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BENTHIC INVERTEBRATE COMMUNITY STRUCTURE AS AFFECTED
BY FOREST SUCCESSION AFTER CLEAR-CDT LOGGING ON
PRINCE OF WALES ISLAND, SOUTHEAST ALASKA

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6’01 cJCIRC/DmoDuepes—n 15

BENTHIC INVERTEBRATE COMMUNITY STRUCTURE As AFFECTED BY
FOREST SUCCESSION AFTER CLEAR-CUT LOGGING ON PRINCE OF WALES
ISLAND, SOUTHEAST ALASKA
By

Osvaldo Hernandez

A THESIS
Submitted to
Michigan State University
in partial fulﬁllment of the requirements
for the degree of
MASTER OF SCIENCE
Department of Entomology

2001

ABSTRACT
BENTHIC INVERTEBRATE COMMUNITY STRUCTURE AS AFFECTED BY
FOREST SUCCESSION AFTER CLEAR-CUT LOGGING ON PRINCE OF WALES
ISLAND, SOUTHEAST ALASKA
By

Osvaldo Hernandez

The primary industries of Prince of Wales Island are toursim, timber harvesting
and sport and commercial ﬁsheries. Because timber harvesting is a disturbance that
affects both physical and biological characteristics of adjacent streams and rivers, the
effects of clear-cutting on spawning and rearing habitats of commercial and sport ﬁsh
species have been investigated. However, many small ﬁshless headwater streams in
upland forests, with potential sources of benthic invertebrates as major food items for
economically important ﬁsh species have received little attention. In an effort to assess
the effects of timber harvest practices in upland forests, benthic invertebrate community
structure was contrasted among four dominant forest management conditions and
instream habitats. Timer harvest caused increases in invertebrate richness, densities and
biomass relative to old grth conditions, particularly in second growth managements
with an alder-dominated riparian vegetation. Large woody debris and gravel habitats
supported high densities and biomass of invertebrates. In addition, large woody debris
also supported a richer and more diverse fauna than either cobble or gravel substrates.
Alternatives to clear-cut harvesting should be employed for the maintenance of wood

recruitment into streams and grth of red alder along riparian margins.

Este trabajo es para mi familia que se encuentra muy lejos de mi y que quiero mucho.
También para mis gran amigos Robert y Hook que me dieron mis primer experiencias en

las ciencias.

iii

ACKNOWLEDGMENTS

I would like to thank Michigan State University’s Entomology department for the
opportunity to ﬁnther my education, and my committee members Dr. E. Graﬁus, Dr. M.
Kaufman, Dr. M. S. Wipﬂi, and Dr. R. W. Merritt who provided excellent guidance. I
also thank Derek Bush, Takashi Gomi, and Brittany Graham for help in the ﬁeld, and
Edward McCoy for help in the lab. I would also like to thank a number of contributors
who funded my tuition and living expenses, in part, while at Michigan State University:
Urban Affairs Program, College of Natural Sciences, Department of Entomology, and the
Bill and Melinda Gates Foundation. Lastly, I thank the Paciﬁc Northwest Research
Station, USDA Forest Service, Juneau, AK 99801 USA, for research funding and

logistical support.

iv

TABLE OF CONTENTS

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

SUMMARY OF SELECTED STUDIES EXAMINING FOREST HARVEST
EFFECTS ON STREAM ATTRIBUTES IN NORTH AMERICA

Introduction .......................................................................................................................... 1
Eastern North America ............................................................................................... l
Coweeta Hydrologic Laboratory ................................................................................. 3
Western North America .............................................................................................. 5
Alsea Watershed Study ............................................................................................... 5
HJ. Andrews Experimental Forest ............................................................................. 6
Washington State ........................................................................................................ 7
British Columbia ......................................................................................................... 8
Alaska ......................................................................................................................... 9

Literature Cited .................................................................................................................. 14

CHAPTER 2

BENTHIC INVERTEBRATE COMMUNITY STRUCTURE AS AFFECTED
BY FOREST SUCCESSION AFTER CLEAR-CUT LOGGING ON PRINCE OF
WALES ISLAND, SOUTHEAST ALASKA

Introduction ........................................................................................................................ 19
Methods ............................................................................................................................. 21
Study Sites ................................................................................................................ 21
Physical Measurements ............................................................................................. 22
Experimental Design and Analysis ........................................................................... 23
Macroinvertebrate Sampling ..................................................................................... 24

Results ................................................................................................................................ 25

Physical Measurements ............................................................................................. 25
Macroinvertebrate Richness and Diversity Among Management Conditions ......... 26
Macroinvertebrate Densities and Biomass Among Management Conditions .......... 26
Functional Feeding Group Proportions Among Management Conditions ............... 28
Macroinvertebrate Richness and Diversity Among Instream Habitats ..................... 29
Macroinvertebrate Densities and Biomass Among Instream Habitats ..................... 29
Functional Feeding Group Proportions Among Instream Habitats .......................... 30
Discussion .......................................................................................................................... 31
Management Conditions ........................................................................................... 31
Instream Habitats ...................................................................................................... 34
Conclusion ......................................................................................................................... 37
Literature Cited .................................................................................................................. 57
Appendix 1 ......................................................................................................................... 62

vi

LIST OF TABLES

CHAPTER 1
Table 1. Review and summary of selected studies examining forest harvest effects

on stream attributes ............................................................................................................ 11

