. Effects of logging on
macroinvertebrates in streams with and without buffer strips. Canadian
Journal of Fisheries and Aquatic Sciences. 37: 1076 -1085.

Piccolo,J.J. and WIpﬂi,M.S. 2002. Does red alder (Alnus rubra) in upland riparian
forests elevate macroinvertebrate and detritus export from headwater
streams to downstream habitats in southeastern Alaska? Canadian
Journal of Fisheries and Aquatic Sciences 59: 503-513.

Triska F.J., J.R.Sedell, and B.Buckley 1975. The processing of conifer and
hardwood leaves in two coniferous forest streams: ll Biochemical and
nutrient changes. Verh.lnt.Ver.Limnol. 19: 1628-1639.

Vannote,R.L., Minshall,G.W., Cummins,K.W., Sedell,J.R., and Cushing,C.E.
1980. The river continuum concept. Canadian Journal of Fisheries and
Aquatic Sciences. 37: 130-137.

CHAPTER 2
BENTHIC MACROINVERTEBRATE RESPONSE TO EXPERIMENTAL

MANIPULATION OF LIGHT AND NUTRIENTS IN SECOND ORDER STREAMS
OF NORTHERN CALIFORNIA

INTRODUCTION

Coastal waterbodies of the Paciﬁc Northwest are characterized as
oligotrophic (Bisson and Bilby 1998), particularly with respect to phosphorus
(Welsh et al. 1998). It has been proposed that in general, this condition has been
exacerbated due to declining salmonid stocks (Larkin and Slaney 1997) that
historically provided an estimated 95% of the N and P in those systems to a
marine derivation (Gresh et al. 2000), before nitrogen-ﬁxing red alder became the
dominant riparian tree. Fertilization has been proposed and used as a
management technique with the goal of reversing or ameliorating this trend. The
idea is that fertilization would lead to greater primary production and cascade up
the food web, ultimately increasing the number of salmonid spawners that will
return and perpetuate the cycle. There is evidence from southeast Alaska that
stream biota, at multiple trophic levels, assimilate these marine derived nutrients
(Chaloner et al. 2002), where the effects of enrichment have been evaluated. For
example, salmon carcass and carcass analog (salmon cakes - dried salmon
pellets) enrichment to stream tributaries in Alaska were shown to result in higher
mean condition, production and lipid content of cutthroat trout (Oncrhynchus
clam) than in un—enriched stream sections (WIpﬂi et al. 2004). Although, these

positive responses to enrichment have been documented, effects on stream

chemistry and biota have been inconsistent and context dependent (Janetski et
al. 2009; WIIzbach et al. 2005).

In addition to nutrient limitation, many of these coastal streams have
undergone a history of timber harvest of their riparian vegetation, which has led
to a change in their composition, to a dominance of hardwoods (particularly red
alder Alnus rubra) with youngest stands having the highest levels of canopy
cover (Russell 2009). This dense canopy cover, which limits primary production,
may further be problematic to salmonid production, as autotrophic pathways are
important to salmon growth in the spring and summer (Bilby and Bisson 1992;
WIlzbach et al. 2005). Removal of riparian canopy has led to increases in
periphyton production (Hansmann and Phinney1973; Duncan and
Brusven1985a; Feminella et al. 1989; Hetrick et al. 1998a) that enhances benthic
invertebrate abundances and biomass (Burns 1972; Newbold et al. 1980;
Hawkins et al. 1982; Duncan and Brusven 1985b, Nislow and Lowe 2006). This
enhanced invertebrate production may be due partly to the generally higher
protein content and digestibility of algae and algal-based detritus than most
incoming terrestrial plant matter (T riska et al. 1975). A change in the relative
importance from allochthonous organic matter sources upon which small order
streams depend (Vannote et al. 1980) to autochthonous organic matter, results in
changes to the invertebrate community functional composition from dominance
by shredders and gathering-collectors to scrapers (Gregory et al. 1987). These

changes in community composition of benthic invertebrates likely result in greater

food availability for ﬁsh by way of greater invertebrate abundances or possibly
through behavioral differences such as increases in invertebrate drift.

In an effort to determine the relative effects of increased light, nutrients, or
both, on macroinvertebrate communities, we augmented light (via canopy
removal) and nutrient levels (via salmon carcass additions) in previously
harvested second order forested streams with a predominantly red alder riparian

canopy in northern California.

METHODS

Study Area

The study was conducted in coastal northern California (Del Norte and
Humboldt counties) between October 2001 and October 2003. The area typically
experiences warm dry summers and cool wet winters, with average yearly
temperatures ranging from 5.5 - 165°C and average yearly rainfall ranging
between 170 to 2000m. Study reaches were established upstream of anadromy
on 2 tributaries of the Klamath (Tarup and Tectah) and 4 tributaries of the Smith
River (South Fork Rowdy, Savoy, Little Mill and Peacock) basins (Figure 2.1).
Study reaches on second to third order streams had similar catchment areas
(Table 2.1). Riparian vegetation consisted primarily (>80%) of red alder (Alnus
rubra); while surrounding vegetation included coastal redwood (Sequoia
sempervirens), Douglas-ﬁr (Pseudotsuga menziesir), Sitka spruce (Picea

sitchensis), big leaf maple (Acer macrophyllum), tan oak (Lithocarpus

10

densiﬂorus), and California bay laurel (Umbellulan’a califomica). Resident
salmonids were cutthroat trout (Oncorhynchus clarkr), and rainbow trout l

steelhead (Oncorhynchus mykiss) and hybrids (WIIzbach et al. 2005).

Experimental Design

In an effort to augment nutrients and light to study reaches, salmon
carcasses were added to streams, and riparian hardwoods were removed from
the riparian margin. The ﬁrst phase of the study followed a split-plot experimental
design, with carcass treatment (carcass, no carcass) as the whole-plot factor and
canopy treatment (open, closed) as the sub-plot factor. Each of the streams
consisted of two 100m study reaches separated by a 150-200m buffer. Three of
the six streams were randomly selected to receive carcass addition, and a study
reach within each stream was randomly selected to undergo canopy removal
(Table 2.1). The design resulted in three replicate study reaches of four
treatments; control (C), augmented nutrients (N), augmented light (L), and
augmented light and nutrients (L+N).

Carcass addition of Chinook salmon (Oncorhynchus tsawytscha)
carcasses to the three streams was conducted in January of both 2002 and
2003. Salmon carcasses for Tarup creek were procured from California
Department of Fish and WIldlife's Iron Gate Fish Hatchery on the Klamath River.
Carcasses for Little Mill and Peacock creeks were collected from the privately

owned Rowdy Creek Fish Hatchery and from spawned carcasses on the Smith

11

River. Carcasses were stored frozen until introduced at a level of approximately 1
kgom‘z. Carcasses were anchored throughout study reaches with rebar in areas
of slack water and near debris dams. Grab samples of water were collected in
2002 from each study reach at 2, 6, 15 and 22 weeks after carcass addition and
analyzed for multiple water chemistry parameters (WIlzbach et al. 2005).

The canopy treatment was carried out in December 2001 after leaf fall,
and involved falling red alder and other hardwoods from a 20m wide band on
each side of the stream for the 100m length of each randomly selected stream
reach. A 2-man team with a chainsaw and a winch felled the trees to minimize
disturbance and direct the cut trees away from the stream. Prior to canopy
treatment, the potential available sunlight reaching streambeds was estimated at
20m intervals of all study reaches using a Solar PathﬁnderTM. Photosynthetically
active radiation (PAR) was measured simultaneously in open and closed canopy
reaches of a stream using a Li-Cor® quantum sensor model Ll-19OS after
canopy removal. Stream temperatures were monitored using Stowaway®

TidbiTTM submersible data loggers (VVIlzbach et al. 2005).
Macroin vertebrate Sampling

Macroinvertebrates were sampled once pre-treatment in October 2001
and 4 times post-treatment (March and July 2002, and July and October 2003). A

total of three macroinvertebrate samples were collected from each study reach at

each sampling event. Individual samples were collected from cobble habitats

12

from randomly selected rifﬂes within a study reach. Other habitats were sampled,
but have not yet been analyzed. Individual samples were collected with a D-
frame aquatic net placed immediately downstream of agitated habitat for 30
seconds. Samples were rinsed through a 250-um sieve and transferred into
Whirl-Paks®, preserved in 80% ethanol, and returned to the lab for sorting under
a dissecting scope. Invertebrates were picked from samples, counted, measured
to the nearest millimeter, and identiﬁed mostly to generic level for lnsecta (except
for Chironomidae) using Merritt & Cummins (1996; Merritt et al. 2008).
Chironomidae were identiﬁed to subfamily and non-insect invertebrates were not
identiﬁed beyond order level. A total of 180 samples were collected and
analyzed.

Invertebrate biomass was calculated using INVERTCALC software (Merritt
et al. 2002). Richness was measured as mean number of taxa present. Diversity
was calculated based on invertebrate biomass using the Shannon-Weiner
diversity index (Hauer and Resh 1996), and percent Chironomidae was also
calculated based on invertebrate biomass. Macroinvertebrates were assigned a
functional group status (shredders, scrapers, ﬁltering-collectors, gathering-
collectors, and predators) according to Merritt and Cummins (1996; Merritt et al.
2008). Percent functional group and functional feeding group ratios (P/R,
CPOM/FPOM, Drift Food) were calculated based on biomass (Merritt et. al.
2002)

The response variables biomass, and functional feeding group ratios

(CPOM/FPOM and Drift Food) were analyzed by a split-split-plot ANOVA, with

13

carcass treatment (carcass, no carcass) as the whole-plot factor canopy
treatment (open, closed) as the sub-plot factor and sampling event (Oct 2001,
March 2002, July 2002, July 2003, and Oct 2003) as the sub-sub-plot factor.
Richness, diversity, percent functional group, and P/R ratio were analyzed
similarly, however, only Oct 2001 and Oct 2003 sampling events were evaluated,
for comparison pre and post manipulation during the same time of year.
Additionally, four pair-wise comparisons of macroinvertebrate biomass treatment
means, pre and post-treatment, were conducted using Bonferroni adjusted alpha
levels of 0.0125 per test (0.05/4). Response variable residuals were tested for
normality, and results were transformed where necessary. Means were
separated using Fishers LSD at p = 0.05. Although response variables were
transformed, they have been presented in untransformed fashion in graphs and

tables.

