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thesis entitled

MACROINVERTEBRATE COMMUNITY RESPONSE TO
RIPARIAN FIED ALDER WITHIN HEADWATER
STREAMS OF SECOND-GROWTH FORESTS IN
SOUTHEAST ALASKA

presented by

Ryan K. Kimbirauskas

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MACROINVERTEBRATE COMMUNITY RESPONSE TO RIPARIAN RED ALDER
WITHIN HEADWATER STREAMS OF SECOND-GROWTH FORESTS 1N
SOUTHEAST ALASKA

by

Ryan K. Kimbirauskas

A THESIS

Submitted to
Michigan State University
In partial fulfillment of the requirements
For the degree of

MASTER OF SCIENCE
Department of Entomology

2004

ABSTRACT
MACROINVERTEBRATE COMMUNITY RESPONSE TO RIPARIAN RED ALDER
WITHIN HEADWATER STREAMS OF SECOND-GROWTH FORESTS IN
SOUTHEAST ALASKA
by
Ryan K. Kimbirauskas
Recent declines in Pacific Northwest salmon returns and the subsequent loss of

in-stream nutrients have increased consideration for watershed restoration practices to
include Red alder (Alnus rubra Bong). Red alder is an aggressive pioneer species that
colonizes disturbed soils, particularly within riparian corridors of young-growth forests
following clearcut timber harvesting. Although riparian red alder appears to add
nutrients to aquatic ecosystems, the thinning and removal of alder within regenerating
forests is still practiced. In an effort to improve scientific knowledge about the ecological
role of red alder on aquatic ecosystems, we compared benthic macroinvertebrate
colonization of wood substrates among 13 headwater streams with a range riparian alder
(0-53%). Research was conducted within the Maybeso Creek and Harris drainages on
eastern Prince Of Wales Island, southeast Alaska. Alder wood, especially pieces in more
advanced decay, generally supported greater density and diversity of macroinvertebrates
than did conifer wood. Collectors and shredders responded to the presence of riparian
alder, and streams with more alder in riparian habitats produced greater mean
macroinvertebrate density and biomass. Results from this study suggested that the
presence of alder along headwater corridors enhances macroinvertebrate productivity and
may subsidize lost nutrients within Pacific Northwest watersheds, thus managing forested
uplands to include red alder should be considered as a management tool to improve

saImonid production.

ACKNOWLEDGEMENTS

I would first like to thank my parents, Paul and Cinda for their love, and unconditional
support throughout this process; and my sister, Paula for her friendship and encouragement.
I would also like to acknowledge my friends that kept me grounded. Christian for his
humor, Brandon for his open door, Eric for traveling the world with me, and Mollie Balls
for being a constant source of energy and inspiration. Of course, this could not have been
half the experience without Rich. Thank you for the opportunities, the dedication to my

development as a student and educator, and the constant entertainment.

TABLE OF CONTENTS

LIST OF TABLES .................................................................................. v
LIST OF FIGURES ................................................................................ vi
CHAPTER 1

BACKGROUND OF PREVIOUS RESEARCH EXAMINING THE EFFECTS OF
CLEAR-CUT LOGGING ON HEADWATER HABIT ATS AND ASSOCIATED
AQUATIC ECOSYSTEMS IN THE PACIFIC NORTHWEST

Introduction ........................................................................................... 1
Temperature & Thermal Recovery2
Nutrients & Macroinvertebrate Response ................................................ 3
Recruitment & Distribution of Wood Debns4
Summary & Management Implications ................................................... 5

Literature Cited ........................................................................................ 7

CHAPTER 2

MACROINVERTEBRATE COMMUNITY RESPONSE TO RIPARIAN RED
ALDER WITHIN HEADWATER STREAMS OF SECOND-GROWTH FORESTS IN
SOUTHEAST ALASKA

Introduction ........................................................................................ 12
Methods ............................................................................................. 16
Stand Selection & Study Sites ........................................................... 16
Wood Collection & Experimental Design ............................................. 18
Macroinvertebrate Sampling & Analysis ............................................. 19
Results .............................................................................................. 2O
Macroinvertebrates Community Composition & Functional Feeding
Groups ..................................................................................... 20
Macroinvertebrate Density, Biomass & Diversity .................................. 22
Discussion ......................................................................................... 23
Conclusion ......................................................................................... 29
Literature Cited .................................................................................... 47
Appendix 1 ........................................................................................... 56

iv

LIST OF TABLES

CHAPTER 2

Table 1. Physical and biological characteristics of study sites ............................. 31

Table 2. Checklist of taxa collected and percent relative density and abundance
collected over the entire study period (2001-2002) .......................................... 32

LIST OF FIGURES

Figure 1. Red alder colonization following soil disturbance along cleared stream banks,
abandoned logging roads, and landslides within the Maybeso experimental Forest on
Prince Of Wales Island, southeast Alaska ....................................................... 36

Figure 2. Maybeso Experimental Forest, eastern Prince of Wales Island, Southeast
Alaska (132° 67’W, 55° 49’N) ..................................................................... 37

Figure 3. Study sites within the Maybeso Creek and Harris River drainages on
Prince Of Wales Island, southeast Alaska ....................................................... 38

Figure 4. Pieces of wood representing four treatments: early decay alder (EA), late
decay alder (LA), early decay conifer (EC), and late decay conifer (LC) .................... 39

Figure 5. Percent distribution of the five most common families (and inset bar graph of
midge subfamilies) collected from wood surfaces between 2001 and 2002 ................ 40

Figure 6. Percent relative abundance of functional feeding groups collected from the
surfaces of wood substrates between 2001 and 2002 ........................................... 41

Figure 7. Functional feeding groups relative abundance in streams with low
(0-9%, n=4), medium (10-35%, n=4), and high (36-53%, n=5) percentages of
riparian red alder .................................................................................... 42

Figure 8. Mean macroinvertebrate (A) density and (B) biomass in streams with low,
medium, and high percentages of riparian red alder ............................................. 43

Figure 9. Mean macroinvertebrate (A) count and (B) biomass collected from the surfaces
of naturally collected alder and conifer wood between 2001 and 2002 ...................... 44

Figure 10. Shannon-Weiner diversity of macroinvertebrates collected from the surfaces
of alder and conifer wood (A) and from recent and late decay classes (B) between 2001
and 2002 .............................................................................................. 45

Figure 11. Mean macroinvertebrate density (A) and biomass (B) collected from the

surfaces of wood substrates in recent and late stages of decay. Totals are based on
collections of both alder and conifer wood between 2001 and 2002 ......................... 46

vi

CHAPTER 1

BACKGROUND OF PREVIOUS RESEARCH EXAMININ G THE EFFECTS OF
CLEAR-CUT LOGGING ON HEADWATER HABITATS AND ASSOCIATED
AQUATIC ECOSYSTEMS IN THE PACIFIC NORTHWEST

INTRODUCTION

Forest management practices have improved dramatically since large-scale
commercial forestry began in the 1950’s. Among these improvements are shifts from
strict timber management to practices that preserve biodiversity and maintain long-term
sustainability of all forest resources. In the Pacific Northwest, considerable attention has
been aimed towards the effects of clearcutting on large streams that contain fish,
particularly Pacific salmon. In recent years, there has been a growing concern with the
management of small, headwater streams and their associated riparian habitats.
Currently, the small, often fishless, streams that drain forested watersheds receive little or
no protection during logging operations and management is based on limited scientific
knowledge. Here I present a brief synopsis of recent studies that address these issues. I
will comment on the direct and indirect effects of clearcutting on 1) in-stream
temperature and thermal recovery, 2) macroinvertebrate community structure and
production, and 3) wood recruitment into headwater streams. These findings are
primarily from sites in the Pacific Northwest and central British Columbia and mostly

published within the last decade.