CHAPTER 2
Table 1. Average daily maximum, minimum, and differences in streamwater

temperature (°C) 1999 ....................................................................................................... 39

Table 2. Percentage of instream large woody debris, cobble and gravel habitats
within management conditions. Do not equal 100 percent in cases where ﬁne

woody debris or bedrock substrates were present .............................................................. 39

Table 3. Checklist of taxa collected from upland forest headwater streams of

Maybeso Experimental Forest and adjacent Harris River watershed Prince of

Wales, Southeast Alaska. ................................................................................................... 40

Table 4. Percent distribution of Chironomidae subfamilies collected from large

woody debris, cobble and gravel habitats in each management condition ........................ 41

vii

LIST OF FIGURES

CHAPTER 2

Figure 1. Maybeso Experimental Forest, Prince of Wales Island, Southeast Alaska ........ 43

Figure 2. Mean algal AF DM among management conditions 1999. Means with

different letters are signiﬁcantly different (p<0.05) .......................................................... 44

Figure 3. Mean richness of invertebrates among management conditions from (a)
1998 and (b) 1999. Means with different letters are signiﬁcantly different

(p<0.05) .............................................................................................................................. 45

Figure 4. Mean Shannon-Weiner diversity of invertebrates among management
conditions from (a) 1998 and (b) 1999. Means with different letters are

signiﬁcantly different (p<0.05) .......................................................................................... 46

Figure 5. Mean densities of invertebrates among management conditions from (a)

large woody debris and cobble habitats 7-15 July 1998; and (b) gravel habitats l 1-

14 June 1999. Means with different letters are signiﬁcantly different (p <0.05). ............. 47

viii

LIST OF FIGURES (CONTINUED)

Figure 6. Mean dry mass of invertebrates among management conditions from (a)
1998 large woody debris and cobble habitats; (b) 1999 large woody debris and
cobble habitats; and (c) 1999 gravel habitats. Means with different letters are

signiﬁcantly different (p<0.05). ......................................................................................... 48

Figure 7. Percentage of functional feeding groups, based on biomass, present

among management conditions 7-15 July 1998 ................................................................. 49

Figure 8. Percentage of functional feeding groups, based on biomass, present

among management conditions 11-14 June 1999 .............................................................. 50

Figure 9. Mean richness of invertebrates among habitats from (a) 7-15 July 1998
and (b) 11-14 June 1999. Means with different letters are signiﬁcantly different

(p<0.05) .............................................................................................................................. 51

Figure 10. Mean Shannon-Weiner diversity among habitats from (a) 7-15 July
1998 and (b) 11-14 June 1999. Means with different letters are signiﬁcantly

different (p<0.05) ............................................................................................................... 52

ix

LIST OF FIGURES (CONTINUED)

Figure 11. Mean densities of benthic invertebrates on large woody debris and
cobble habitats across all management conditions. 7-15 July 1998 and 11-14 June

1999 .................................................................................................................................... 53

Figure 12. Mean dry mass of invertebrates among large woody debris and cobble
habitats. 7-15 July 1998. Means with different letters are signiﬁcantly different

(p<0.05) .............................................................................................................................. 54

Figure 13. Percentage of functional feeding groups, based on biomass, present

among instream habitats 7-15 July 1998. .......................................................................... 55

Figure 14. Percentage of functional feeding groups, based on biomass, present

among instream habitats 11-14 June 1999. ........................................................................ 56

CHAPTER 1

SUMMARY OF SELECTED STUDIES EXAMINING FOREST HARVEST EFFECTS
ON STREAM ATTRIBUTES IN NORTH AMERICA

INTRODUCTION

The effects of forest clear-cutting upon physical and biological characteristics of
streams have been well studied from eastern to western United States and British
Columbia. These studies have been conducted with a number of designs, from before and
after studies to comparative and long-term studies. Often, experimental forests have been
established for this purpose. The following consists of a brief introduction into several
studies at different localities examining forest harvest effects on stream attributes, and are
summarized in Table 1.
EasImNQEthAmCLLQa

Rishel et a1. (1982) conducted a study at the Leading Ridge Watershed Research
Unit in Pennsylvania to investigate changes in stream temperatures. They compared
temperatures of headwater streams whose riparian vegetation was clear-cut and herbicide
treated to streams with a buffer strip and streams with undisturbed riparian vegetation and
found signiﬁcant increases in daily maximum and signiﬁcant decreases in daily minimum
temperatures relative to the reference streams. Diurnal ﬂuctuations increased four times
from the control, and the highest temperature recorded was 32°C.