RESULTS

Carcass addition had no signiﬁcant effect on the following water chemistry
parameters tested, SIOz, nitrogen (TN, N05, and NH4*), or phosphorus (TP and
P043) (WIlzbach et al. 2005). Availability of potential sunlight did not differ among
streams or study reaches prior to canopy removal, however, canopy removal
often led to a tenfold increase in PAR, (Ambrose et al. 2004). Additionally,
canopy removal resulted in higher NH] levels in open-canopy reaches, however,

winter, summer and maximum weekly average temperatures did not differ

14

between open and closed study reaches. Small temperature increases (1 .5°C)
were noted at the downstream end of open-canopy reaches (WIIzbach et al.

2005)

Macroinvertebrate Biomass

There were no differences in mean macroinvertebrate biomass among
study reaches during pre-treatment sampling (all p z 0.60). Mean
macroinvertebrate biomass did not differ between streams with and without
carcasses (p = 0.64), or between open and closed-canopy reaches (p = 0.08).
However, biomass did differ among sampling dates with lowest levels at
pretreatment sampling (Figure 2.2a). Additionally, biomass was greater in
streams with carcasses than in streams without carcasses during March 2002
(Figure 2.2b). Results of pair-wise comparisons of macroinvertebrate biomass
treatment means, pre and post-treatment, show no difference within C, N, or L
treatments, and greater post-treatment biomass in L+N augmented streams (p =
0.0025) (Figure 2.3).

Mean °/o Chironomidae did not differ among study reaches during pre-
treatment sampling (all p > 0.21), or between streams with and without carcasses
(p = 0.07). Canopy treatment (p = 0.004) and sampling event (p < 0.0001) did
affect % Chironomidae, with closed-canopy and post-treatment samples having a
greater percentage of Chironomidae biomass (Figure 2.4a). Additionally, the

interaction of canopy and sampling event treatments also resulted in signiﬁcantly

15

greater (all p < 0.0001) % Chironomidae in closed-canopy reaches sampled in

October 2003 (Figure 2.4b).

Richness and Diversity

A total of 83 insect genera and three chironomid subfamilies were
collected from study reaches of the Smith and Klamath River basins of Northern
California, in addition to eight non-insect taxa (Table 2.2). Mean taxa richness did
not differ signiﬁcantly among study reaches during pre—treatment sampling (all p
2 0.14). Mean taxa richness did not differ between streams with and without
carcasses (p = 0.47), or between open and closed—canopy reaches (p = 0.50).
However, mean taxa richness was greater during the October 2001 (pre-
treatment) sampling event, than during October 2003 (p < 0.0001)(Figure 2.5a).
There were no signiﬁcant interaction effects on taxa richness. Shannon-Weiner
diversity did not differ signiﬁcantly between streams with and without carcasses
(p = 0.12), between open and closed-canopy reaches (p = 0.91), or among
sampling events (p = 0.32) (Figure 2.5b). Additionally, there were no signiﬁcant

interaction effects on diversity (all p > 0.06).

Functional Analysis

Percent functional group composition for scrapers (Sc), ﬁltering-collectors

(Fc), gathering-collectors (Go) and predators (Pr) did not differ between streams

16

with and without carcasses (all p > 0.19), or between open and closed-canopy
reaches (all p > 0.14). Additionally, there were no signiﬁcant interactions on
percent composition for these groups (all p > 0.05). However, there were
signiﬁcant differences on percent composition during pre and post-treatment
October sampling effects on these groups (all p < 0.05)(Figure 2.6). The
predators were the most dominant functional group present at both pre and post-
treatment sampling events, comprising about 40% of the total biomass.
Calineun'a (Perlidae) and Rhyacophila (Rhyacophilidae) were the dominant (Pr)
taxa; each had between 30-50% more biomass in the post-treatment samples.
Scraper biomass was overall greater in post-treatment samples. The snail Juga
(Pleuroceridae) was the dominant (Sc) taxon in both pre and post-treatment
samplings with three times as much biomass in the post-treatment sampling.
Similarly, ﬁltering-collector biomass was greater in post-treatment samples. The
(Fc) caddisﬂy Hydropsyche (Hydropsychidae) was the dominant taxa in both pre
and post-treatment samples with ten times more biomass in post-treatment
samples. Gathering-collector biomass was greater overall in pre-treatment
samples, with Oligochaeta being the dominant taxon with similar levels of
biomass in both pre and post-treatment samplings. There was greater biomass of
both Cinygmula (Heptageniidae) and Baetis (Baetidae) in post-treatment
samples. Percent shredder (Sh) composition did not differ between streams with
and without carcasses (p = 0.79), or between open and closed-canopy reaches
(p = 0.71), or between pre and post-treatment sampling events (p = 0.46) (Figure

2.6). However, there was a signiﬁcant interaction between sampling event and

17

canopy type (p = 0.008). There was a greater percentage of shredders in
October 2003 than in October 2001 within closed-canopy reaches (p = 0.03), as
a result of more Pteronarcys (Pteronarcyidae), Amphinemoura and Malenka
(Nemouridae) biomass in closed-canopy reaches in post-treatment samples than
in closed-canopy reaches in pre-treatment samples. There were also a greater
percentage of shredders within closed-canopy reaches than in open-canopy
reaches during October 2003 (p =0.02) (Figure 2.7), as a result of more
Pteronarcys and the Nemouridae stoneﬂies Zapada, Malenka and
Amphinemoura, in post-treatment closed-canopy reaches than in post-treatment
open-canopy reaches.

Analysis of mean P/R ratios resulted in no signiﬁcant main effects of
carcass (p = 0.63), canopy (p = 0.30), or sampling event (p = 0.60) treatments, in
addition to no signiﬁcant interactions (all p > 0.16)(Table 2.3). Similarly there was
no main effect of carcass and canopy treatments on CPOM/FPOM (p = 0.92 and
p = 0.49) or Drift Food (p = 0.33 and p = 0.33) ratios. Sampling event (p =
0.0035), and the interaction of sampling event and carcass treatment (all p <
0.05) led to signiﬁcant differences in CPOM/FPOM ratios (Figure 2.8). Similarly,
sampling event (p = 0.0008) (Figure 2.9), and the interaction of sampling event
and carcass treatment (p = 0.04) led to signiﬁcant differences in Drift Food ratios

(Figure 2.10).

18

DISCUSSION

Macroinvertebrate Biomass

Overall study design limitations (lower degrees of freedom at the
wholeplot level) led to greater signiﬁcant ﬁndings at the subplot (primarily
sampling event) and interaction levels of treatments in this study. Invertebrate
biomass varied signiﬁcantly among sampling events, potentially due to seasonal
differences in standing stock biomass. However, biomass was signiﬁcantly
greater at all sampling dates post-treatment, including the ﬁnal sampling event
that occurred in the same month as the pre-treatment sampling event. Sampling
events yielded similar results in taxonomic composition, according to Jaccard
coefﬁcients, no two sampling events were less than 76% similar. Additionally, the
top ﬁve dominant taxa comprised between 40-55% of the total biomass across
sampling dates (Table 2.4), suggesting all sampling dates were dominated
similarly by these same taxa. The stoneﬂy Calineuria (Perlidae) was present
across all sampling dates as the most biomass dominant organism, comprising
11-17% of the total biomass. A positive response by perlid stoneﬂies to carcass
additions and indirect uptake of nutrients occurring about two months after the
addition also was recorded in Washington (Claeson et al. 2006). This evidence
and the ﬁnding that the greatest percentage of salmon derived nutrients occurs in

macroinvertebrates three months after additions (Honea and Gara 2009), lends

19

support to our ﬁnding of greater invertebrate biomass in streams with carcass
addition than in streams without carcass addition in March 2002.

Analysis of invertebrate biomass using Bonferroni adjusted pair-wise
comparisons of treatment means pre versus post-manipulation reveals a gradual
pattern of increased biomass from C to N, and L reaches. Previous studies have
shown greater densities of Chironomidae and Ephemeroptera following nutrient
additions to experimental stream channels (Perrin and Richardson 1997; Kiffney
and Richardson 2001). This study, conducted in natural streams with a red alder
riparian canOpy failed to show a signiﬁcant increase in invertebrate biomass
subsequent to carcass additions. Carcass additions to streams with a red alder
riparian canopy have been shown not to affect stream water chemistry or quality
(Edmonds and Mikkelsen 2006; WIlzbach et al. 2005). Previous studies have
documented greater abundances of invertebrates in streams with clearcut
canopies (Hawkins et al. 1982), in addition to greater biomass of invertebrates in
open versus closed canopies (Hetrick et al. 1998b). Our study resulted in greater
amounts of invertebrate biomass in light-augmented treatment reaches, although
the difference was not signiﬁcant (p = 0.09). However, we did show positive
additive effects on invertebrate biomass through the augmentation of both light
and nutrients, suggesting the need to consider light as a limiting factor in marine
or salmon derived nutrient study designs.

Chironomidae abundance and biomass have been shown to have a
positive relationship with marine derived nutrients (Lessard and Merritt 2006). In

addition, chironomid density has been shown to increase in nutrient enriched

20

experimental streams (Kiffney and Richardson 2001). Our study showed no such
relationship to carcass addition, again a potential limitation of the study design.
Additionally, our study showed lower % Chironomidae biomass in open versus
closed canopy stream reaches, opposite of what is commonly described.
Chironomidae have been shown to increase three times after tree harvest
(German and Moring 1993), and low canopy cover has been shown to result in
increased numbers of Chironomidae larvae (Nislow and Lowe 2006). There was
a greater percentage of Chironomidae, in terms of biomass, during the post-

treatment sampling particularly in the closed canopy reaches.