TEMPERATURE & THERMAL RECOVERY

Among the many physical responses of headwater streams to clearcutting, stream
temperature dynamics are the most extensively studied. This is primarily due to the
negative impacts that even slight increases in stream temperatures can generate on
aquatic organisms, particularly cool-water fish species. Macdonald et al. (2003a) studied
the effects of various riparian harvesting practices on the temperature of first-order
streams in northern British Columbia, Canada. The harvesting practices represented a
range of possible forest management options for headwater streams as outlined by the
Forest Practices Code of British Columbia. Results from this study showed that the daily
maximum temperature increased 4-6 °C following harvest—even when partial riparian
vegetation was intact. They also found that these temperatures remained elevated for 5
yrs following harvest. In the western Cascades of Oregon, the thermal recovery period
following canopy removal was reported to be as long as 15 yrs within small streams
(Johnson and Jones 2000). Story et al. (2003) compared stream temperature in cleared
and shaded reaches of small streams in central British Columbia and found that stream
temperatures generally cooled downstream, where the canopy was complete, and the
magnitude of cooling depended to a large extent on fluctuations in streamflow from
upstream reaches. Moore et a1. (2003) found similar trends in stream cooling in two
small tributaries downstream of clearings associated with logging. In southeast Alaska,
Hernandez et al. (2004) compared the temperatures of headwater streams that flowed
through old-growth, second-growth, and recently clearcut forests. Results showed that
daily maximums were higher in the old-growth condition, however the greatest

differences in maximum and minimum temperatures were found in the clearcut condition.

Future research with regard to thermal recovery and riparian disturbance will investigate
the role of stream substrates and hyporheic flow on temperature fluxes in headwater

streams (Macdonald et al. 2003b, Moore et al. 2003, Story et al. 2003).

NUTRIENT S & MACROINVERTEBRATE RESPONSE

Headwater streams deliver nutrients to downstream reaches and regulate energy
flow in aquatic systems. In the Pacific Northwest, many of these small streams drain into
larger systems that support Pacific salmon. Several studies in this region have examined
the effects of timber harvest on the accumulation, storage, and transport of organic matter
in forested watersheds (Bilby and Bisson 1992, Culp and Davies 1985, Hall et al. 1987).
Recent interest has been directed towards the effects of logging on macroinvertebrate
communities in headwater streams and the drift of this food source to juvenile salmon.
Anderson (1992) provided a detailed account of macroinvertebrate response to
disturbance in small, upland streams of the Pacific Northwest and found that logging
activities alter the food source for macroinvertebrates and therefore, change the
functional feeding group composition of invertebrate communities. Greater solar inputs
and increased periphyton growth is common within recently cleared streams and may
lead to initial increases in macroinvertebrate production (Cole et al. 2003, Fuchs 1999,
Hernandez et al. 2004). Fuchs et al. (2003) found similar results in small streams of the
central interior of British Columbia and suggested that, with the onset of canopy closure,
macroinvertebrate production may return to prelogging conditions as early as 10 years
following timber harvest. Wipfli (1997) identified terrestrial macroinvertebrates

originating from riparian vegetation in second-growth forests as a major food source for

downstream juvenile salmon and later found that drifting invertebrates provide enough
biomass annually to support up to 2000 juvenile salmonids per kilometer of river (Wipfli
and Gregovich 2002). In regenerating forests of southeast Alaska, the presence of
riparian alder along headwater streams appears to enhance macroinvertebrate production
(Lesage et al. 2003, Piccolo and Wipfli 2002, Wipfli and Musslewhite 2004). Price et al.
(2003) compared invertebrate communities in headwater streams of old-growth and
recently clearcut forests of British Columbia and found that stream size, persistence of
flow, and canopy cover were good predictors of invertebrate abundance. They also
suggested that forest management practices should offer protection during logging

operations for intermittent, as well as, perennial streams.

RECRUITMENT & DISTRIBUTION OF WOOD DEBRIS

Wood produced from riparian vegetation is critical to the physical and biological
attributes of forested streams and in some small streams appears to play a more
prominent role than previously understood. In the Oregon Coast Range, May (2002)
found that large wood entering headwaters during landslides could be delivered
downstream to fish-bearing reaches. In southest Alaska, Martin and Benda (2001)
quantified wood recruitment and redistribution mechanisms at the watershed scale and
provided information on how and where to protect sources of wood debris that enter
headwater streams. May and Gresswell (2003) investigated the recruitment and
redistribution of large wood in headwater streams of the southern Coast Range of

Oregon, and concluded that stream size and slope stability strongly influence the

processes that recruit and redistribute wood in low—order channel networks. They
suggested that the recruitment of large wood to high gradient streams may not be possible
in intensely harvested basins that lack streamside buffers. A substantial proportion of the
volume and number of pieces of large wood in Cummins Creek Watershed in western
Oregon is up-slope derived wood (Reeves et al. 2003). These results were similar to
those found in Redwood National Park, California (Benda 2002) and Olympic National
Park, Washington (Benda et al. 2003). In western Washington, Ralph et al. (1994) and
McHenry et al. (1998) reported less large wood in streams that flowed through second—
growth forests compared to pre-harvest records. Both concluded that the removal of
streamside vegetation was the reason for the reduction in recruitment of wood into
streams. For 15 steep, headwater streams in southeast Alaska and demonstrated that
timber harvesting within riparian habitats influences the recruitment, distribution, and
accumulation of woody debris in forested watersheds (Gomi et al. 2001). Hernandez et
al. (2004) reported similar results and also found differences in the recruitment of large

wood into streams of alder and conifer dominated second-growth forests.

SUMMARY & MANAGEMENT IMPLICATIONS

Although small streams account for nearly 90% of the total channel networks in
forested watersheds of the Pacific Northwest (Gomi et al. 2002), only recent efforts have
been directed towards understanding their ecological role in aquatic ecosystems. Current
management of headwater habitats, particularly in the context of timber harvesting, is

therefore based on limited scientific knowledge and has lead to inconsistent practices and

generated debate over the degree at which these areas should be managed. Recent
interest has been given to the potential of managing forests to include red alder (Alnus
rubra Bong.)as a way of improving timber, wildlife and aquatic resources. The focus of
my research was to examine the role of riparian red alder on macroinvertebrate
communities within headwater streams that flow through second-growth forests of
southeast Alaska. The development of new policy regarding the management of
headwater habitats depends on increased knowledge of upland habitats and a compromise

between ecological and economic interests.

LITERATURE CITED

Literature Cited

Anderson, NH. 1992. Influence of disturbance on insect communities in Pacific
Northwest streams. Hydrobiologia. 248: 79-92.

Benda, L.E., Bigelow, P., and Worsley, W. 2002. Recruitment of instream large wood in
old-growth and second-growth redwood forests, northern California, USA. Can. J. For.
Res. 32: 1460-1477.

Benda, L.E., Veldhuisen, C., and Black, J. 2003. Influence of debris flows on the
morphological diversity of channels and valley floor, Olympic Peninsula, Washington.
Geol. Soc. Am. Bull. In press.

Bilby, RE. and Bisson, P.A.1992. Allochthanous versus autochthanous organic matter
contributions to the trophic support of fish populations in clear-cut, and second-growth
forests in southwestern Washington. Can. J. Fish. Aquat. Sci. 48: 2499-2508.