Silsbee and Larson (1983) investigated numerous physical parameters (Tablel)
and responses of benthic macroinvertebrates in second order streams to clear-cut logging
in the Great Smokey Mountains National Park in Tennessee and North Carolina. Their

treatment streams had been clear-cut in the early 19008 (1905 to 1926) and their control

streams had previously been uncut. They found that unlogged streams had less numbers
of invertebrates than clear-cut streams, and that scraper and shredder functional groups
were dominant in these systems. They attributed these results to differences in the
riparian vegetation community.

Martin et al. (1985) and Noel et al. (1986) conducted studies of the
biogeochemistry of streamwater in New England after a clear-cut. Their approach
involved comparing streams whose riparian vegetation was clear-cut (2 years previous) to
reference streams (uncut for 35 years) within areas of different vegetation communities
(coniferous, Northern Hardwoods and Central Hardwoods). Streams with clear-cut
riparian vegetation within the Northern Hardwood forests showed greatest differences in
stream chemistry after a clear-cut. They found greater numbers of macroinvertebrates
(Ephemeroptera and Diptera) in the clear-cut streams and attributed this to increases in
temperature and light, leading to increases in food supply for invertebrates by
accelerating the mineralization of organic matter as well as by increasing algal growth.

Grifﬁth and Perry (1991) compared leaf pack processing rates in second order
streams ﬂowing through 20 year and 80-year-old forests with differing vegetation types.
The 80-year-old forests were dominated by Red Oak (Quercus rubra), Sugar Maple
(Acer saccharum) and American Beech (F agus grandifolia) while the 20-year-old forest
was dominated by black cherry (Prunus serotina) and black birch (Betula lenta). Sugar
Maple leave leaf packs were used and they found leaf pack processing rates were faster in
the 20-year-old forests, which they attributed to signiﬁcantly greater densities of total

macroinvertebrates, shredders and collector gatherers than in the 80-year-old forests.

In Maine, a 1.2 km section of the east branch of the Piscataquis River was studied
before and after a clear-cut in 1982 for timber harvesting effects on ﬁsh diet and
production (Garrnan and Moring 1993). As part of the ﬁsh diet studies, the investigators
found signiﬁcant changes in the macroinvertebrate community. Mean densities were
higher in the spring and lower in the fall than they were the previous year (before
harvesting). After the harvest, Ephemeroptera, Plecoptera and Odonata were signiﬁcantly
less abundant while Chironomidae abundance increased threefold.

The responses of macroinvertebrates to different degrees of timber harvesting also
have been investigated in pools of small headwater intermittent streams of the Quachita
National Forest in Arkansas (Brown et al. 1997). The authors found signiﬁcantly higher
total densities and signiﬁcantly lower diversity in harvested streams, as well as an
increase in the ratio of shredders to collectors possibly due to increases in CPOM from
harvesting techniques.

W

The effects of clear-cutting on second order streams have been studied
extensively at the Coweeta Hydrologic Laboratory in Franklin, North Carolina. The two
primary streams under study were the Big Hurricane Branch whose watershed was clear-
cut in 1977, and Hugh White Creek whose watershed has been undisturbed since 1924.
Many faunal changes occurred in the benthic macroinvertebrate community that was
sampled on a monthly basis (Webster et a1. 1983). Greater abundances of collector-
gatherer mayﬂies (Baetis spp. and Ephemerella spp.) were found in the logged stream.

The dominant shredder, Peltoperla maria, declined in numbers and was virtually non-

existent within three years of the harvest. The changes in the macroinvertebrate
community were attributed to changes corresponding to their food availability.

Other studies conducted at Coweeta include Gurtz and Wallace’s (1984)
investigation of substrate-mediated responses of macroinvertebrates to logging
disturbance. Of the four substrates (rock face, cobble, pebble and sand) investigated, rock
face was the preferred substrate in the clear-cut stream and cobble was the preferred
substrate in the reference stream. They concluded that larger substrates, requiring more
energy to move, had increased numbers of macroinvertebrates colonizing them. Wallace
and Gurtz (1986) concluded that Baetr's spp. mayﬂies respond quickly to take advantage
of increases in autochthonous production, therefore, being important to energy processing
in the disturbed stream.

Stone and Wallace (1998) investigated the effects of sixteen years of forest
succession on benthic macroinvertebrate community structure, as well as the efﬁcacy of
ﬁve indices in determining recovery. Macroinvertebrate abundance, biomass and
secondary production were greater in the disturbed stream than in the reference stream.
The authors found that the percentage of scrapers initially present increased after logging,
following the increase in algae production, and then declined with the decline in primary
production. The trend in shredders present was an initial decline following the decrease in
allochthonous inputs and successive increase in percentage following the return of
allochthonous inputs. Once again, changes in macroinvertebrate community structure
were directly corresponding to changes in the type and quantity of food available to them.
Of the indices examined, percent Baetis, shredder to scraper ratios, and the North

Carolina Biotic Index showed the greatest ability to detect differences between the logged

and the reference stream. They all showed recovery or no difference between clear-cut
and reference streams after sixteen years of forest succession.
WistemNQrLRAmerica