Richness and Diversity

Invertebrate taxa richness and diversity did not respond signiﬁcantly to
experimental addition of salmon analogs (dried salmon pellets) in Idaho (Kohler
et al. 2008). However, invertebrate diversity has been shOwn to decline during
salmon spawning (Lessard and Merritt 2006), presumably the result of
disturbance during salmon spawning. We did not assess invertebrate diversity
during carcass addition. The difference in mean richness pre and post-treatment
in this study resulted from some rare taxa that appeared in between 1-8 of the
total number of samples (Moselia, Polycentropus, Hydatophylax, Ceratopsyche,

Rhantus).

21

Functional Analysis

A dominance of predators in study reaches suggests a need for a food
base with rapid turnover rates (Merritt et al. 2002), such as chironomid larvae,
whose turnover rates have been shown to decrease when in the presence of
nutrient enrichment (Ramirez and Pringle 2006). Greater scraper biomass in
post-treatment sampling was presumed the result of increased periphyton growth
expected after canopy removal (Hetrick et al. 1998a). The absence of signiﬁcant
differences in periphyton ash-free dry mass pre and post-manipulation in this
study (Ambrose et al. 2005) may have resulted from increased scraper feeding
pressure or scouring. Collectors have been implicated as important for the
transfer of marine or salmon derived nutrients through food webs (Chaloner and
WIpﬂi 2002) although in some cases Heptageniidae and Baetidae have been
shown to have greater abundances and biomass in control reaches of marine
derived nutrient studies (Lessard and Merritt 2006). The positive response by
Cinygmula and Baetis in this study likely reﬂect a lack of disturbance by salmon
migration, as our carcass introductions were experimental rather than natural.
Shredder biomass only differed at the interaction level (sampling event*canopy
treatment) of the study. Invertebrate samples from cobble habitats of closed-
canopy stream reaches had greater shredder biomass than the same habitat in
open-canopy reaches, likely because of the greater availability of food resource

in the form of red alder riparian litter.

22

Due to sample variability, there were no signiﬁcant differences in P/R
functional group ratios as was expected between open and closed-canopy
reaches. Mean P/R ratios by treatment and sampling event Show heterotrophy
dominated stream reaches, although autotrophy appeared to dominate during the
October 2003 sampling event across all treatments during leaf drop (Table 2.3).

CPOM/F POM functional group ratio varied by sampling event with March
2002 and July 2002 having the smallest ratios, as a result of low numbers of
large individuals (ex. Ptronarcys). The March sampling event was marked by a
small overall shredder biomass (75mg) versus >140mg at other sampling events,
while collector biomass was large in July 2002 (3900mg) versus < 2300mg at
other sampling events. Streams without carcass addition had more (ca. 100mg)
shredder biomass than streams with carcass additions, particularly during pre
and post-treatment October sampling events.

Drift Food functional group ratio was largest during the July 2002 sampling
event. The biomass of the mayﬂies Baetis and Acentrella (Baetidae) was 2-5
times greater during this sampling event resulting in twice as much behavioral
drifter biomass than other sampling events. In streams without carcass additions,
primarily Baetis (Baetidae), and to a lesser part Paraleptophlebia
(Leptophlebiidae), are responsible for the greater biomass of behavioral drifters
during March 2002, July 2002 and October 2003 when compared to the pre-
treatment sampling event. The difference in Drift Food functional ratio between

streams with and without carcasses in March 2002 was attributable to accidental

23

drifters such as Pteronarcys (Pteronarcyidae), Calineuria (Perlidae) and

Rhyacophila (Rhyacophilidae), in streams with carcasses.

Conclusion

Results of this study revealed no Signiﬁcant main effect of carcass addition
on benthic macroinvertebrate community structure and function in these low
order coastal California streams. However, invertebrate biomass was signiﬁcantly
greater in streams reaches 2-3 months following carcass additions, suggesting a
need to re-evaluate carcass addition effects on a more focused Shorter
timeframe and a larger spatial scale than in this study. A two year follow up study
to Wllzback et al. 2005 was conducted by Harvey and Wllzbach (in prep) in
which the design was to add no carcasses to the upper reaches (both canopy
treatments) and carcasses to all the down stream reaches (both canopy
treatments). Again, this two-year study showed the same ﬁsh results - no carcass
effect on ﬁsh growth. The lack of carcass addition effects may likely reﬂect
limitations of the study's experimental design, in which the subplot factors of
canopy removal and sampling date have greater power. In 2-3 months, the effect
would likely be largely downstream well out of the experimental area. An ongoing
study on nutrient (P) spiraling in the study streams (Harvey and Hill in prep) has
shown streams are very non-retentive, i.e. they leak phosphorus rapidly. So the
physical, chemical, and biological conditions to effectively take up phosphorus

are absent with or without carcasses.

24

Similarly, removal of riparian hardwoods did not lead to signiﬁcant main
effects in all but one response variable (% Chironomidae). PainNise comparisons
pre and post-treatment however did reveal a signiﬁcant effect of augmenting both
light and nutrients on mean invertebrate biomass. These results suggest the
importance of considering light availability as a component of studies
investigating management options that aim to increase salmonid ﬁsh production

through cascading food web pathways.

25

 

 

 

ow Ev 530.. >>..Nrmo¢NF \ z..3.vm°Pv £_Ew\ o.m 32%
E On 885 >>..m~.mov~:z..ormmo$ ES? 3 .. Exam so“. :58
N6 YN 826.. >>..m~.movwr \ 35.803 £_Ew\ 0m .. xooommn.
md RN 826.. 3:239.va \ ZRNNva cng\ v.m ____2 2::
3 RN 5&2 3.98.8, \ ZSENOS 58525 3 .. 9:3
we ad 526.. 5.9.303? \ 2:539st ﬁmEmv: mg :998.

p.320 :30 comma Ewmm \ ANExV Emobw
Agrcoiﬁw Rocco :30 282654 33:23 moi EmEcono

 

.5363 3880 5:5 9:35 3550 .. .mozmtouomcmco Emobm >35 .FN p.98.

26

Table 2.2. Checklist of taxa collected from cobble habitats within study
reaches of the Smith and Klamath River basins of Northern California.

(a- premaipulation, b- postrnanipulation, c- open canopy, d- closed
canopy, e- control, f- nutrient, g- light, h- Iight+nutrient, Sc- scraper,

Gc- gathering collector, Fc- ﬁltering collector, Sh- shredder, Pi- Piercer, Pr-
predator, Pa- parasite, A- Accidental Drifter, B- Behavioral drifter)

 

 

Drift
Family Genus Presence FFG type
Ephemeroptera

Ameletidae Ameletus a b c d e f g h Gc B
Baetidae Acentrella a b c d e f g h Gc B
Baetis abcdefgh Gc B

Ephemerellidae Attenella a b c d e g h Gc A
Caudatella a c d e Go A

Drunella a b c d e f g h Sc A

Ephemerella a b d f Go A

Senate/Ia a b c d e f g h Go A

Timpanoga b c g h Go A

Heptageniidae Cinygma a b c d f g h Sc A
Cinygmula a b c d e f g h Go A

Epeorus a b c d e f g h Gc A

Heptagenia a b c d f 9 SC A

lronodes a b c d e f g h Sc A

Rhithrogena a b c d e f g h Gc A

Leptophlebiidae Paraleptophlebia a b c d e f g h Gc B

Plecoptera

27

Table 2.2 (cont)

 

(D

(OCOOCQ

Sh
Gc
Pr
Pr
Sh
Sh
Sh
Sh
Sh
Sh
Sh
Sh
Pr
Pr
Pr
Pr
Pr

Sh

>>>>>>>>>>>>>>>>>>

Capniidae

Chloroperlidae Kathroperta
Suwallia
Sweltsa

Leuctridae Despaxia
Mose/la
Pedomyia

Nemouridae Amphinemura
Malenka
Nemoura
Zapada

Peltoperlidae Soliperta

Perlidae Calineuria
Doroneuria
Hesperoperla

Perlodidae lsoperta
Osobenus

Pteronarcyidae Pteronarcys

Odonata
Gomphidae
Megaloptera
Corydalidae Oroherrnes

Table 2.2 (cont)

 

Sialidae
Trichoptera
Apatanidae

Brachycentridae

Calamoceratidae
Glossosomatidae

Hydropsychidae

Hydroptilidae
Lepidostomatidae
Leptoceridae

Limnephilidae

Philopotamidae
Polycentropodidae
Rhyacophilidae
Sericostomatidae

Uenoidae

Sialis

Apatania
Brachycentrus
Micrasema
Heteroplectron
Glossosoma
Ceratopsyche
Hydropsyche
Parapsyche
Ochrotn‘chia

Lepidostoma

Cryptochia
Ecclisomyia

H ydatoph ylax
Wonnaldia
Polycentropus
Rh yacophila
Gumaga
Neophylax

Neothremma

29

O'

U' 0'

U" U' 5' U' U'

U'

Pr

Sc
Fc
Sh
Sh
Sc
Fc
Fc
Fc
Gc
Sh
Sc
Sh
Gc
Sh
Fc
Pr
Pr
Sh
Sc
Sc

>>>>>m>>>>>>>>>>>>>>

Table 2.2 (cont.)

 

Hymenoptera
Scelionidae
Coleoptera

Dytiscidae

Elmidae

Hydrochidae
Hydrophilidae

Psephenidae

Diptera

Ceratopogonidae

Chironomidae

Laccophilus
Oreodytes
Rhantus
Ampumixis
Cleptelmis
Dubiraphia
Lara
Narpus
Optioservus
Rhizelmis
Zaitzevia
Hydrochus
Paracymus

Acneus

Atn'chopogon
Orthocladinae

Tanypodinae

30

Pa

Pr
Pr
Pr
Gc
Gc
Gc
Sh
Gc
Sc
Sc
Sc
Sh
Gc
Sc

Pr
Pr
Gc

Pr

>>>>>>>>>>>>>>

>CD>>

Table 2.2 (cont)

 

Tan ytarsini
Dixidae Dixa
Empididae Chelifera

Clinocera

Hemerodromia

Pelecorhynchidae Glutops-

Psychodidae
Maun'na
Simuliidae Simulium
Stratiomyidae Nemotelus
Thaumaleidae
Tipulidae Antocha
Dicranota
Hexatoma
Pedicia
Non-Insect
Collembola
Turbellaria
Oligochaeta
Hydracarina
Copepoda
Ostracoda

31

U U U U

U U U

U U U

0.0.0.