Cole, M.B, Russell, K.R., and Mabee, T.J. 2003. Relation of headwater
macroinvertebrate communities to in-stream and adjacent stand characteristics in

managed second-growth forests of the Oregon Coast Range mountains. Can J. For. Res.
33(8): 1433-1443.

Culp, J .M., and Davies, R.W. 1985. Responses of benthic macroinvertebrate species to
manipulation of interstitial detritus in Carnation Creek, British Columbia. Can. J. Fish.
Aquat. Sci. 42: 139-146.

Fuchs, SA. 1999. Responses of macroinvertebrate community composition to changes in
stream abiotic factors after streamside clear-cut logging in the central interior of British
Columbia. M.Sc. thesis, The University of British Columbia, Vancouver, BC.

Fuchs, S.A., Hinch, S.G., and Mellina, E. 2003. Effects of streamside logging on stream

macroinvertebrate communities and habitat in the sub—boreal forests of British Columbia,
Canada. Can. J. For. Res. 33: 1408-1415.

Gomi, T., Sidle, R.C., Bryant, M.D., and Woodsmith, RD. 2001. The characteristics of
woody debris and sediment distribution in headwater streams, southeastern Alaska. Can.
J. For. Res. 31: 1386-1399.

Hall, J .D., Brown, G.W., and Lantz, R. 1987. The Alsea Watershed Study: A
Retrospective. In Streamside Management: Forestry and Fishery Interactions. Edited by
Salo EO and Cundy TW. College of Forest Resources, University of Washington,
Contribution 57, Seattle. pp. 399-416.

Hernandez, O., Merritt. R.W., and Wipfli, MS. 2004. Benthic invertebrate community
structure is affected by forest succession after clear-cut logging southeast Alaska.
Hydrobiologia. In press.

Johnson, S.L., and Jones, J .A. 2000. Stream temperature responses to forest harvest and
debris flows in he western Cascades, Oregon. Can. J. Fish. Aquat. Sci. 57(Suppl. 2): 30-
39.

LeSage, CM. 2003. Headwater riparian invertebrate community changes in response to
red alder stand composition in southeastern Alaska. Thesis for the Degree of MS,
Michigan State Univeristy, East Lansing, Michigan.

Macdonald, J.S., Beaudry, E.A., MacIsaac, EA, and Herunter H.E. 2003a. The effects of
forest harvesting and the best management practices on streamflow and suspended
sediment concentrations during snowmelt in headwater streams in sub-boreal forests of

British Columbia, Canada. Can. J. For. Res. 33: 1397-1407.

Macdonald, J .S., MacIsaac, EA, and Herunter H.E. 2003b. The effect of variable-
retention riparian buffer zones on water temperatures in small headwater streams in sub-
boreal ecosystems of British Columbia, Canada. Can. J. For. Res. 33: 1371-1382.

McHenry, M.L., Schott, E., Conrad, RH, and Grette, GB. 1998. Changes in the quantity
and characteristics of large woody debris in streams of the Olympic Peninsula,
Washington, USA. (1982-1993). Can. J. Fish. Aqaut. Sci. 55: 1395-1407.

Martin, D.J., and Benda, LE. 2001. Patterns of instream wood recruitment and transport
at the watershed scale. Trans. Am. Fish. Soc. 130: 940-958.

May, CL. 2002. Debris flows through different forest age classes in the central Oregon
Coast Range. J. Am. Water Resour. Assoc. 38(4): 1097-1113.

May, C.L., and Gresswell, RE. 2003. Large wood recruitment and redistribution in
headwater streams in the southern Oregon Coast Range, U.S.A. Can. J. For. Res. 33:
1352-1362.

Moore, R.D., Macdonald, J .S., Herunter H. 2003. Downstream thermal recovery of
headwater streams below cutblocks and logging roads. Can. Tech. Rep. Fish. Aquat. Sci.
In press.

Piccolo, J .J ., and Wipfli, MS. 2002. Does red alder (Alnus rubra) along headwater
streams increase the export of invertebrates and detritus from headwaters to fishbearin g
habitats in southeastern Alaska? Can. J. Fish.Aquat. Sci. 59(3): 503-513.

Price, K., Suski, A., McGarvie, J ., Beasley, B., and Richardson, J .S. 2003. Communities
of aquatic insects of old-growth and clearcut coastal headwater streams of varying flow
persistence. Can. J. For. Res. 33: 1416-1432.

Ralph, S.C., Poole, G.C., Conquest, L.L., and Naiman, R]. 1994. Stream channel
morphology and woody debris in logged and unlogged basins of western Washington.
Can. J. Fish. Aqaut. Sci. 51: 37-51.

Reeves, G.H., Burnett, K.M., and McGarry, E.V. 2003. Sources of large wood in the
main stem of a fourth-order watershed in Coastal Oregon. Can. J. For. Res. 33: 1363-
1370.

Story, A., Moore, RD, and Macdonald, IS. 2003. Stream temperatures in two shaded
reaches below cutblocks and logging roads: downstream cooling linked to substrate
hydrology. Can J. For. Res. 33: 1383-1396.

Wipfli, MS. 1997. Terrestrial invertebrates as salmonid prey and nitrogen sources in
streams: contrasting old-growth and young-growth riparian forests in southeastern
Alaska, USA. Can. J. Fish. Aquat. Sci. 54: 1259-1269.

Wiplfli, MS. and Gregovich, DP. 2002. Export of invertebrates and detritus from
fishless headwater streams in southeastern Alaska: implications for downstream salmonid
production. Freshwater Biology. 47(5): 957-969.

10

Wipfli, MS. and Musslewhite, J. 2004. Density of red alder (Alnus rubra) in headwaters
influences invertebrate and detritus subsidies to downstream fish habitats in Alaska.
Hydrobiologia. 520: 153-163.

11

CHAPTER 2

MACROINVERTEBRATE COMMUNITY RESPONSE TO RIPARIAN RED ALDER
WITHIN HEADWATER STREAMS OF SECOND-GROWTH FORESTS IN
SOUTHEAST ALASKA

INTRODUCTION

Riparian zones are the interfaces between terrestrial and aquatic environments and
represent one of the most dynamic habitats of forested watersheds. The water, sediment,
and wood that enter small streams from riparian habitats are critical to the structure and
functional stability of headwaters and their downstream reaches (Carline and Spotts 1998,
Haigh et al. 1998, Naiman et al. 1992, Wipfli and Gregovich 2002). Wood produced
from riparian vegetation controls embankment erosion, stabilizes channel flow, provides
habitat for invertebrates and fish, and regulates nutrient and energy flow (Bilby and
Likens 1980, Cummins et al. 1989, May 2002, Reeves et al. 2003, Wallace et al. 1997,
Webster et al. 1992). Disturbing riparian habitats alters the physical characteristics of
adjacent streams and disrupts the energy flow in aquatic ecosystems (Bisson et al. 1987,
Bryant 1985, Dudley and Anderson 1982, Philips and Kilambi 1994). In the Pacific
Northwest, the freshwater ecosystems that support juvenile salmon appear to be nutrient
limited (Ashley and Slaney 1997); disturbing headwater habitats may reduce nutrients
downstream and limit the production of salmon (Hayes et al. 2000, Resh et al. 1988,

Wipfli and Gregovich 2002).