Newbold et al. ( 1980) investigated the effects of logging, with and without buffer
strips, on numerous streams across northern California. Their ﬁrst objective was to
establish macroinvertebrate community differences between logged and reference sites.
Of three indices of dissimilarity, only Euclidean distance showed a signiﬁcant logging
effect. Diversity of macroinvertebrates was lower and total density was higher in the
logged streams, which the authors attributed to high densities of Baetis, Nemoura spp.
and Chironomidae. Their second objective was to determine what effects buffer strips of
differing widths had on logging effects. They found signiﬁcant logging effects with
narrow (<30m) buffer strips and no signiﬁcant logging effects with wide (>30m) buffer
strips.
W

The Alsea Watershed Study as described by Hall et a1. ( 1987) was a long-term
study of the effects of timber harvesting, with and without buffer strips, on physical and
biological characteristics of headwater streams along the Oregon coast. The sampling was
conducted seven years prior and seven years after timber harvest on three watersheds.
The ﬁrst watershed was completely clear-cut, the second was cut with buffer strips and
the third was left as a control. They found slight physical characteristic changes in the
watershed with buffer strips, and large changes in the suspended sediments, dissolved
oxygen, and temperature in the clear-cut watershed. However, these changes returned to

prelogging levels as riparian vegetation returned. The biological aspect of their study

included periphyton responses (Hansmann and Phinney 1973), however, focused on ﬁsh
population responses to timber harvest (Connolly and Hall 1999).
WWW

Forest and stream interactions have been studied extensively at the H.J. Andrews
Experimental Forest in Oregon. Rothacher (1970) and other follow-up studies (Harr et a1.
1982 and Hicks et al 1991) compared changes between water yield in streams whose
riparian vegetation was entirely clear-cut and another that was patch cut. Within a larger
study looking at community structure of periphyton communities, Lyford and Gregory
(1975) found standing crop and rates of colonization of periphyton to be greater in a
clear-cut section of Mack Creek than in a forested area of the same stream. They believed
light was the limiting factor determining growth rates of algae in those cascade mountain
streams. Hawkins et a1. (1982) addressed the relative importance of differences in
riparian vegetation, instream substrates and gradient in benthic invertebrate communities
of streams at the H.J. Andrews Experimental Forest. The canopy types in their study
consisted of clear-cut, second grth deciduous, and old growth conifer. Substrate
composition varied with gradient. High gradient streams (~ 10%) had primarily boulder
and gravel substrates, while substrates in low gradient streams (~1%) consisted of cobble,
gravel, and sand. They found canopy type to be more important in inﬂuencing
invertebrate communities than substrate composition, with greater abundances of
invertebrates in streams with clear-cut riparian vegetation. Anderson (1992) examined the
inﬂuence of disturbance on invertebrate communities of Paciﬁc North West streams
where be compared aquatic insect adult emergence in a 3rd order stream ﬂowing through:

1) 450 year old coniferous forest; 2) recent clear-cut; and 3) second grth deciduous

riparian canopy 40 years after a clear-cut. He found that streams in old growth forests had
the highest richness and greatest evenness among Ephemeroptera, Plecoptera and
Trichoptera, and biomass of populations were similar across treatments. He also found
strong grazer dominance in the clear-cut streams and a shift to detritivores using
allochthonous materials in the second growth streams. Studies of peak ﬂow responses to
clear-cutting (Thomas and Megahan 1998) and stream temperature responses to forest
harvest (Johnson and Jones 2000) also have been conducted at the Andrews Experimental
Forest.

W:

The effects of forest harvest on stream characteristics also have been studied in
Washington. Bilby and Bisson (1987) compared emigration and production of stocked
coho salmon fry in streams ﬂowing through old grth and recently clear-cut streams.
Bilby and Ward (1991) contrasted the characteristics and function of large woody debris
in streams draining old growth, clear-cut and second growth forests. They measured
abundance, size and species of wood across the three riparian types. In addition, they
documented the number and type of pools associated with large woody debris, the
number of pieces of large woody debris that formed waterfalls and sediment storage
created by large woody debris. The authors found their second growth stands, which were
composed largely of red alder, do not supply enough large woody debris to streams, and
that which is supplied is not as effective at inﬂuencing channel structure as coniferous
large woody debris. Bilby and Bisson (1992) sought to determine whether autochthonous
organic matter was more important to ﬁsh than allochthonous organic matter in streams

draining old grth and clear-cut forests. They measured leaf litter inputs, periphyton,

ﬂuvial organic matter sources (dissolved organic matter, coarse particulate organic
matter, ﬁne particulate organic matter), discharge, water temperature, nutrients, ﬁsh
production and diet, as well as invertebrate drift. Both ﬁsh diet and drift samples
consisted predominantly of invertebrates belonging to scraper and collector-gatherer
functional groups in both old grth and clear-cut streams. Invertebrates relied heavily
on algae and algal-based detritus as a primary food source, lending to the overall
hypothesis that autochthonous organic matter sources are more important to ﬁsh
production in Washington streams (Bilby and Bisson 1992).