COC0¢0¢0¢0

(00(0

F 0
Go
Pr
Pr
Pr
Pr
Gc
Sc
Fc
Gc
Sc
Gc
Pr
Pr
Pr

Gc
Pr
Gc
Pr
Fc

Gc

>>>>w>w>w>>>>wm

>UJ>>>>

Table 2.2 (cont)

 

Amphipoda

Gammaridae a b c d e f g h Sh B
Gastropoda

Pleuroceridae Juga a b c d e f g h Sc A

 

32

Table 2.3. Mean P/R ratio analog, calculated from macroinvertebrate biomass
by sampling event and treatment. Ratios (>0.75) are autotrophic, (<0.75)
heterotrophic. No statistically signiﬁcant differences.

 

Sampling Event
Treatment (OCT01) (MARCH02) (JULY02) (JULY03) (OCT03)

 

C 0.27 0.46 0.36 0.24 0.76
N 0.41 0.52 0.58 0.61 0.70
L 0.50 0.46 0.56 0.37 0.83
L+N 0.24 0.32 0.33 8.33 1.64

 

33

Table 2.4. Percent of total biomass of top ﬁve taxa per sampling event

 

FFG (OCT01) (MARCH02) (JULY02) (JULY03) (OCT03)

 

 

Calineuria Pr 14.88 17.08 17.40 11.26 13.41
Oligochaeta Gc 8.54 6.27 7.41 6.24
Rhyacophila Pr 9.15 10.07 7.78

Pteronarcys Sh 14.00 1 1.62
Turbellaria Ge 7. 35 5.79
Hesperoperta Pr 6. 89 6.42

Juga Sc 6.91 8.30
lronodes Sc 8. 05

Cinygmula Gc 8.04
Drunella Sc 6.1 2

Hydropsyche Fc 4.75
Narpus Gc 5.70

Percent Dominance 55.48 44.41 42.45 40.30 46.82

 

34

 

LOWER SMITH RIVER

z)—

 

 

LOWER KLAMATH RIVER

   

 

 

 

Figure 2.1. Study reaches within the lower Smith and Klamath River Basins of
northern California (after VIﬁlzbach et al. 2005).

35

(II)
II
D

 

soot (a)

 

250{

 

 

2002

 

l-———l
l———-l

150;

 

I——I

 

 

 

 

 

 

 

 

 

 

 

l 1 I I

 

(b) I Carcass
D No Carcass

Mean Macroinvertebrate Biomass (mg) +/- 1 SE.

 

 

 

 

 

 

O_ l I l l l
PRE(OCT01) (MARCH02) (JULY02) (JULY03) (OCT03)
Sampling Event

Figure 2.2. Mean macroinvertebrate biomass (:l: 1 SE.) by (a) sampling event.
Bars with same letters are not signiﬁcantly different from each other, Fishers LSD
(p = 0.05). And (b) sampling event by carcass interaction, March 2002 t = 2.19,
dh9p=005

36

 

.5
O
O

 

- Pre
'3 Post

N OD 0)
01 8 01
O O
IIILJIILJJAIJLA

2005

 

-L
U"
(I:

J J

 

100g

0:
‘5’

Mean Macroinvertebrate Biomass (mg) +/- 1 SE.

 

 

 

O
I I

 

 

 

 

 

 

 

   

L+N
Treatment

F lgure 2.3. Painrvise comparison of pre and post-treatment macroinvertebrate
biomass treatment means (1 1 SE). L+N signiﬁcant t = 3.08, df=150, p =0.0025.

37

 

(a)

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

0.1
‘T : l T
I I“. l
0 2
N .
.9 0.5:
g z B A B A
c n-
O V I I I I
E A Open Closed (OCT01) (come)
0 v-
‘c‘ «(hi I
0 <
2 .
m 5‘
n. : i
c .
8 4;
5 :
3‘:
2;
: 1L
' T
14 J.
1 B B B A
0 I
(OCT01)'Open ' (OCT01)’Closed (OCT03)"Open ' (OCTO3)'Closed

Figure 2.4. Mean percent Chironomidae by (a) the main effects of canopy type,
and sampling event, and by (b) interaction between canopy and sampling event
treatments. Bars with same letters are not signiﬁcantly different from each other,
Fishers LSD (p = 0.05).

38

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

i (a)
404; T
' 4 .L
LIJ :
(0' 35*.
- : 1-
W i
8 251
C
8 i
E 20?
C :
3 15‘.
2 :
10a
5% A B
u! o ‘ .
a) 3
T.- ‘ (b)
7
r‘ 2.5- I
at, T .L
g 1
9 2—
d) 1
.2 I
0 1
E 1.5-
.E i
o
3, :
8 I“.
C
C
2 i
a) 0.5
c i A A
a: .
I” .
E 0 I
PRE(OCT01) POST(OCT03)

Figure 2.5. (a) Mean Richness and (c) Mean Shannon-Weiner Diversity pre and
post-treatment (: 1 SE). Bars with same letters are not signiﬁcantly different
from each other (p < 0.05).

39

 

6%

24%
46%

7%

   

PRE(OCT01) POST(OCT03)

I Shredder |:| Filtering Collector" E Predator"
I Scraper* Gathering Collector*

Figure 2.6. Functional feeding group composition pre (October 2001) and pos-
treatment (October 2003) across all treatments. * Indicates signiﬁcantly different
pre and post-treatment (ANOVA, p < 0.05).

40

 

(a)

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Hi
(0
‘7 A
T .
S 0 I
3 PRE(OCT01 ) POST(OCT03)
E “i (bl
Y}, 10—
c -I
:3. .
33 i
c 8‘
(U .
o
2 -I

6-

4-

. -L
2-
j A B
0 l
Closed Canopy Open Canopy

Figure 2.7. Interaction effect on percent Shredder (a) pre (October 2001) and
post-treatment (October 2003) within closed-canopy reaches, and (b) post-
treatment open versus closed-canopy reaches (:t: 1 SE). Bars with same letters
are not signiﬁcantly different from each other, Fishers LSD (p = 0.05).

41

 

0.18: I (a)
0.16—

 

 

 

9.0
R33
III
l——-—r——I
—I
I,—

 

 

 

 

 

 

 

 

(b)

 

 

I
I

B Fri—i i—Iii—I A AB

 

 

O
I

Mean CPOM / FPOM Ratio +/- 1 SE.

 

 

 

 

 

 

 

 

 

 

025—; I (C)
i l

 

 

 

 

 

 

 

 

0.05—f
,3 A I I B I . I B35 I I Q, A
(OCTOI) (MARCH02) (JULY02) (JULY03) (OCT03)

Figure 2.8. Mean CPOM / F POM ratio (a: 1 SE.) calculated from
macroinvertebrate biomass by (a) sampling event, (b) sampling event by carcass
streams, and (c) sampling event by no carcass streams. Bars with same letters
are not signiﬁcantly different from each other, Fishers LSD (p = 0.05).

42

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

UJI

a, 0.35 4

T 0.3; T

.9 : l

g :

D 2

g 02—: l T
0 4 T

u . l
-- 015- 1

§ : T

x : .L

73 0.1 1

.9 ~

> :

g 0.05-

m 3 B B A B B
c a

8 0 I I I r

5 (OCT01) (MARCH02) (JULY02) (JULY03) (OCT03)

Figure 2.9. Sampling event effect on mean behavioral / accidental drift ratio (z 1

SE). Bars with same letters are not signiﬁcantly different from each other,
Fishers LSD (p = 0.05).

43

 

0.45

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

(a)
0.4? “i

0.355
LLI I
m. 0.345 I .-
‘,' 0.25:
r i l T
.9 02—;
2 I
E 015—:
9 0.1 -: .11:
9 :
é 0.05—j B A A A
'8 o ‘ I I I
E (OCT01) (MARCH02) (JULY02) (OCT03)
- 0.35 .
g : (b)
'5 03-: T
(D e
c -I
3 0.255 1
§ 02-? "I"
E 0.15% .I.

0.1 —
0.05—f A B
0 i ,

 

No Carcass Carcass

Figure 2.10. Interaction effect (sampling event by carcass treatment) on mean
behavioral / accidental drift ratios (s: 1 SE.) in (a) no carcass streams, and (b) in
March 2002 between carcass and no carcass streams. Bars with same letters
are not signiﬁcantly different from each other, Fishers LSD (p = 0.05).

44

 

LITERATURE CITED

45

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increased light and salmon carcass introduction in northern California
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Bilby, RE. and PA. Bisson 1992. Allochthonous versus autochthonous organic-
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Bisson, PA. and RE. Bilby. 1998. Organic matter and trophic dynamics. In River
ecology and management: lessons from the Paciﬁc coastal ecoregion.
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Burns, J.W. 1972. Some effects of logging and associated road construction on
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Chaloner, D.T., Martin, K.M., VIﬁpﬂi, M.S., Ostrom, PH. and GA. Lamberti. 2002.
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Chaloner, UT and MS. Viﬁpﬂi. 2002. Inﬂuence of decomposing Paciﬁc salmon
carcasses on macroinvertebrate growth and standing stock in
southeastern Alaska streams. Journal of The North American
Benthological Society. 21(3): 430-442.