Timber harvesting has been identified as a both a short and long-term disturbance

that alters the energy base upon which forested headwaters and downstream food webs

12

are dependent (Alaback 1982, Duncan and Brusven 1985, Stone and Wallace 1998,
Wipfli 1997). Clearcutting, a method of harvesting and regenerating timber in which all
trees are cleared from a site, can immediately reduce total benthic macroinvertebrate
densities up to 50% (Hartman et al. 1996). Removing riparian vegetation also directly
eliminates input of wood debris into adjacent streams and reduces input of large wood for
many years (Bragg 1997, Bryant and Sedell 1995, May and Gresswell 2003, McHenry et
al. 1998). Maintaining in-stream wood habitat is advantageous for large numbers of
benthic invertebrates that could potentially benefit the diet of fish downstream
(Hernandez et a1. 2004, Wipfli 1997). Small streams are more easily disrupted than
larger rivers and clearcutting and the process of forest regeneration in headwaters may
alter downstream production (Plotnikoff 1994). Because red alder is a common riparian
hardwood of regenerating forests in the Pacific Northwest, recent interest has been
generated with respect to its effect on macroinvertebrate communities and associated
food webs (Haggerty et al. 2004, Richardson and Shaughnessy in press, Wipfli and

Gregovich 2002).

Red alder is an aggressive pioneer species with a range extending from southern
California to southeast Alaska (Harrington 1990). The process of clearcutting creates a
disturbance where mineral soils are exposed (Newton and Cole 1994) and the shade
intolerant alder quickly colonize riparian habitats along cleared stream banks, abandoned
logging roads and log transfer sites (Harrington 1990, Hulten 1968, Newton et al.
1968)(Figure 1). Red alder may remain at these locations up to 80 years before being

outgrown by conifers (Hibbs et al. 1994). Historically, forest managers regarded red

13

alder as an undesirable species and attempted to remove it from riparian and upland forest
stands; however, its ability to supply the soil with fixed nitrogen (Briggs et al. 1978,
Gordon et al. 1979, Trappe et al. 1968) encouraged its use as a management tool to
enhance productivity of young-growth conifer forests in the Pacific Northwest (Binkley
1981, DeBell et al. 1978). The presence of red alder in riparian habitats also appears to
benefit associated aquatic ecosystems (Allan et al. 2003, Irons et al. 1988, Wipfli 1997,
Wipfli et al. 2002) and may increase benthic macroinvertebrate abundance (Hernandez et
al. 2004, Wipfli et al. 2003). Therefore, managing upland young-growth forests to

include red alder may have substantial effects on salmon production in southeast Alaska.

Large-scale commercial logging and clearcutting in southeast Alaska began in the
19503 (Swanston 1967). During this period, management of riparian debris was
inconsistent (Bragg and Kershner 1999) and harvesting timber and developing logging
roads took priority over all other uses and resources of the forest (USDA 1997).
Clearcutting is still the most common practice of removing trees in southeast Alaska;
however, current policies include protection for streams and rivers that support salmon
(USDA 1999). For example, riparian buffer strips are required on fish-bearing streams
during logging operations to prevent loss of allochthanous inputs to aquatic ecosystems;
however, many of the small fishless streams within forested areas receive little or no
protection during a clearcut (USDA 1997). Headwater streams account for 70% - 90% of
watershed channel systems in southeast Alaska (Gomi et al. 2002, Swanston 1967), and
food generated from these headwaters plays a major role in regulating salmonid

production (Hayes et al. 2000). Wipfli (1997, et al. 2002) found that over half of the prey

14

biomass ingested by juvenile salmonids in southeast Alaska is terrestrial invertebrate prey
that originates from upstream riparian vegetation. Because headwater streams drain into
larger systems containing salmon, the immediate and long-term effects of clearcuttin g

along headwaters need to be addressed.

Wood is critical to the formation of debris dams and retention of organic materials
in streams (Bilby and Likens 1980, Cummins and Klug 1979), therefore it is important to
understand the strength properties and decay patterns of wood in streams. Wood in
terrestrial environments is broken down by saprophytic fungi and wood-boring
invertebrates causing weight and strength loss (Lesage 2003, Worrall 1999). Aerobic
fungal hyphae do not penetrate saturated wood surfaces and most aquatic invertebrates
that contribute to the degradation of wood do not bore or penetrate deeply (Anderson et
al. 1978, Chergui and Pattee 1991). Therefore, logs similar in size and species decay
slower in streams than in terrestrial environments (Bilby et al. 1999). Studies comparing
the breakdown of red alder and conifer wood in streams of the Pacific Northwest showed
that alder decays more rapidly than conifer, and after three years begins to lose structural
integrity from advanced decay and breakage (Cederholm et al. 1997, Keim et al. 2000).
In Alaska, all types of wood in streams are expected to decay slower than more southern
states due to cooler average air and water temperatures (Wipfli et al. 2002). Interest in
wood decomposition in aquatic environments has primarily involved the importance of
wood strength and decay on the stream structure and fish habitat. Less is known about the
role of decaying wood as a source for macroinvertebrate community recruitment and

colonization.

15

My overall objective was to assess the ecological role of red alder in benthic
macroinvertebrate community structure in headwater streams of regenerating forests in
southeast Alaska. I compared and contrasted macroinvertebrate density, biomass, and
diversity associated with red alder and conifer wood in headwater streams with a range of
riparian red alder. The hypotheses I tested were: 1) all macroinvertebrate community
attributes would be greater on red alder than on conifer wood; and 2) these same
attributes would increase as the presence of riparian red alder increased. In addition to
the role of alder in and along streams, I was interested in potential effects that the stages
of wood decay might have on benthic macroinvertebrate communities. I tested the null
hypothesis that there is no difference in macroinvertebrate density, biomass and diversity

on pieces of wood in early stages of decay versus wood in more advanced decay.

METHODS/MATERIALS

I. fland Selection & Studv Sites

Research was conducted on Prince of Wales Island, southeast Alaska (132°67’W,
55°49’N)(Figure 2). This region contains more extensive tracts of virgin, old growth
forest than anywhere else in the United States (USDA 1997) and offers exceptional
opportunities to study ecological interactions between red alder, site conditions and
stream communities in an evolutionary context. Study sites were in the Maybeso Creek
and Harris watersheds (46 km2 and 108 kmz, respectively) on the eastern end of Prince of

Wales Island (Figure 3). This area was heavily logged in the 19503, and in the 19603

16

local and federal governments established the Maybeso Experimental Forest to study the
impacts of commercial logging on wood production, wildlife, and fisheries (Wipfli et al.
2002). The varying amounts of riparian alder and standing old growth within the
Maybeso Experimental Forest made this an ideal site to complete our research objectives.
Sampling sites for this project were limited to fishless headwater streams within forested
stands of 5-10 ha with elevations less than 150 m. The primary sample site selection

criterion was riparian tree species composition to provide a range of 0-53% red alder.

Aerial photographs, US. Forest Service district files and field visits were used to
select 13 headwater streams for this study. To reduce variability, all streams were 3rd
order or less, had channel slopes between 10-28 degrees, and bankfull widths between
0.7-3.4 m (Wipfli et al. 2002). Stream flow in 2002 ranged from 1-116 L's'l (Table 1).
Sampling was restricted to 250-300 In reaches within each stream. Riparian understory
vegetation and tree species composition was measured within the designated sampling
area at each stream as described by Wipfli et. al. (2002). Salmonberry (Rubus spectabilis
Pursh.), Devils Club (Optopanax horridum Miquel.), Skunkcabbage (Lysichitum
amerz’canum Hutten and S. J .), ferns, and mosses dominated the riparian understory.
Western hemlock (Tsuga heterophylla (Raf.) Sarg.), Alaska yellow cedar
(Chamaecyparis nootkatensis (D. Don) Spach.), and Sitka spruce (Picea sitchensis
(Bon g.) Carriere.) were common coniferous species, and the dominant hardwood was red
alder. Streams were from stands with mixed alder-conifer vegetation that represented a

range of riparian red alder from 0-53 % (Table l).