Carnation Creek is a small drainage on the west coast of Vancouver Island,
British Columbia, where long-term (15 years) study of timber harvest effects on stream
attributes was initiated in 1971. The study was divided into three phases. The ﬁrst phase,
from 1971 to 1975, was the prelogging monitoring phase. Phase two, from 1976 to 1981,
involved studies during logging and road construction, and phase three consisted of
postlogging monitoring. Three different logging methods were studied including: 1)
leaving a buffer strip along the stream margin; 2) clear-cutting with great care not to
disturb the stream; and 3) clear-cutting without regard for the stream. Hartman et a1.
(1987) provided an excellent synthesis of published Carnation Creek studies up to 1986,
describing changes in physical characteristics of the watershed and responses of ﬁsh to
those changes. Hartman et a1. (1996) followed up the review of published Carnation
Creek results to 1996, and the implication for ﬁsheries managers. In the review they
describe how timber harvesting had negative impacts on macroinvertebrate densities.

Densities of seven select taxa were lower in the clear-cuts and total macroinvertebrate

densities were at 41-50% of prelogging and unlogged control levels. In an effort to
determine how the distribution of macroinvertebrates is affected by interstitial detritus
quality and quantity, Culp and Davies (1985) experimented with four different mixtures
of detritus (no detritus, low hemlock, low alder, high alder) in the main channel of
Carnation Creek. They found that total macroinvertebrate densities were higher in the
low alder treatment and that total biomass of macroinvertebrates was greater in both alder
treatments than in the no detritus and low hemlock treatments.
Alaska

The research focus of timber harvest effects on ﬁsheries in Southeast Alaska has
changed from the 19505 to the present (Gibbons et al., 1987; Murphy and Milner, 1997).
Beginning in the 19505, when timber harvesting began to ﬂourish in southeast, research
focused on harvesting effects on salmon spawning habitat. Post 1960, research switched
focus to harvesting effects on rearing habitat. In the 19903, with revision of the Alaska
Forest Resources and Practices Act, streamside buffer strips were required to protect
salmonid habitat, and research focus will likely be aimed at monitoring these buffer strips
and their effectiveness. Research on the effects of timber harvesting on benthic
macroinvertebrates has determined that changing light levels are important in Southeast
Alaska streams. Increases in algal production following clear-cutting resulted in
increased benthic invertebrate abundance, while the dominant functional feeding group
(collector-gatherer) remains unchanged (Duncan and Brusven 1985). Research on effects
of canopy type on benthic macroinvertebrate and detritus export from headwater streams,
due to previous timber management in Southeast Alaska, (Piccolo and Wipﬂi, submitted)

has shown streams with a young grth (35 yr old) red alder-dominated canopy exported

signiﬁcantly more macroinvertebrates than did streams with young growth (35 yr old)

conifer-dominated canopy.

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LITERATURE CITED

14

Literature Cited

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

Bilby, RE. and Bisson, RA. 1987. Emigration and production of hatchery coho salmon
(Oncorhynchus kisutch ) stocked in streams draining an old-growth and clear-cut
watershed. Canadian Journal of Fisheries and Aquatic Sciences 44: 1397-1407.

Bilby, RE. and Bisson, RA. 1992. Allochthonous versus autochthonous organic matter
contributions to the trophic support of ﬁsh populations in clear-cut and old-
growth forested streams. Canadian Journal of Fisheries and Aquatic Sciences 49:
540-551.

Bilby, RE. and Ward, J .W. 1991. Characteristics and function of large woody debris in
streams draining old-grth clear-cut, and second-grth forests in southwestern
Washington. Canadian Journal of Fisheries and Aquatic Sciences 48: 2499-2508.

Brown, A.V., Aguila, Y., Brown, KB, and Fowler, W.P. 1997. Responses of benthic
macroinvertebrates in small intermittent streams to silvicultural practices.
Hydrobiologia 347: 119-125.

Connolly, P.J. and Hall, JD. 1999. Biomass of Coastal Cutthroat Trout in Unlogged and
Previously Clear-Cut Basins in the Central Coast Range of Oregon. Transactions
of the American Fisheries Society 128: 890-899.

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, MA. 1985. Energy dynamics of three low-order southeast
Alaska streams: Allochthonous processes. Journal of Freshwater Ecology. La
Crosse 3: 233-248.

Garman, GO and Moring, IR. 1993. Diet and annual production of two boreal river
ﬁshes following clearcut logging. Environmental Biology of Fishes. The Hague
36: 301-311.

15

Gibbons, D.R., Meehan, W.R., Koski, K.V., and Merrell, T.R. Jr. 1987. History of studies
of ﬁsheries and forestry interactions in southeastern Alaska. In Streamside
Management: Forestry and Fishery Interactions. Edited by Salo ED. and T.W.
Cundy. College of Forest Resources, University of Washington, Contribution 57,
Seattle pp. 297-329.

Grifﬁth, MB. and Perry, SA. 1991. Leaf pack processing in two Appalachian mountain
streams draining catchments with different management histories. Hydrobiologia
220: 247-254.

Gurtz, ME. and Wallace, 1B. 1984. Substrate-mediated response of stream invertebrates
to disturbance. Ecology 65: 1556-1561.