Claeson, S.M., Li, J.L., Compton, J.E and PA. Bisson. 2006. Response of
nutrients, bioﬁlm, and benthic insects to salmon carcass addition.
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Duncan, W.F.A. and MA. Brusven. 1985a. Energy dynamics of three low-order
southeast Alaskan streams: Autochthonous production. Journal of
Freshwater Ecology. La Crosse 3: 155-166.

Duncan, W.F.A. and MA. Brusven. 1985b. Benthic macroinvertebrates in logged
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Invertebrate Biology. 4: 125-132.

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Edmonds, R.L. and K. Mikkelsen. 2006. Inﬂuence of salmon carcass placement
in red alder riparian areas on stream chemistry in lowland western
Washington. North American Journal of Fisheries Management. 26(3):
551-558.

Feminella, J.W., Power, ME. and V.H. Resh. 1989. Periphyton responses to
invertebrate grazing and riparian canopy in three California coastal
streams. Freshwater Biology 22: 445-457.

Garman, G. and J. Moring. 1993. Diet and annual production of 2 boreal river
ﬁshes following clear-cut logging. Environmental Biology of Fishes. 36(3):
301 -31 1 .

Gregory S.V., Lamberti, G.A., Errnan, D.C., Koski, K.V., Murphy, ML. and JR.
Sedell. 1987. Inﬂuence of forest practices on aquatic production. In
Streamside management: forestry and ﬁshery interactions. Edited by ED.
Salo and T.W. Cundy. Institute of Forest Resources, Contribution 57,
University of Washington AR-10, Seattle, WA pp. 233-255.

Gresh, T., Lichatowich, J. and P. Schoonmaker 2000. An estimation of historic
and current levels of salmon production in the northeast Paciﬁc
ecosystem: evidence of a nutrient deﬁcit in the freshwater systems of the
Paciﬁc Northwest. Fisheries. 25(1): 15—21.

Hansmann, E.W. and H. K. Phinney. 1973. Effects of logging on periphyton in
coastal streams of Oregon. Ecology 54: 194-199.

Hauer, F .R. and V.H. Resh. 1996. Benthic Macroinvertebrates. In Methods In
Stream Ecology. Edited by PR. Hauer and GA. Lamberti. San Diego,
Academic Press: 339-370.

Hawkins, C.P., Murphy, ML. and NH. Anderson. 1982. Effect of canopy,
substrate composition, and gradient on the structure of macroinvertebrate

communities in cascade range streams of Oregon. Ecology. 63: 1840-
1856.

Hetrick, N.J., Brusven, M.A., Meehan, W.R., and TC Bjornn. 1998a. Changes in
solar input, water temperature, periphyton accumulation, and
allochthonous input and storage after canopy removal along two small
salmon streams in Southeast Alaska. Transactions of the American
Fisheries Society 127: 859-875.

Hetrick, N.J., Brusven, M.A., Bjornn, T.C., Keith, RM. and W.R. Meehan. 1998b.
Effects Of canopy removal on invertebrates and diet of juvenile Coho

47

salmon in a small stream in southeast Alaska. Transactions of the
American Fisheries Society. 127(6): 876-888.

Honea, J.M. and RI. Gara. 2009. Macroinvertebrate community dynamics:
strong negative response to salmon redd construction and weak response
to salmon-derived nutrient uptake. Journal of the North American
Benthological Society. 28(1): 207-219.

Janetski, D.J., Chaloner, D.T., Tiegs, SD. and GA. Lamberti. 2009. Paciﬁc
salmon effects on stream ecosystems: a quantitative synthesis.
Oecologia. 159: 583-595.

Kiffney, P. and J.S. Richardson 2001. Interactions among nutrients, periphyton,
and invertebrate and vertebrate grazers in experimental channels. Copeia.
2: 422-429.

Kohler, K., Rugenski, A. and D. Taki. 2008. Stream food web response to salmon
carcass analogue addition in two central Idaho, U.S.A. streams.
Freshwater Biology. 53: 446-460.

Larkin, GA. and PA. Slaney. 1997. Implications of trends in marine-derived
nutrient inﬂux to south coastal British Columbia salmonid production.
Fisheries. 22(11): 16-24.

Lessard, J. and R. Merritt. 2006. Inﬂuence of marine-derived nutrients from
spawning salmon on aquatic insect communities in southeast Alaskan
streams. Oikos. 1 13(2): 334-343.

Merritt, R.W. and K.W. Cummins. 1996. Introduction to The Aquatic Insects of
North America. Dubuque, Kendall/Hunt.

Merritt, R.W., Cummins, K.W. and MB. Berg. 2008. Introduction to The Aquatic
Insects of North America. Dubuque, Kendall/Hunt.

Merritt, R.W., Cummins, K.W., Berg, M.B., Novak, J.A., Higgins, M.J., Wessell,
K.J. and J.L. Lessard. 2002. Development and application of a
macroinvertebrate functional-group approach in the bioassessment of

remnant river oxbows in southwest Florida. Journal of the North American
Benthological Society. 21(2): 290-310.

Newbold, J.D., Erman, DC, and KB. Roby 1980. Effects of logging on

macroinvertebrates in streams with and without buffer strips. Canadian
Journal of Fisheries and Aquatic Sciences. 37: 1076 -1085.

48

Nislow, K.H. and W.H. Lowe. 2006. Inﬂuences of logging history and riparian
forest characteristics on macroinvertebrates and brook trout (Salve/inus
fontinalis) in headwater streams (New Hampshire, USA). Freshwater
Biology. 51(2): 388-397.

Perrin, C. and J. Richardson. 1997. N and P limitation of benthos abundance in
the Nechako River, British Columbia. Canadian Journal of Fisheries and
Aquatic Sciences. 54(11): 2574-2583.

Ramirez, A. and CM. Pringle. 2006. Fast growth and turnover of chironomid
assemblages in response to stream phosphorus levels in a tropical
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Russell, W. 2009. The inﬂuence of timber harvest on the structure and
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Triska, F .J., Sedell, JR. and B. Buckley. 1975. The processing of conifer and
hardwood leaves in two coniferous forest streams: II Biochemical and
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1980. The river continuum concept. Canadian Journal of Fisheries and
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R. Naiman and R. Bilby. New York, Springer: 69-94.

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riparian canopy opening and salmon carcass addition on the abundance
and growth of resident salmonids. Canadian Journal of Fisheries and
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salmon-based food webs: contrasting effects of salmon carcass and
salmon carcass analog additions on stream-resident salmonids.
Transactions of the American Fisheries Society. 133: 1440-1454.

49

CHAPTER 3
RED ALDER (Alnus rubra) LEAF PACK MASS LOSS AND
MACROINVERTEBRATE RESPONSE TO EXPERIMENTAL MANIPULATION

OF LIGHT AND NUTRIENTS IN SECOND ORDER STREAMS OF NORTHERN
CALIFORNIA.

INTRODUCTION

Leaf litter is a primary energy source for streams and rivers, particularly
temperate headwater streams with a deciduous riparian canopy (Peterson and
Cummins 1974). Low order and headwater streams heavily shaded by a riparian
canopy are dependent upon these allochthonous sources of energy, as a heavily
shaded canopy serves to limit available sunlight to the streambed and therefore
limits primary production. Macroinvertebrate shredders are adapted to take
advantage of the litter resource, and convert it from coarse particulate organic
matter (CPOM) to ﬁne particulate organic matter (F POM) that is used by ﬁltering
and gathering collectors downstream (Merritt and Cummins 1996; Merritt et al.
2008; Vannote et al. 1980). This breakdown of leaf litter is an important process
in streams (Benﬁeld 1996), because it involves the conversion of plant biomass
to the animal biomass of higher trophic levels.

Leaf litter mass loss rates have been shown to vary depending on
microbial conditioning (Crowl et al. 2006), leaf species (Peterson and Cummins
1974), leaf species assemblages (Kominoski and Pringle et al. 2007), nutrient
enrichment, and riparian condition. Nutrient enrichment has resulted in mixed

effects on leaf litter mass loss, depending on the amount of nutrient enrichment

50

(Knorr et al. 2005). Studies have reported no effect on leaf mass loss (Ferreira et
al. 2007), as well as no effect on leaf pack invertebrate richness or abundance
(Ferreira et al 2006). Conversely, some studies have shown a positive effect on
leaf mass loss from nutrient enrichment (Yanai 2005; Paul 2006), and yet others
have reported suppression of leaf mass loss as a result of nutrient enrichment
(Zhang 2003). Riparian condition (light intensity) has similarly yielded mixed
results with respect to litter mass loss. For example, lshikowa et al. (2007)
reported effects of clear-cutting on leaf mass loss to be species speciﬁc, while

F ranken et al. (2005) reported no effect of light intensity on leaf mass loss, and
Bates et al. (2007) reported 37% greater leaf mass loss in streams with cut
riparian vegetation when compared to those with uncut riparian vegetation. It is
therefore important to determine and document the effects of natural stream
manipulations on organic matter processing dynamics. Two manipulations
proposed to stimulate stream productivity and ultimately salmonid growth and
abundance are stream fertilization by way of carcass addition and increasing light

by way of canopy removal.

METHODS

Study Area

This study was conducted in coastal northern California (Del Norte and

Humboldt counties) between October 2001 and October 2002. The area typically

51

experiences warm dry summers and cool wet winters, with average yearly air
temperatures ranging from 5.5 - 165°C and average yearly rainfall ranging
between 170 to 200cm. Study reaches were established upstream of anadromy
on 2 tributaries of the Klamath (Tarup and Tectah) and 4 tributaries of the Smith
River (South Fork Rowdy, Savoy, Little Mill and Peacock) basins (Figure 3.1).
Study reaches on second to third order streams had similar catchment areas
(Table 3.1). Riparian vegetation consisted primarily (>80%) of red alder (Alnus
rubra); while surrounding vegetation included coastal redwood (Sequoia
sempervirens), Douglas-ﬁr (Pseudotsuga menziesir), Sitka spruce (Picea
sitchensis), big leaf maple (Acer macrophyllum), tan oak (Lithocarpus
densiﬂorus), and California bay laurel (Umbellularia califomica). Resident
salmonids were cutthroat trout (Oncorhynchus clarkr), and rainbow trout /

steelhead (Oncorhynchus mykiss) and hybrids (Wllzbach et al. 2005).