17

H. Wood Collection & Experimental Design

In order to survey the macroinvertebrates associated with wood, pieces of
naturally occurring alder and conifer (approx. 5 x 20 cm) representing early and late
decay classes were randomly collected from all 13 streams (except one site that had no
alder) during May and July of 2001 and 2002. A piece of wood was considered late
decay if a thumbnail could easily penetrate the surface—all others were considered early
decay. In addition to collecting macroinvertebrates from naturally occurring wood, we
also collected macroinvertebrates from wood that had been secured to substrate holders

and was experimentally added to streams as describes below.

Wood to be secured to substrate holders was collected from standing snags within
the Maybeso Experimental Forest. Boles from alder and conifer were initially divided
into early and late decay classes based on color, weight, and presence or absence of bark
and branches. Boles had uniform diameters ranging from 3 - 5 cm and lengths of 1 - 6 m.
The longer pieces were later cut into 20 cm sections. The middle section and end pieces
from each bole were dried, weighed, and water displacement was used to test for
consistency across the density gradient and to confirm the stage of decay. Boles that
showed irregularities in density and sections containing knots were excluded from the
study. Wood pieces representing four treatments: early decay alder, late decay alder,
early decay conifer, and late decay conifer were then randomly arranged and secured to
plastic substrate holders (Figure 4). A total of nine substrate holders were placed in each

of six streams in a randomized complete block design. The sub-plot factor was wood

18

type and consisted of the treatments: EA, LA, EC, and LC. The whole-plot factor was
riparian alder-conifer species composition and represented a range of riparian red alder
from 1.5 - 47%. Substrate holders were placed in a variety of habitats, however
placement was often restricted to shallow pools due to low water levels. All holders were
set July 7-14, 2000 and remained in the streams for either 1 or 2 yrs. Three substrate
holders from each site were randomly selected and collections took place in early July of

2001 and 2002.

III. Macroinvertebrate Samplin g and Aflvsis

Macroinvertebrates were sampled from both added wood substrates and naturally
occurring woody debris. Individual pieces were pulled from the stream and transferred to
a 19 L bucket for processing. Each piece of wood was rinsed with a hand-pumped
portable pressurized sprayer and the contents of the bucket were strained through a 250-
micron sieve. Invertebrates were washed into 250-ml Whirlpaks® with 80-percent
ethanol and returned to the lab for sorting under a dissecting microscope.
Macroinvertebrates were picked from each sample and identified to the lowest taxonomic
unit possible using Merritt and Cummins (1996a). Measurements of the total length were
made for each identified specimen and length-weight regressions were used to estimate
individual biomass (Benke et al. 1999, Hodar 1996, Smock 1980). Wood surface area
was estimated from length and diameter using the equation for the surface area of a
cylinder, and macroinvertebrate biomass m2 and number rn'2 were calculated for each

piece of wood. Diversity was measured using the Shannon-Weiner diversity index

19

(Hauer and Resh 1996). Functional feeding group ratios were calculated to help assess

stream ecosystem parameters (Merritt and Cummins 1996b).

Results from the dependant variables of invertebrate density, biomass m2, and
diversity were tested for normality, log (x + 1) transformed when necessary, and analyzed
for significance using SAS (SAS Institute 1996). Multiple T-tests and ANOVA were
generated contrasting the dependant variables with percent riparian alder, wood type
(conifer or red alder), and decay class (early or late decay). Percent riparian alder was
grouped into three bins of low (0-9%, n=4), medium (IO-35%, n=5) and high alder (36-
53%, n=4) and Bonferroni adjustments were made for the analysis. Pairwise
comparisons were generated for all dependant variable results with a significant ANOVA
(p < 0.05)(Sokal and Rolf 1969) using Tukey’s Studentized Range (HSD) post-hoe test.

All graphs and tables are presented using nontransformed data.

RESULTS

Macroinvertebrates Community Composition & Functional Feeding Groups

 

Over 10, 000 macroinvertebrates from 14 different orders were recorded from
wood samples. At least 54 insect taxa were identified, but of those taxa a relatively small
number comprised most of the total count and biomass densities (Table 2). Total
chironomid taxa comprised 56% of the total relative abundance, and chironomids in the

subfamily Orthocladiinae accounted for 70% of all chironomids (Figure 5). Nearly half

20

of the total count biomass was from only two genera—a heptageniid mayfly, Cinygma
and a rhyacophilid caddisfly, Rhyacophila. Functionally, collector-gatherers
(Chironomidae, Baetidae, Leptophlebiidae) were the most abundant group collected
(71%), followed by predators (Rhyacophilidae, Chloroperlidae, Empididae)(11%),
shredders (Nemouridae, Leuctridae)(10%), scrapers (Heptageniidae)(8%), and collector-

filterers (Simuliidae)(1%)(Figure 6).

Overall, functional feeding group colonization of alder and conifer wood was not
different. A slight increase in scrapers on alder wood (79%) compared to conifer wood
(59%) was observed in May 2001, however, by July 2001 the relative abundance of
scrapers was similar (58%). During the same period in 2001, the relative abundance of
predators on alder increased from 6% in May to 36% in July. This was primarily due to
an increase in numbers of Rhyacophila on pieces of alder wood. The relative abundance
of predators and the caddisfly Rhyacophila were consistant on conifer wood throughout
2001 and 2002 (34%). The amount of riparian alder did appear to influence the
distribution of some feeding groups and individual taxa. Over the entire sampling period,
collector-gatherer relative biomass increased from 9% to 20% as the percentage of
riparian alder increased from low (0-9%) to high (36-53%). The realtive biomass of
collector-filterers, exclusively prosimulium, also increased with gretaer amounts of
riparian alder. The relative biomass of scrapers, on the other hand, decreased from 71%
to 56% as the percentage of riparian alder increased, and shredder biomass also decreased
as the amount of riparian alder increased. Total predator distributions did not appear to

be influenced by the presence of riparian alder in this study (Figure 7); however, the

21

caddisfly Rhyacophila was positively correlated with an increasing presence of riparian

alder.

II. Macroinvertebgte Density, Biomass & Diversity

Overall, streams with more riparian alder generally supported greater density and
biomass of macroinvertebrates than did streams with low riparian alder. The densities of
macroinvertebrates associated with natural wood were significantly greater (p> 0.05) in
streams with high percentages of riparian alder (>36%) than in streams with low riparian
alder (<10%)(Figure 8A). Densities of macroinvertebrates collected from added wood
were also greater in streams with more riparian alder, however these differences were not
significant (p< 0.05). Likewise, the mean biomass of macroinvertebrates collected from
added wood was greater in streams with more alder, but were not significantly different.
The presence of riparian alder did have an effect on the biomass of macroinvertebrates
collected from naturally occurring wood (p< 0.05; Figure 8B), but variability among
groups led to difficulty in clearly establishing differences among individual levels of
alder. The diversity of macroinvertebrates collected from added wood was slightly
greater in streams with more riparian alder, however, these differences were not
significant and the diversity of macroinvertebrates collected from natural wood showed

no differences with lower or higher amounts of riparian alder.