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.

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

Harr, R.D., Levno, A., and Mersereau, R. 1982. Sreamﬂow changes after logging 130-
year-old douglas ﬁr in two small watersheds. Water Resources Research 18: 637-
644.

Hartman, G.F., Scrivener, J .C., Holtby, LB, and Powell, L. 1987. Some effects of
different streamside treatments on physical conditions and ﬁsh population
processes in Carnation Creek, a coastal rain forest stream in British Columbia. 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. 330-372.

Hartman, G.F., Scrivener, J .C., and Miles, M.J. 1996. Impact of logging in Carnation
Creek, a high-energy coastal stream in British Columbia, and their implication for
restoring ﬁsh habitat. NationalResearch Council of Canada, Ottawa, ON
(Canada).

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

16

Hicks, B.J., Beschta, R.L., and Harr, RD. 1991. Long-term changes in streamﬂow
following logging in western Oregon and associated ﬁsheries implications. Water
Resources Bulletin 27: 217-226.

Johnson, S.L. and Jones, IA. 2000. Stream temperature responses to forest harvest and
debris ﬂows in western Cascades, Oregon. Canadian Journal of Fisheries and
Aquatic Sciences. 57: 30-39.

Lyford Jr., J .H. and Gregory, S.V. 1975. The dynamics and structure of periphyton
communities in three cascade mountains. Verh.Internat.Verein.Limnol. 19: 1610-
1616.

Martin, C.W., Noel, D.S., and F ederer, CA. 1985. Clearcutting and the biogeochemistry
of streamwater in New England. Journal of Forestry 83: 686-689.

Murphy, ML. and Milner, AM. 1997. Alaska timber harvest and ﬁsh habitat. In
Ecological Studies 119. Edited by AM. Milner and M.W. Oswood. Springer-
Verlag, New York pp. 229-263.

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

Noel, D.S., Martin, C.W., and Federer, CA. 1986. Effects of forest clearcutting in New
England on stream macroinvertebrates and periphyton. Environmental
Management 10: 661-670.

Piccolo, II. and Wipﬂi, MS. 2001. Does red alder (Alnus rubra) in upland forests
elevate macroinvertebrate and detritus export from headwater streams to
downstream habitats in Southeast Alaska. (Submitted to Canadian Journal of
Fisheries and Aquatic Sciences)

Rishel, G.B., Lynch, IA, and Corbett, ES. 1982. Seasonal Stream Temperature Changes
Following Forest Harvesting. Journal of Environmental Quality 11: 112-116.

Rothacher, J. 1970. Increases in water yield following clear-cut logging in the Paciﬁc
Northwest. Water Resources Research 6: 653-658.

17

Silsbee, DC. and Larson, G.L. 1983. A comparison of streams in logged and unlogged
areas of Great Smoky Mountains National Park. Hydrobiologia 102: 99-111.

Stone, MK. and Wallace, J .B. 1998. Long-term recovery of a mountain stream from
clear-cut logging: The effects of forest succession on benthic invertebrate
community structure. Freshwater Biology 39: 151-169.

Thomas, RB. and Megahan, W.F. 1998. Peak ﬂow responses to clear-cutting and roads
in small and large basins, western Cascades, Oregon: A second opinion. Water
Resources Research 34: 3393-3403.

Wallace, J .B. and Gurtz, ME. 1986. Response of Baetis mayﬂies (Ephemeroptera) to
catchment logging. American Midland Naturalist 115: 25-41.

Webster, J.R., Gurtz, M.E., Hains, J.J., Meyer, J .L., Swank, W.T., Waide, J .B., and
Wallace, J.B. 1983. Stability of stream ecosystems. In Stream ecology:
Application and testing of general ecological theory. Edited by J.R.Bames and
G.W.Minshall. Dep. Biol., Virginia Polytech. Inst. State Univ., Blacksburg, VA
24061 pp. 355-396.

18

CHAPTER 2
BENTHIC INVERTEBRATE COMMUNITY STRUCTURE AS AFFECTED BY
FOREST SUCCESSION AFTER CLEAR-CUT LOGGING ON PRINCE OF WALES
ISLAND, SOUTHEAST ALASKA
INTRODUCTION

Clear-cut logging, a timber harvesting method in which the forest canopy is
removed, is a disturbance that affects both physical and biological characteristics of
adjacent streams and rivers. One of the major changes occurring after a clear-cut is the
decrease of the allochthonous energy base (Duncan and Brusven 1985a) on which
headwater streams and benthic macroinvertebrates are dependent (Vannote et a1. 1980).
The effects of the disturbance on benthic macroinvertebrates have either been studied
before and after the harvest (Garman and Moring 1993; Hall et al. 1987; Hartman et al.
1996) or in comparative studies with a reference stream (Anderson 1992; Hawkins et al.
1982; Newbold et al. 1980). However, because the riparian vegetation is brought to a
stage of secondary succession (Alaback 1984), it also has been studied as such (Stone and
Wallace 1998) by comparing macroinvertebrate community structure in streams with
riparian vegetation in different stages of succession (Haefner and Wallace 1981; Murphy
et al. 1981).