Experimental Design

In an effort to augment nutrients and light reaching study reaches, salmon
carcasses were added to streams, and riparian hardwoods were removed from
the riparian margin. The study followed a split-plot experimental design, with
carcass treatment (carcass, no carcass) as the whole-plot factor and canopy
treatment (open, closed) as the sub-plot factor. Each of the streams consisted of
two 100m study reaches separated by a 150-200m buffer. Three of the six

streams were randomly selected to receive carcass addition, and a study reach

52

within each stream was randomly selected to undergo canopy removal (Table
3.1). The design resulted in three replicate study reaches of four treatments;
control (C), augmented nutrients (N), augmented light (L), and augmented light
and nutrients (L+N).

Addition of Chinook salmon (Oncorhynchus tsawytscha) carcasses to the
three streams was conducted in January 2002. Salmon carcasses for Tarup
creek were procured from California Department of Fish and Wildlife's Iron Gate
Fish Hatchery on the Klamath River. Carcasses for Little Mill and Peacock creeks
were collected from the privately owned Rowdy Creek Fish Hatchery and from
spawned carcasses on the Smith River. Carcasses were stored frozen until
introduced at approximately 1 kg-m'z. Carcasses were anchored throughout
study reaches with rebar in areas of slack water and near wood debris dams.
Grab samples of water were collected in 2002 from each study reach at 2, 6, 15
and 22 weeks after carcass addition and analyzed for multiple water chemistry
parameters (WIlzbach et al. 2005).

The canopy treatment was carried out in December 2001 after leaf fall,
and involved falling red alder and other hardwoods from a 20m wide band on
each side of the stream for the 100m length of each randomly selected stream
reach. A 2-man team with a chainsaw and a winch felled the trees to minimize
disturbance and direct the cut trees away from the stream. Prior to canopy
treatment, the potential available sunlight reaching streambeds was estimated at
20m intervals along all study reaches using a Solar PathﬁnderTM.

Photosynthetically active radiation (PAR) was measured simultaneously in open

53

and closed canopy reaches of a stream using a Li-Cor® quantum sensor model
Ll-1908 after canopy removal. Stream temperatures were monitored using

Stowaway® Tidbil'TM submersible data loggers (WIlzbach et al. 2005).

Leaf Packs

Individual leaf litter packs were constructed using red alder leaves at abscission.
Leaves were air-dried, weighed to 5-g packs, rewetted, loosely fastened together
with a quilting gun and tethered two to a brick for introduction into streams. A
total of 720 leaf packs (60 per stream reach) were constructed and deployed 60
per stream reach in both late spring (May-June) and early fall (August-
September) 2002. Leaf packs were collected at day-2 (10 leaching packs), day-
15 (5 packs), day-23 (5 packs), and day-30 (10 packs), in addition to 30 initial
packs to account for handling loss. Handling loss packs were subjected to all
conditions, however, time in the water was approximately 5 seconds. Individual
packs were removed from bricks, placed in a zip-lockTM bag, and transported to
the lab in an ice chest. In the lab, leaf packs were rinsed with de—ionized water
and air-dried to a constant mass. After weighing, a sub-sample of each leaf pack
was collected for ash—free dry mass (AF DM) determination. Percent AF DM leaf
pack remaining and decay coefﬁcients (k) were computed following Benﬁeld
(1996). Leaf pack-associated macroinvertebrates were collected from day-15,
day-23 and day-30 samples and preserved in 70% ethanol for later identiﬁcation

and enumeration. Invertebrate biomass was calculated using INVERTCALC

54

software (Merritt et al. 2002), and richness was measured as number of taxa
present. Invertebrate diversity was calculated based on invertebrate biomass
using the Shannon-Weiner diversity index (Hauer and Resh 1996, Hauer and
Resh 2007). Invertebrates were assigned to a shredder, scraper, ﬁltering-
collector, gathering-collector or predator functional group status (Merritt and
Cummins 1996; Merritt et al. 2008). Percent of each functional group, shredder
biomass and shredder diversity were also calculated based on biomass (Merritt
et al. 2002). Shredder richness was measured as number of taxa present.
Differences in decay coefﬁcients (k) among experimental treatments (C,
N, L, L+N) as well as invertebrate response variables were assessed by analysis
of covariance (ANCOVA) with time (days) as the oovariate (Barlocher 2005).
Initial values at day_0 were excluded from analysis. Response variable residuals
were tested for normality, and results were transformed where necessary. Means
were separated using Fishers LSD at p = 0.05. Although response variables were

transformed, they are presented in untransformed fashion in ﬁgures and tables.

RESULTS

Water Chemstry / PAR / Temperature

Carcass addition had no signiﬁcant effect on the following water chemistry
parameters tested, SiOz, nitrogen (TN, N03', and NH4‘), or phosphorus (TP and

P043) (Wllzbach et al. 2005). Availability of potential sunlight did not differ among

55

streams or study reaches prior to canopy removal, however, canopy removal
often led to a tenfold increase in PAR, (Ambrose et al. 2004). Additionally,
canopy removal resulted in higher NH4+ levels in open-canopy reaches, however,
winter, summer and maximum weekly average temperatures did not differ
between open and closed study reaches. Small temperature increases (1 .5°C)
were noted at the downstream end of open-canopy reaches (Wllzbach et al.

2005).

% AFDM remaining / Decay coefﬁcients (k)

Leaf mass loss was faster in fall the than in the spring 2002. Leaf packs in
the control stream reaches had the greatest % AFDM remaining at the end of
both spring (55% +/-1.7) and fall (29% +/-1.9) experiments. Leaf packs in the (L)
treatment reaches had the least percent remaining in both spring (38.7% +/- 3.5)
and fall (11.8% +l-1.9) experiments (Figure 3.2). In the Spring, carcass addition
had a signiﬁcant effect (F 1317:2231 p <0.0001), resulting in greater experimental
leaf pack mass loss than in streams without carcass addition (Table 3.2).
Invertebrate biomass in streams with carcass additions was dominated primarily
by the scraper Juga (Pleuroceridae) and the Trichoptera shredder Hetemplectron
(Table 3.3).

In the fall, the interaction of carcass and canopy treatments had a signiﬁcant
effect (F1,3o4=8.36 p =0.004) on decay rates, with faster breakdown of leaf packs

in the (L) treatment than in the (C) treatment (Table 3.2). Fall invertebrate

56

biomass was dominated by Juga sp., and was approximately 2-5 times greater in

(L) than in (C) treatments (Table 3.4).

Macroin vertebrates

A total of 74 insect genera and three chironomid subfamilies, and 8 non-
insect taxa were collected from red alder leaf packs in study (Table 3.5). The
caddisﬂies (Trichoptera), with representatives from 11 families and 21 genera
had 9 shredder taxa. Stoneﬂies (Plecoptera) also had 9 representative shredder

taxa.

Spring 2002

Leaf pack invertebrate biomass did not differ signiﬁcantly between
streams with and without carcass additions, between stream reaches with open
or closed canopies, or among experimental treatments (C, N, L and L+N).
However, an interaction with the covariate (time*carcass; F1,g4=5.28, p=0.0238)
revealed a pattern of decreasing invertebrate biomass in streams with carcass
additions, and increasing invertebrate biomass in streams without carcass
additions (Figure 3.3a). Neither main nor interactive effects of the study
signiﬁcantly affected invertebrate richness (all p>0.05). Invertebrate diversity was
signiﬁcantly greater in streams without carcass additions than in streams with

carcass additions (F1,5o=5.34, p=0.025). Additionally, an interaction with the

57

oovariate (time*carcass; F1,94=4.79, p=0.031) revealed a pattern of increasing
invertebrate diversity in streams with carcass additions (Figure 3.3b).

Neither main nor interactive effects of the study signiﬁcantly (all p>0.05)
affected percent functional group (shredder, scraper, gathering collector, ﬁltering

collector or predator) or shredder (biomass, richness or diversity).

Fall 2002

Leaf pack invertebrate biomass did not differ signiﬁcantly between
streams with and without carcass additions, between stream reaches with open
or closed canopies, or among experimental treatments (C, N, L and L+N). There
were interactions with the covariate. Leaf pack invertebrate biomass decreased
through time in streams with carcass additions, and increased through time in
streams without carcass additions (time*carcass; F1,153=4.0, p=0.0472) (Figure
3.4a). Furthermore, leaf pack invertebrate biomass decreased through time in
stream reaches with a closed canopy (time*canopy; F1,150=6.67, p=0.01) (Figure
3.4b). Neither main nor interactive effects of the study signiﬁcantly affected
invertebrate richness (all p>0.05). Invertebrate diversity did not differ between
streams with and without carcass additions, between stream reaches with open
or closed canopies, or among experimental treatments (C, N, L, L+N). However,
there were interactions with the covariate. Invertebrate diversity decreased
through time in streams without carcass additions and increased through time in

streams with carcass additions (time*carcass; F1,153=4.71, p=0.03) (Figure 3.5a).

58

Furthermore, invertebrate diversity increased through time in all but the (L)
treatment (time*carcass*canopy; F1_157=7.19, p=0.008) (Figure 3.5b).

Although not signiﬁcantly different (F1,47=3.82, p=0.0565), mean shredder
biomass was smallest in the (C) treatment and largest in the (L) treatment
(Figure 3.6). Shredder richness was not signiﬁcantly different between streams
with and without carcass additions, between open and closed canopies, or
among experimental treatments (C, N, L, L+N). Additionally, shredder diversity
was signiﬁcantly greater (F1,159=5.77, p=0.0174) in closed than in open canopy

reaches.