22

Wood type and decay class had some effect on benthic macroinvertebrate density,
biomass, and diversity. Both density (Figure 9A) and diversity (Figure 10A) of
macroinvertebrates collected from natural and added wood substrates were greater on
alder than on conifer, but statistically these differences were not significant. However,
when converted to biomass, naturally occuring pieces of conifer wood supported a
significantly greater mass of benthic macroinvertebrates than naturally occuring alder
wood (Figure 9B; p< 0.01). Mean biomass of macroinvertebrates collected from added
wood substrates was not significantly different and was greater on alder. Overall, pieces
of wood in later stages of decay generally supported greater density, biomass (Figure 11),
and diversity than did pieces of wood in more recent stages of decay, however, only
diversity of macroinvertebrates (associated with natural wood in later stages of decay was

significantly different p< 0.05)(Figure 10B).

DISCUSSION

Results from naturally collected wood showed that the presence of riparian alder
in second-growth forests appeared to influence the density of benthic macroinvertebrates
associated with wood surfaces in southeast Alaskan headwater streams. Anderson et al.’s
(1978) findings in two Oregon watersheds were similar and he suggested that the higher
in-stream macroinvertebrate densities were due increased organic inputs from alder
vegetation. Culp and Davies (1985) also found higher macroinvertebrate abundance
associated with alder litter. On Prince of Wales Island, Hernandez et al. (2004) found

that streams with alder-dominated young-growth riparian vegetation supported higher

23

densities of benthic macroinvertebrates on gravel, cobble, and wood substrates. Wipfli
and Musselwhite (2004) found that the density of macroinvertebrates collected in 24-hour
drift samples increased with the presence of riparian alder, and Wipfli and Gregovich
(2002) indicated that the abundance of terrestrial insects collected from stream drift
significantly increased as the percent basal area of alder increased along riparian zones.

In a study concurrent with this one, Lesage et al. (2003) compared macroinvertebrates
colonizing terrestrial litter and wood along the same streams. Results from their study
showed that invertebrate density, taxa richness, and biomass increased when the

percentage of riparian alder increased.

Naturally collected wood debris from streams also showed an increase in
macroinvertebrate biomass associated with an increasing presence of alder in riparian
forests. This increase in biomass may be the result of greater nutrient availability. Alder
is a better food source for herbivorous invertebrates and aquatic shredders than is conifer
litter (Irons et al. 1988, Webster and Benfield 1986) and alder leaves are processed more
quickly than conifer needles (Cummins et al. 1989, Lesage et al. 2003, Sedell et al.
1975). Streams with alder in riparian forests deliver more alder litter to
macroinvertebrate communities than streams with more conifer and this may lead to
increased production of benthic macroinvertebrates (Stone and Wallace 1998). Piccolo
and Wipfli (2002) identified alder in forested riparian zones as critical habitat for
terrestrial invertebrates that fall prey to fish, and Wipfli (1997, et al. 2002) found that
over half of the prey biomass ingested by juvenile salmonids in southeast Alaska was

drifting terrestrial invertebrates that originated from upstream riparian vegetation. In an

24

ensuing study, Wipfli and Gregovich (2002) found that the biomass of drifting
macroinvertebrates in headwater streams increased as the presence of alder increased in
riparian forests. In southeast Alaska, where freshwater ecosystems appear to be nutrient
limited (Ashley and Slaney 1997, Wipfli and Gregovich 2002), prey biomass generated
from riparian alder vegetation and its associated litter may improve stream productivity.
Past management practices in southeast Alaska include removing alder from regenerating
forests, however results from recent studies in the Pacific Northwest and southeast
Alaska show that including an alder component to young-growth upland forests generates
more prey biomass to downstream salmon and may increase salmonid production
(Hernandez et al. 2004, LeSage 2003, Wipfli and Gregovich 2002, Wipfli and

Musselwhite 2004).

This research did not show a relationship between macroinvertebrate diversity and
an increasing presence of alder in riparian habitats on either added wood substrates or
naturally collected wood in headwater streams. Previous studies in the Pacific Northwest
have shown that benthic macroinvertebrate diversity was greater in old-growth than in
either recently clearcut or second-growth forest riparian habitats (Anderson 1992,
Newbold et al. 1980). However, on Prince of Wales Island, Hernandez et al. (2004) found
that the diversity of stream macroinvertebrates was significantly greater in alder
dominated second-growth than in old-growth watersheds and suggested that the greater
diversity was due to changes in food resources and the availability of more labile
allochthonous inputs generated from deciduous alder vegetation. Lesage (2003) found

that terrestrial invertebrate richness and diversity was significantly greater in riparian

25

habitats where alder was present and concluded that increasing the presence of alder may
contribute to increased production of higher tropic levels. We did not find a difference
in macroinvertebrate diversity, however the mean biomass of the trichopteran
Rhyacophila (Rhyacophilidae)—the most common predator collected—did increase with
the presence of riparian alder. This suggests that the presence of alder in riparian
corridors may contribute to an increased production of specific higher trophic levels in

aquatic organisms.

Macroinvertebrate colonization of wood surfaces showed that mean biomass was
significantly greater on conifer compared to alder wood debris. The greater biomass on
conifer wood was primarily due to an increase in the presence of two taxa: the
heptageniid mayfly Cinygma and the rhyacophilid caddisfly Rhyacophila. Mayflies in
this genus are largely restricted to wood substrates (Anderson et al. 1978), and were
observed in this study scraping wood surfaces—presumably feeding on periphyton and
detritus. The caddisfly Rhyacophila, one of the larger invertebrates collected, was often
found associated with protective niches on wood surfaces and may have been more
common on conifer due to its cracking nature during decomposition (Wipfli et al. 2003).
Both mean density and diversity of macroinvertebrates were greater on alder, however
differences were not statistically significant due to large variation between samples.
Contributing to the variation in mean densities between conifer and alder were higher
counts of midge taxa on some individual pieces of alder. Because chironornids tend to
be ubiquitous, it is difficult to identify the specific role alder plays in their colonization,

but the presence of xylophagous midges in the subfamily Orthocladiinae and large

26

numbers of pupae and tubes of midges in the subfamily Tanytarsani, suggests that the
wood surfaces are being used as both food source and refuge. Results from this study did
indicate that macroinvertebrate colonization of wood surfaces appears to be more closely

associated with stage of decay than wood type.

Overall, macroinvertebrate density, biomass and diversity was greater on wood in
advanced stages of decay. Dudley and Anderson (1982) suggested that wood as a food
resource becomes available to more taxa as surface decay progresses, and Golladay and
Webster (1988) found that functional feeding groups using wood were directly associated
with the decay of wood surfaces. There was evidence of feeding on wood in our study,
however most of the macroinvertebrates colonizing wood were probably using this
habitat for resting, refuge, and oviposition. Wood in later stages of decay provided more
area of this habitat. Evidence that the decay of wood surfaces may be a greater
contributor to macroinvertebrate colonization than wood type was found in the results of
our experiments with added wood substrates. Although two decay classes of conifer and
alder were used as added wood, the surfaces of all wood pieces used were smooth and
very similar structurally. The similarities in wood surfaces may explain why there were
no differences in colonization between wood types. The rhyacophilid caddisflies
commonly collected within cracks and crevices of natural wood was further evidence that
decay influenced colonization patterns. Anderson et al. (1978) described several taxa
from streams in the Pacific Northwest that used the protective niches on wood surfaces
for oviposition, as a nursery for early instars, molting, pupation, emergence, and as a

source of prey. McLachlan (1970) found that mayflies more readily colonized

27

submerged wood with bark previously damaged compared to newer pieces with solid
bark. High densities of baetid mayflies were collected from wood surfaces in our study,
and both conifer and alder wood classified as late decay supported greater mean densities.
Whether the decaying process conditions wood surfaces for feeding or creates more
interstitial spaces for refuge, it appears as though wood with greater surface decay and
more available interstitial spaces are more attractive to benthic macroinvertebrates.
Managing young-growth forests to include Red alder may improve benthic
macroinvertebrate habitat, because alder generally dies standing (Wipfli et al. 2002) and
produces more litter in advancing stages of decay than do conifers (Zavitkovski and

Newton 1971).