Prince of Wales Island is the third largest island in the United States. It is located
within the Tongass National Forest in the southeast panhandle of Alaska. The primary
industries of the island are tourism, timber harvesting and sport and commercial ﬁsheries.
Because many streams and rivers on the island are located within Sitka spruce (Picea
sitchensis) and western hemlock (T suga heterophylla) forests, they are subjected to

forestry management practices such as clear-cut logging. Because timber harvesting is a

19

disturbance that could adversely affect the ﬁshing industry, the effects of clear-cutting on
spawning and rearing habitats of commercial and sport ﬁsh species such as coho
(Onocorhynchus kisutch), pink (0. gorbuscha), chum (0. keta), and sockeye (0. nerka)
salmon have been investigated (Murphy and Milner 1997; Duncan and Brusven 1985b).
However, because the topography of Prince of Wales and southeast Alaska is
mountainous, there are many small ﬁshless headwater streams in the upland forests
connecting to larger ﬁsh bearing streams that have received little attention. These small
headwater tributaries can be important sources of ﬂuvial organic matter to the larger
streams and its biota, and potential sources of benthic invertebrates (Wipﬂi and
Gregovich 2001) that are major food items for economically important ﬁsh species.
Therefore, knowledge of the effects of timber harvesting on benthic macroinvertebrates
in these headwater streams is important for the more effective management of both forest
and aquatic resources.

Prince of Wales has been logged extensively since 1950 (Swanston 1967). As a
result, there are streams within forest management conditions with differing types of
dominant vegetation in different stages of forest succession. This study contrasted
streams within four different forest management conditions. First, recent clear-cut (CC)
conditions that were in their ﬁfth year of regeneration had riparian vegetation comprised
primarily of shrubs. Second growth areas varied in canopy composition. Canopy
composition ranged between two extremes where either red alder (Alnus rubra) or
coniferous trees were dominant along the riparian margin. This study contrasted the

extremes, alder-dominated second grth (SA) and conifer-dominated second growth

20

(SC), in stands that had been regenerating between 35 and 40 years. Lastly, coniferous
trees dominated the old growth (OG) condition, which has previously not been cut.

In addition to studying the effects of timber harvest based on forest succession, I
also wanted to assess the effects of substrate type on macroinvertebrate response to
timber harvest. Gurtz and Wallace (1984) concluded that more stable substrates were
capable of mediating disturbance effects. Common substrates in these streams are large
woody debris, cobble and gravel.

My overall objectives were to: 1) contrast headwater benthic invertebrate
community structure between the four forest management conditions (CC, SA, SC, 0G);
and 2) evaluate which instream habitat (large woody debris, cobble or gravel) supported

greater taxa richness, diversity, densities, and biomass of invertebrates.

METHODS
LMLSIIES

The study was conducted on 12 streams in the Maybeso Experimental Forest and
the adjacent Harris River watershed in the Tongass National Forest, Prince of Wales
Island, in southeast Alaska (Figure l). Vegetation of the island is classiﬁed as a temperate
rainforest with annual precipitation ranging between 150 to 500 cm, and air temperatures
range from —20°C to 36°C (Duncan and Brusven 1985c; Harris et al. 1974). All stream
sites were ﬁrst order, high gradient, headwater streams of the Harris and Maybeso
catchments, and they were sampled upstream of logging roads once between 7-15 July

1998 and once between 11-14 June 1999.

21

Riparian vegetation differed across management conditions. Vegetation in clear-
cut condition was in its ﬁfth year of regeneration with salmonberry (Rubus spectabilis)
and Alaskan blueberry ( Vaccinium spp.) as the dominant riparian species. The alder-
dominated second growth condition, between 35 and 40 years old, consisted of dense red
alder along the riparian margin with a mixture of some conifers and an understory of
ferns and mosses. The conifer-dominated second growth condition, also between 35 and
40 years old, was predominantly Sitka spruce and western hemlock with some red alder
and a fern understory. The old growth condition, that had never been cut, had similar
vegetation to the conifer-dominated second growth, but the trees were more mature and
less dense. There also was a more extensive understory comprised of devils club

(OpIOpanax horridus), skunk cabbage (Lysichition americanum) and ferns.

ILEhYsicalMeasuLements

Physical measurements were taken in streams across the four management
conditions in 1999. Water temperatures were taken simultaneously with Onset® Optic
StowAway Temperature Loggers for a three week period. Discharge was measured using
a Marsh-McBimey® ﬂow meter by the velocity-area method described in Gore (1996).
Nitrates were measured with a Hach® nitrate ﬁeld test. The percentage of large woody
debris, cobble and gravel habitats was quantiﬁed within 25m in each of the 12 streams.
Algal AF DM on clay tiles was determined at two streams of each management condition

using the methods in Steinman and Lamberti (1996).