DISCUSSION

Although red alder leaf litter pack mass loss occurred faster in fall than in
the spring within our experimental study reaches, lack of seasonal replicates
prevents formal statistical comparison. There was a larger amount of invertebrate
biomass in fall leaf litter packs relative to spring, but the opposite was true of
shredder biomass. Perhaps, the greater mass loss in the fall was due to physical
abrasion by scrapers, such as Juga which dominated the invertebrate biomass,
feeding on surface bioﬁlm, as has been reported for some marine gastropods
(Profﬁtt and Devlin 2005). Furthermore, leaf litter pack mass loss is likely the
result of greater shredder biomass. Although not statistically signiﬁcant, faster
leaf litter pack mass loss in the spring is due to the dominance of the shredder

Heteroplectron in streams with carcass additions as well as in the fall in the open

59

canopy no carcass (L) treatment. Shredder biomass and taxa richness is
important to leaf litter mass loss rates (Jonsson et al. 2001). The greater
shredder biomass on leaf packs in open canopy reaches may have been the
result of lack of available natural leaf litter and therefore a concentration of
shredder individuals on the leaf packs in the vicinity. An important aspect to note
is that leaf mass loss rates were increased in all experimental treatments relative
to control stream reaches, which may affect overall stream food web dynamics.
Leaf pack invertebrate richness was not affected by experimental
treatments in this study, as has been previously reported in studies investigating
logging effects on leaf processing (Kreutzweiser et al. 2008), perhaps a result of
our low-impact approach to canopy removal. However, leaf pack invertebrate
diversity was affected during both seasons of study, being overall lower in
streams with carcass additions or changing through time. Invertebrate diversity
on experimental leaf packs was primarily affected by the overwhelming presence

ofJuga.

CONCLUSIONS

Experimental augmentation of light, nutrients, and both light and nutrients
reaching the stream of the study reaches resulted in greater red alder leaf pack
mass loss in both late spring and early fall 2002. Although canopy removal in
addition to carcass additions had a positive effect on benthic invertebrate

biomass of cobble habitats, it's effects on red alder leaf litter mass loss and

60

invertebrate community structure should also be taken into consideration. Further

investigation of this data is needed.

61

 

 

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62

Table 3.2. Daily decay rates (k) of red alder leaf packs from experimental study
reaches of the Smith and Klamath River basins, northern California. (C=
controls, N= augmented nutrients, L= augmented light, L+N= augmented light
and nutrients). k values with same letters are not signiﬁcantly different from
each other, Fishers LSD (p =0.05).

 

 

Spring Fall
k R2 k R2

Controls (C) 0.0195 0.81 0.0443 b 0.75
Nutrient (N) 0.0308 0.63 0.0595 ab 0.74
Light (L) 0.0326 0.64 0.0818 a 0.79
Light and Nutrient (L+N) 0.0303 0.70 0.0520 ab 0.61
Carcass 0.0302 a 0.61

No Carcass 0.0245 b 0.53

 

63

Table 3.3. Biomass (mg) and percent dominance of the most

abundant taxa: nutrient treatment (carcass, no carcass) by time
(Day 15, Day 23, Day 30), from Spring experimental red alder (Alnus rubra)

 

 

 

leaf packs.
Carcass No Carcass
Day Day Day Day Day Day
1 5 23 30 1 5 23 30

Biomass (mg)
Juga 4038 1479 980 1357 1313 1286
Calineuria 671 846 876
Heteroplectron 564 2431 1 169 476 694
Paraleptophlebia 936 498 434
H ydatoph yIax 775

% Dominance
Juga 62 21 23 35 26 25
Calineuria 1 0 1 7 1 7
Heteroplectron 9 35 27 12 14
Paraleptophlebia 1 3 12 1 1
Hydatophylax 1 5

 

Table 3.4. Biomass (mg) and percent dominance of the most abundant
taxa in (C) and (L) treatments by time (Day 15, Day 23, Day 30)
from fall experimental red alder (Alnus rubra) leaf packs.

 

 

 

Control Light
Day Day Day Day Day Day
1 5 23 30 1 5 23 30

Biomass (mg)
Juga 1669 1818 1538 2935 10055
Paraleptophlebia 141 310 270 219
Heteroplectron 139 163 301 605 915 323
Calineuria 1 50
Psychoglypha 640
Hydatophylax 140

% Dominance
Juga 56 54 50 64 90
Paraleptophlebia 1 9 9 9 2
Heteroplectron 1 9 5 9 20 20 3
Calineuria 21
Psychoglypha 22
H ydatoph ylax 3

 

65

Table 3.5 Checklist of taxa collected from experimental red alder (Alnus
rubra) leaf packs from study reaches of the Smith and Klamath River basins
of northern California. ( a- Spring, b— Fall, Sc- scraper, Gc— gathering
collector, F c- ﬁltering collector, Sh- shredder, Pi- piercer, Pr- predator)

 

 

Order Family Genus FFG
Ephemeroptera
Ameletidae Ameletus a b Go
Baetidae Baetis a b Gc
Ephemerellidae Attenella a D Go
Caudatella b Gc
Drunella a b Sc
Ephemerella a Go
Senate/Ia a b Go
Timpanoga b Gc
Heptageniidae Cinygma a D So
Cinygmula a b Go
Epeorus b Gc
lronodes a b Sc
Nixe b Sc
Rhithrogena b Go
Leptophlebiidae Paraleptophlebia a b Gc
Plecoptera
Capniidae b Sh
Chloroperlidae Kathroperla a b Gc
Suwallia a b Pr

66

Table 5 (cont.)

 

Leuctridae

Nemouridae

Peltoperlidae

Perlidae

Pteronarcyidae
Odonata

Gomphidae
Megaloptera

Sialidae
Trichoptera

Apatanidae

Brachycentridae

Sweltsa
Despaxia
Mose/fa
Pedomyia
Malenka
Zapada
Sierraperla
Soliperla
Calineuria
Claassenia
Hesperoperla
lsoperla
Osobenus

Pteronarcys

Sialis

Apatania

Brachycentrus

Micrasema

67

Pr
Sh
Sh
Sh
Sh
Sh
Sh
Sh
Pr
Pr
Pr
Pr
Pr

Sh

Pr

Pr

Sc

Fc

Sh

Table 5 (cont.)

 

Calamoceratidae
Hydropsychidae
Hydroptilidae
Lepidostomatidae

Limnephilidae

Polycentropodidae
Rhyacophilidae
Sericostomatidae

Uenoidae

Coleoptera

Dytiscidae

Heteroplectron
Hydropsyche
Hydroptila
Lepidostoma
Allocosmoecus
Ch yranda
Cryptochia
Dicosmoecus
Ecclisomyia
Homophylax
Hydatophylax
Philocasca
Psychoglypha
Polycentropus
Rh yacophila
Gumaga
Neophylax

Neothremma

Hydaticus

Hydrovatus

Oreodytes

68

Sh
Fc
Pi
Sh
Sc
Sh
Sh
Sc
60
Sh
Sh
Sh
Gc
Pr
Pr
Sh
So

So

Pr
Pr

Pr

Table 5 (cont)

 

Diptera

Elmidae

Hydrochidae

Psephenidae

Ceratopogonidae

Chironomidae

Dixidae

Empididae

Pelecorhynchidae
Psychodidae
Simuliidae

Tipulidae

Ampumixis
Lara
Narpus
Optioservus
Rhizelmis
Hydrochus

Acneus

Orthocladinae
Tan ypodinae
Tan yfarsini
Dixa

Chelifera
Clinocera
Hemerodromia
Oreogeton
Glutops
Pericoma
Simulium
Dicranota

Hexatoma

69

Go
Sh
Gc
So
So
Sh

Sc

Pr
Gc
Pr
Fc
Gc
Pr
Pr
Pr
Pr
Pr
Gc
F c
Pr

Pr

Table 5 (cont.)

 

Non-Insects
Collembola
Turbellaria
Oligochaeta
Hydracarina
Copepoda
Ostracoda
Amphipoda
Gammaridae
Gastropodae

Pleuroceridae

Pedicia

Juga

Pr

Gc
Pr
Gc
Pr
F 0
Go
Sh

Sc

 

7O

 

LOWER SMITH RIVER

r
l A
N

 

 

LOWER KLAMATH RIVER

a.

-.J ,
I“ ‘

I

/ 2/
+¥JUZJM

Figure 3.1. Study reaches within the lower Smith and Klamath River Basins of
northern California (after Vlﬁlzbach et al. 2005).

 

 

 

 

71

100 a)

 

on
2: 8' ‘*
m \
0E: 60 \\
02: -o- C \
D 4' -I- N
u.
‘5) q— L
°\ 20 -- L+N

I

10'
b)

.9.)
C
76
E
a:
[I
2
O
u.
<
o\° 20

O

0 1O 20 30

Time (Days)

Figure 3.2. Percent AFDM experimental red alder leaf pack remaining (+/- 1 SE.)
at day (2, 15, 23, and 30) in study reaches of the Smith and Klamath River
basins during a) spring and b) fall.

72

Invertebrate Biomass (mg) +/- SE.

Invertebrate Diversity (H') +/- SE.

 

 

 

 

 

50 a)
40
30
20
10
T I F r l
10 15 20 25 30 35
2.0] b)
1 .8
15¢ }‘
1 .41
-o- Carcass

1'27 -I- No Carcass
1 .0-

10 15 20 25 30 35

Time (Days)

Figure 3.3. Spring a) dry mass and b) Shannon-Weiner diversity (H') (+/- 1 SE.)
of macroinvertebrates on experimental red alder leaf packs at days (15, 23 and
30) from streams of the Smith and Klamath River basins.

73

 

 

 

 

Hi

03 100

3} -0- Carcass

,.. a)
E) 30 -I- No Carcass

é’ 60

S

a 40 %

8

n 20 *4)

8

9 I I F I u
E

UJI

CD 80

1' -0- Open Canopy b)
’7 -I- Closed Cano

E 60 PY

I’D

U)

E 40

o

a \

d)

‘5 20

.0

8

9 I I 1 I u
E 10 15 20 25 30 35

Time (Days)

Figure 3.4. Fall dry mass (+/- 1 SE.) of invertebrates on experimental red alder
leaf packs at days (15, 23 and 30) from a) streams and b) study reaches of the
Smith and Klamath River basins.