Changes in macroinvertebrate functional feeding groups shifted in relation to
riparian alder vegetation. Collector-gatherer communities, dominated by midge taxa and
the baetid mayfly (Baetis sp.), gradually increased as the percentage of riparian alder
increased. Increases in collector-gatherer groups have been associated with increased
loads of fine particulate organic matter (Cuffney and Wallace 1989, Cumrrrins 1973,
Richardson and Neill 1991), and inputs from riparian alder may contribute to this nutrient
load. Scraper populations, on the other hand, decreased with more riparian alder. In the
light-limited small streams of the Pacific Northwest, clearcutting can result in increased
solar radiation (Bisson and Bilby 1998), which also increases primary production and the
abundance of scrapers (Hernandez et al. 2004, Murphy et al. 1981, Wallace et al. 1988).
The lower percentage of scrapers found in headwaters on Prince of Wales Island was

comparable to results found in other headwater streams in regenerating forests of the

28

Pacific Northwest, most likely due to canopy closure (Cole et al. 2003, Haggerty et al.
2004). Although it was difficult to determine the specific role of alder on
macroinvertebrate functional group relationships from this study, changes in feeding
group compositions are typically the result of changes in food type and its availability
(Merritt and Cummins 1996b, Vannote et al. 1980). It does appear that increased organic
material from riparian alder vegetation provides more food resources for some

macroinvertebrate taxa in headwater streams on Prince of Wales Island.

CONCLUSION

Managing upland forests to include a red alder component may provide more
suitable conditions for macroinvertebrate communities and therefore, improve the
potential of nutrients to reach downstream ecosystems. In southeast Alaska, fresh water
ecosystems are generally nutrient limited (Ashley and Slaney 1997, Wipfli 1997) and the
presence of red alder in riparian habitats could protect or even improve the productivity
of aquatic organisms—including downstream salmonids. Current forest management
plans for southeast Alaska include clearcutting and alder-thining within headwater
habitats (USDA 1999), which may greatly effect macroinvertebrate communities and
food resources for juvenile Pacific salmon (Hayes et al. 2000, Hernandez et al. 2004,
Resh et a1. 1988, Wipfli and Gregovich 2002). Recent declines in Pacific salmon returns
have increased consideration for the development of new policies which protect salmon
and all forest resources. Results from this study, and recent findings in the Pacific

Northwest, suggest that provisioning for red alder in forested uplands should be

29

considered as a management tool to improve downstream salmonid production in

southeast Alaska, and elsewhere.

30

Table 1. Physical and biological characteristics of headwater streams

and adjacent riparian forest on Prince of Wales Island, Alaska, 2000.

 

 

Stream Channel Bankfull Stream Basal
Slopel Widthz Flow3 Area‘
(deg.) (m) (L-s.")3 (% red alder)
Lost Bob 19.5 0.7 7.4 0.0
Upper Good Example 13.6 0.9 3.0 1.5
Cedar 2 18.8 1.7 4.5 3.8
Upper Morning 14.5 1.1 4.7 3.9
Creature Creek 15.2 1.1 36.0 10.0
Gomi 26.1 1.0 5.5 25.0
Mile 22 18.3 1.2 1.5 29.2
Big Spruce 26.3 0.9 0.9 31.5
Cedar 1 19.3 1.6 16.0 35.6
Broken Bridge West 22.5 1.5 3.9 38.7
Cotton 272* 3.4 116.2 43.2
Broken Bridge East 107* 2.2 24 47.3
Brushy 15.5 0.9 1.4 53.3

 

I Channel slope gradient of entire stream; * measured on 25% of stream length
only.

2 Bankfull width — the dominant channel forming flow (Dunne and Leopold 1978).
3 Mean stream flow at transition zone.

4 Percent red alder as a proportion of total stand basal area.

31

Table 2. Checklist of taxa collected and the percent relative density and biomass

for each taxon collected. Totals are combined over the entire study period.

 

% Relative Abundance

 

 

Taxon Numbers Biomass
Collembola
Entomobryiidae 0.7 0. 1
Hypogastruridae <0. 1 <0.1
Sminthuridae 0. 1 <0.1
Coleoptera
Dytiscidae, Liodessus <0.1 <0.1
Elmidae, Narpus <0.1 <0.1
Hydrophilidae, Ametor 0.1 0.5
Ptilodactylidae, Araeopidius <0.1 0.1
Diptera
Ceratopogonidae, C eratopogon <0. 1 <0.1
Ceratopogonidae, Probezzia 0.2 <0.1
Chaoboridae, Eucorethra <0.1 <0.1
Chironomidae 1.2 0.2
Chironomidae, Chironominae 4.2 1.1
Chironomidae, Diamesinae 0.1 <0.1
Chironomidae, Orthocladiinae 39.5 1.3

32

Table 2. (cont.)

Chironomidae, Tanypodinae 2.2 0.6
Chironomidae, Tanytarsani 10.8 2.2
Dixidae, Dixa 0.7 <0.1
Empididae, Chelifera 3.5 0.4
Empididae, Clinocera <0.1 <0.1
Empididae, Oregeton <0.1 <0.1
Psychodidae, Pericoma <0. 1 <0. 1
Pyralidae <0. 1 <0. 1
Sciaridae <0.1 <0.1
Simuliidae, Prosimulium 0.9 3.0
Tipulidae, Dicranota 0.3 0.1
Tipulidae, Hexatoma 0.1 <0.1
Tipulidae, Limonia <0.1 <0.1
Tipulidae, Molophilus <0.1 <0.1
Ephemeroptera
Ameletidae, Ameletus <0.1 <0.1
Baetidae, Baetis 6.7 7.3
Ephemerellidae, Drunella 0.8 1.6
Heptageniidae, Cinygma 4.1 37.6
Heptageniidae, Cinygmula 0.3 19.3

33

Table 2. (cont)

Heptageniidae, Epeorus 1.8 0.4
Heptageniidae, Ironodes <0. 1 1.0
Heptageniidae, Rhithrogena <0.1 <0.1
Leptophlebiidae, Paraleptophlebia 6.4 2.6
Plecoptera
Chloroperlidae, Suwalia 0.1 0.1
Chloroperlidae, Sweltsa 0.9 1.0
Leuctridae, Despaxia 1.3 0.9
Nemouridae, Zapada 7.0 1.7
Trichoptera
Brachycentridae, Micrasema 1.3 0.2
Goeridae, Goeracea 0.2 0.2
Hydropsychidae, Arctopsyche 0.1 0.7
Limnephilidae, Allomyia <0.1 <0.1
Limnephilidae, Chyranda <0.1 0.5
Limnephilidae, Cryptochia 0.2 0.1
Limnephilidae, Ecclisiomya <0.1 <0.1
Limnephilidae, Moselyana <0.1 <0.1
Limnephilidae, Psychaglypha 0.2 1.5

34

Table 2. (cont.)