74

 

 

 

 

 

 

u.i

m 1.6 a)

1

'3; 1.4

3

l2

3 1.2

o

g 1.0 -o- Carcass

g -I- No Carcass

a)

E 0-8 ﬂ I I r l

u.i

(15 2.5-

1'

f 2.0- b)

.é‘ A

a, -

.2 ‘ 4+

D 1.0-

Q

P; 0.5 -0- L+N + L

g 1 -I- N 4- C

E 0.0 I I F F u

- 10 15 20 25 30 35
Time (Days)

Figure 3.5. Fall Shannon-Weiner diversity (H') (+/- 1 SE.) of macroinvertebrates
on experimental red alder leaf packs at days (15, 23 and 30) from a) streams and
b) treatment study reaches of the Smith and Klamath River basins.

75

 

 

ui

m1”

1

a 8'

E

8’ 6'

E

.9 4'

m

8 2

.0

e i

('Ic) I

C N L L+N
Treatment

Figure 3.6. Fall shredder biomass (+/- 1 SE.) on experimental red alder leaf
packs among study treatments in the Smith and Klamath River basins.

76

LITERATURE CITED

77

Literature Cited

Ambrose, H.E., Wllzbach, MA. and K.W. Cummins. 2004. Periphyton response
to increased light and salmon carcass introduction in northern California
streams. Journal of The North American Benthological Society. 23(4):
701-712.

Barlocher, F. 2005. Leaf mass loss estimated by litter bag technique. In Methods
to Study Litter Decomposition: A Practical Guide. Edited by M.A.S. Graca,
F. Barlocher and MC. Gessner. The Netherlands, Springer: 37-42.

Bates, J.D., Svejcar, TS. and RF. Miller. 2007. Litter decomposition in cut and
uncut western juniper woodlands. Journal of Arid Environments. 70(2):
222-236.

Benﬁeld, E.F. 1996. Leaf breakdown in stream ecosystems. In Methods in
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Academic press: 579-589.

Crowl, T.A., Welsh, V., HeartsilI—Scalley, T., and AP. Covich. 2006. Effects of
different types of conditioning on rates of leaf-litter shredding by
Xiphocan's elongata, a Neotropical freshwater shrimp. Journal of the North
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Ferreira, V. and M.A.S. Graca. 2007. Fungal activity associated with
decomposing wood is affected by nitrogen concentration in water.
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Ferreira, V., Gulis, V. and M.A.S. Graca. 2006. Whole-stream nitrate addition
affects litter decomposition and associated fungi but not invertebrates.
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Franken, R.J.M., Waluto, B., Peeters, E.T.H.M., Gardeniers, J.J.P., Beijer, J.A.J.
and M. Scheffer. 2005. Growth of shredders on leaf litter bioﬁlms: the
effect of light intensity. Freshwater Biology. 50: 459-466.

Greenwood, J.L., Rosemond, A.D., Wallace, J.B., Cross, W.F. and HS. Weyers.
2007. Nutrients stimulate leaf breakdown rates and detritivore biomass:
bottom-up effects via heterotrophic pathways. Oecologia. 151 (4): 637-649.

Hauer, F .R. and V.H. Resh. 1996. Benthic Macroinvertebrates. In Methods in

Stream Ecology. Edited by F.R. Hauer and GA. Lamberti. San Diego,
Academic press: 339-369.

78

Hauer, F .R. and V.H. Resh. 2007. Benthic Macroinvertebrates. In Methods in
Stream Ecology. Edited by PR. Hauer and GA. Lamberti. San Diego,
Academic press: 435-463.

lshikawa, H., Osono, T. and H. Takeda. 2007. Effects of clear-cutting on
decomposition processes in leaf litter and the nitrogen and lignin dynamics
in a temperate secondary forest. Journal of Forest Research. 12(4): 247-
254.

Jonsson, M., Malmqvist, B. and P. Hoffsten. 2001. Leaf litter breakdown rates in
boreal streams: does shredder species richness matter? Freshwater
Biology. 46: 161-171.

Knorr, M., Frey, SD. and PS. Curtis. 2005. Nitrogen additions and litter
decomposition: a meta-analysis. Ecology. 86(12): 3252-3257.

Kominoski, J.S., Pringle, C.M., Ball, B.A., Bradford, M.A., Coleman, D.C., Hall,
DB. and MD. Hunter. 2007. Nonadditive effects of leaf litter species
diversity on breakdown dynamics in a detritus-based stream. Ecology.
88(5): 1167-1176.

Kreutzweiser, D.P., Good, K.P., Capell, SS. and SB. Holmes. 2008. Leaf-litter
decomposition and macroinvertebrate communities in boreal forest
streams linked to upland logging disturbance. Journal of the North
American Benthological Society. 27(1): 1-15.

Merritt, R.W. and K.W. Cummins.1996. Introduction to The Aquatic Insects of
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K.J., and J.L. Lessard. 2002. Development and application of a
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remnant river oxbows in southwest Florida. Jounal of the North American
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Paul, M.J., Meyer, J.L. and CA. Couch. 2006. Leaf breakdown in streams
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Peterson, RC. and K.W. Cummins. 1974. Leaf processing in a woodland stream.
Freshwater Biology. 47: 129-141.

79

Profﬁtt, CE. and DJ. Devlin. 2005. Grazing by the intertidal gastropod Melampus
coffeus greatly increases mangrove leaf litter degradation rates. Marine
Ecology - Progress Series. 296: 209-218.

Vannote, R.L., Minshall, G.W., Cummins, K.W., Sedell, JR, and CE. Cushing.
1980. The river continuum concept. Canadian Journal of Fisheries and
Aquatic Sciences. 37(1): 130-137.

Vlﬁlzbach, M.A., Harvey, B.C., White, J.L. and R.J. Nakamoto. 2005. Effects of
riparian canopy opening and salmon carcass addition on the abundance
and growth of resident salmonids. Canadian Journal of Fisheries and
Aquatic Sciences. 62(1): 58-67.

Yanai, S. and K. Kochi. 2005. Effects of salmon carcasses on experimental
stream ecosystems in Hokkaido, Japan. Ecological Research. 20(4): 471-
480.

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80

APPENDIX

81

Appendix 1
Record of Deposition of Voucher Specimens*
The specimens listed on the following sheet(s) have been deposited in the named museum(s) as
samples of those species or other taxa, which were used in this research. Voucher recognition
labels bearing the Voucher No. have been attached or included in ﬂuid-preserved specimens.

Voucher No.: 2009-01

Title of thesis or dissertation (or other research projects):

Museum(s) where deposited and abbreviations for table on following sheets:

Entomology Museum, Michigan State University (MSU)

Other Museums:

Investigator’s Name(s) (typed)
Q§valgo Hernandez

 

 

 

Date 30. April 2009

*Reference: Yoshimoto, C. M. 1978. Voucher Specimens for Entomology in North America.
Bull. Entomol. Soc. Amer. 24: 141-42.

Deposit as follows:
Original: Include as Appendix 1 in ribbon copy of thesis or dissertation.

Copies: Include as Appendix 1 in copies of thesis or dissertation.
Museum(s) ﬁles.
Research project ﬁles.

This form is available from and the Voucher No. is assigned by the Curator, Michigan State
University Entomology Museum.

82

Appendix 1.1

Voucher Specimen Data

1 of 1_2_ Pages

Page

 

Numb“ 83‘ 3

2552:: 29o 5822.2 05 5 58%
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no.2 F 88 x 55. £5 do 888E:
:ms. _. moow> Loam 5E5 .do o‘dodmtom
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e e s h n k P w 9 coxﬂ 55° Lo 36on
s h o t u u u m 9 Lo mom 8 ocoanaoB mm m m .
no .3832

 

83

Appendix 1.1

Voucher Specimen Data

2 of 3 Pages

Page

Boo .oFoLno

 

.Enoons. >mo.oEoEm.
3.99.5 23w $952.2 9: c. awedon
.8 208.8% Ugo: o>onm o£ nozooom

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nos. F «com > 52m 53%. do anodesmx
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nos. F 88 x 52m £.Eo 82.88
<mmFdoomnd
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82.82833
nos. F Foom x 52m. £.Eo do «conﬁrm
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nos. F Foom x 52m £_Eo .do 88:8.
nos. F 88 x 52m £.Eo .do £28.38:
nos. F 58 x and 5.5 .do oEoodm
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.8 6.85:: .o 282 .on .5 .<on
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M w A A N
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84

Appendix 1.1

Voucher Specimen Data

_3_of g Pages

Page

Ema .oFmSo

 

.Enoons. 30.060ch
2.92.5 29o 898.2 o... c. goodou
.2 808.3% 69o: o>onm 05 82081

 

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nos. F 88 x 82”. 5.5 do 9885.35...
$258.52
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cc .3532

 

85

Appendix 1.1

Voucher Specimen Data

ges

iof12_Pa

Page

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86

Appendix 1.1

Voucher Specimen Data

5_ of 1_2_ Pages

Page

QHNO .O~M.=..O

 

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87

Appendix 1.1

0.00 .2050

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3.0.02.5 0.0.o 00020.5. 0;. 0. 200000
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88

Appendix 1.1

Voucher Specimen Data

7_of 3 Pages

Page

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89

Appendix 1.1

Voucher Specimen Data

8_ of g Pages

Page

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90

Appendix 1.1

Voucher Specimen Data

9_ of 12_ Pages

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Appendix 1.1

Voucher Specimen Data

Page .11 of 1; Pages

 

 

 

 

 

 

 

 

 

 

 

 

 

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92

Appendix 1.1

Voucher Specimen Data

Page _1_1_ of g Pages

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93

Appendix 1.1

Voucher Specimen Data

Page 1_2_ of _1_2_ Pages

9.3 .9930

 

88 .5? .8 28

 

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94

    
  

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