 

Philopotamidae, Dolophiloides 0.2 0.1
Philopotamidae, Wormalidia <0.1 <0.1
Ryacophilidae, Rhyacophila 3.4 13.6
Uenoidae, Neophylax 0.2 0.3
54 total taxa 100 100

35

, '~. . _ I . , . _
Second-growth forest- . ~. ‘ v" ‘ g

t ' \
-l-I

5\ r 7Headwater streams
If // x

Former logging roads

\

Land slides
v .v - my

 

Figure 1. Red alder colonization following soil disturbance along cleared stream
banks, abandoned logging roads and landslides within the Maybeso Experimental

Forest on Prince of Wales Island, southeast Alaska.

36

Alaska Canada

 

IS

 

Maybeso
Experimental
Forest,
Prince of
Wales Island

 

 

 

 

Figure 2. Research was conducted within the Maybeso Experimental Forest on eastern

Prince of Wales Island, southeast Alaska (132°67’W, 55°49’N).

37

 

 

 

Prince of Wales Island _

 

 

 

 

 

  

 

Maybeso & Harris Drainages I
s l
a N P
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' ’ - ‘ " ’ LEGEND
\ ,’
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' ' 108 km2 ' ' ' " Drainage boundary

 

 

 

 

 

 

Figure 3. Study sites within Maybeso Creek and Harris River drainages on

Prince of Wales Island, SE Alaska (132°67’W, 55°49’N).

38

 

 

 

| Prince of Wales Island

Maybeso & Harris Drainages
7% Maybeso Creek ,' I ~ .

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38

Figure 3. Study sites within Maybeso Creek and Harris River drainages on

Prince of Wales Island, SE Alaska (132°67’W, 55°49’N).

 

20 cm

 

 

Figure 4. Macroinvertebrates were collected from the surfaces of wood pieces
representing four treatments: early decay alder (EA), late decay alder (LA), early
decay conifer (EC) and late decay conifer (LC). Pieces were randomly arranged and

secured to plastic substrate holders.

39

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10%
\
Predators
1 1%
Dominant Families Within Feeding Groups:
Collector- atherers Predators Shredders Scra ers Collector-fllterers

 

Chironorrridae Rhyacophr'lidae Nemouridae Heptageniidae Simuliidae
Baetidae Chloroperlidae Leuctridae

Leptophlebiidae Empididae

 

Figure 6. Relative abundance of functional feeding groups and the dominant families

within each of the five feeding groups represented.

41

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m E
5 g 400
0 .—
2 e
g 200
no
5 0
Low Med. High
Alder Alder Alder

Figure 8. Mean macroinvertebrate density (A) and biomass (B) on naturally
occurring wood in streams with low (0-9%, n=4), medium (IO-35%, n=4), and
high (36—53%, n=5) percentages of riparian red alder. Means with * are
significantly different (p< 0.05). Percent riparian alder did significantly
influence biomass (p=.047), but the individual levels of alder could not be

identified.

2000 : A)

 

 

1500 i

i

500 «

 

 

 

 

 

 

Count denSIty
(no. macroinvertebrates m2 t SE)

1000 ' B)

800 - *

 

§

400 ‘

29’

 

 

 

 

O

‘ ;

Alder Conifer
Wood Wood

 

 

Biomass densrty
(mg macroinvertebrates m2 t SE)

Figure 9. Mean macroinvertebrate (A) count and (B) biomass collected from the
surfaces of naturally collected alder and conifer wood between 2001 and 2002.

Means with * are significantly different (p=0.05).

3.5 - A)

 

 

 

 

 

 

 

 

 

 

 

 

3.0 -
2.5 -
LIJ 2.0 s
(D
“ ’5? 1.5 .
é‘S
9 c
a) — 1.0 ~
.2 3‘
O z)
9 g, .5 I
m ._
a9 .
t: 0:3 Alder Conifer
G) ._
E G.)
E: 35 - B)
0 c
(U _
2 g 3.0 *
o
“g 3,5, 2.5 -
g v
CD 2.0 r
1.5 -
1.0 r
.5 ‘
0

 

 

 

 

 

 

Recent Late
Decay Decay

Figure 10. Shannon-Weiner diversity on (A) naturally occurring alder wood versus
naturally occurring conifer wood and (B) recent versus late decay classes of naturally
occurring wood pieces. Totals are from wood collected over the 2001 and 2002

sampling seasons. Means with * are significantly different (p=0.05).

45

3000 I (A)

2500 ~

2000-

 

1500i

1000—

Count densrty
(no. macroinvertebrates m2* SE)

 

 

 

 

Biomass densrty
(mg macroinvertebrates m2*SE)

Recent Late
Decay Decay

Figure 11. Mean macroinvertebrate (A) density and (B) biomass collected from
naturally occurring alder and conifer wood in recent and late stages of decay. Results

based on collections made between 2001 and 2002.

46

LITERATURE CITED

47

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55

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.: 2004-05

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

Macroinvertebrate Community Response to Riparian Red Alder within
Headwater Streams of Second-Growth Forests in Southeast Alaska

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

Entomology Museum, Michigan State University (MSU)

Other Museums:

lnvestigator's Name(s) (typed)
Ryan K. Kimbirauskas

 

 

Date December 13. 2004
*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.

56

Appendix 1.1

Voucher Specimen Data

 

 

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Appendix 1.1

Voucher Specimen Data

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58

Appendix 1.1

Voucher Specimen Data

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0 FOON >50 000505 0002.0... 0.50.000 0050002500
5000000000
0F .OON >50 0000.05 0000>05. 030.000.020.00. 005.000.000.000
N FOON >50 0000.05 0000>05. 00000.50... 005500200...
F NOON >5... 0000.05 0002.05. 000000... 005500050...
0005.000 .<mm...00¢m§wz0m
000.0. 00.0.5 .0 005.039.... 6.0:
or. 02.00000 000 0000
6 .
.m d .0 00.00.50 00050000 .2 0.00 .0000 00x0. .020 .0 00.0000
0 .0 a 2 .0 e
0.. r m m. e D.
0 m 0 0 m m m 0
M .m 0 A A P N m E
”.0 .0252

 

59

Appendix 1.1

Voucher Specimen Data

Ema .9930

 

.Ezmmaz >mo_oEoEm

36.52:: 29w 59:22 9: E «688
.2 mcoE_8am 35: 9on 05 8282“.
moéoom .oz 85:;

 

 

38 2 c8588 25

 

 

mmxngnéx .x :85

638 @252 228582.
3388: = $85 .mcoEnum $3

 

 

Page 4 of 4 Pages

58 22.
38 22.
58 22.
68 22.
58 22,
«SN 22.
«com 22.
58 22.
58 22.
58. 22.
68 22.

Y—Y‘V—FF v-v-C'DV-NCO

03590 08ng
mom—=90 08ng
39:90 omega—2
mmmEmE omega—2
womESQ Omega—2
@9590 08ng
3235 80ng
3,285 Omega—2
$9590 0896.2
39.55 03ng
39:90 833.2

956. 3.25 .o 8.5%. .x< .<w:

92382.”. 323383.“.
«635.03 3289822.”.
mz§.mmzo>mn_ magzamcE:
wSEomzoow mmv___;am:E_._
«28:50 $2.283:
35;ch $2.285:
£522 $2.285:
magmamaa omEcgmaofif
223882 322963.31
«8880 98:80
mEmmSoS mmchmoEoSm
<mmEOIOEP

 

Museum where

deposited

 

Other

100+ m 6

MM. % 0.8 S
U u pm N 9
dd U ya 9
AAPNLE

 

 

 

 

 

 

$2388 98 8m:
5 382.8 mcoE_o¢am .2 Emu .23

 

:93. 850 .o 86me

 

 

 

”.6 69:32

 

60

   

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