22

IIIE . 112' 151'

The experimental design of the study was a split-plot design; management
condition was the whole-plot factor, habitat was the sub-plot factor. Three streams were
sampled upstream of logging roads in each of the four management conditions. Within
each stream, three random macroinvertebrate samples were collected in 1998 while two
random samples were collected in 1999, from each of the following habitats: 1) large
woody debris, characterized by being of cedar origin, >10 cm diam. and conditioned (in
the water long enough to be suitable for invertebrate colonization); 2) completely
submersed rifﬂe cobble (64-256 mm diam.); and 3) gravel between 2-16 mm diameter. A
total of 108 samples were collected from the 12 headwater streams in 1998, and a total of
72 samples were collected from 12 headwater streams in 1999.

Results were tested for normality and log transformed or square root transformed
where necessary. Multiple ANOVAs were generated contrasting richness, diversity,
density and biomass, vs. management conditions (clear-cut, alder-dominated second
growth, conifer-dominated second growth, and old growth) and habitat (large woody
debris, cobble, gravel). ANOVAs were generated for the analysis of nitrates, discharge
and algal AF DM vs. management conditions (SAS Institute 1996). Although results were

transformed, they will be presented in untransformed fashion in graphs and tables.

III I l . l S l'
The large woody debris samples were randomly selected from the ﬁrst six pieces

encountered upstream of logging roads. Length and diameter measurements were taken to

estimate surface area. Each piece of large woody debris was collected and placed in a 19

23

L bucket and pressure sprayed with water from a hand-pumped lawn sprayer to remove
invertebrates. The macroinvertebrate sample was then rinsed through a 250-micron sieve
and transferred into a 250-ml Whirl-Pak®, preserved in 80% ethanol, and returned to the
lab for sorting under a dissecting scope. All invertebrates were picked from each sample,
counted, identiﬁed to the lowest possible taxon, mostly generic level (except
Chironomidae) using Merritt and Cummins (1996a). Chironomidae were subsampled and
identiﬁed to subfamily.

Cobble samples were removed from submersed cobble of rifﬂe areas. Each
cobble habitat was selected from the center of each of the ﬁrst three rifﬂe areas
encountered upstream of logging roads. Sample collection and processing were similar to
that of large woody debris. The cobble were labeled and returned to the lab to estimate
surface area. Surface area was determined by wrapping the cobble in foil paper, removing
the foil, and tracing it onto paper. Surface area was calculated from the paper using a Li-
Cor® portable leaf area meter Model-Li-3000.

Gravel core samples were collected from the center of each of the ﬁrst three
gravel areas encountered upstream of logging roads. Samples were collected with a core
sampler (6 cm x 6 cm x 6 cm). The core sampler was inserted into the gravel area to a
depth of 6 cm and the contents scooped out with a 250-micron mesh net. The entire
sample was transferred into a whirl-pak and returned to the lab for processing as above.

Invertebrate density was estimated from abundance and surface area calculations
for the large woody debris and cobble samples and was converted to number per lmz.
Density was calculated from abundance and sample volume for the gravel core samples

and was converted to number per 1m3. Dry mass of invertebrates was estimated

24

according to Benke et al. (1999). Richness was measured as total number of taxa present
and diversity was measured using the Shannon-Weiner diversity index (Hauer and Resh
1996). Macroinvertebrates were designated a ﬁrnctional feeding group status (shredders,
ﬁltering-collectors, gathering-collectors, scrapers and predators) according to Merritt and
Cummins (1996b). Dry mass of Oligochaeta were not determined and therefore omitted

from biomass and functional group analysis.

RESULTS
LlhxsisalMeasuremems

Daily temperature maximums were greatest in the old grth condition and
lowest in the conifer-dominated second growth. Daily temperature minimums were
lowest in the conifer dominated second growth condition and highest in the old grth
condition. Greatest differences in maximum and minimum daily temperatures were found
in the clear-cut condition and smallest differences in daily temperatures were found in
both conifer-dominated second growth and old growth conditions (Table 1).

Discharge was not signiﬁcantly different across management conditions, and
nitrate data were incomplete for comparisons across management conditions. Instream
habitat quantiﬁcation showed that cobble and gravel habitats comprised the greatest
proportion of available habitat for invertebrate colonization (Table 2). The conifer-
dominated second grth management condition had the largest proportion of large
woody debris with 20.9%, followed by old growth (12.1%), alder-dominated second

growth (5.4%) and clear-cut (3.3%) (Table 2).

25

Algal AFDM was greatest in the clear-cut (0.06 mg cm'z) management condition.
It was signiﬁcantly greater than in the old growth (0.013 mg cm'z) (p<0.05), alder-
dominated second grth (0.015 mg cm’z) (p<0.05), and conifer-dominated second

growth (0.013 mg cm'z) (p<0.05) (Figure 2).

U1-..'I.. 1' . rr‘ an |°‘ ' in!!! uqru‘rr‘r o ' or

A total of 38 genera were collected from representatives of the Ephemeroptera,
Plecoptera, Trichoptera and Diptera from the headwater streams of the Maybeso and
Harris River watersheds (Table 3). In addition, three subfamilies of Chironomidae were
identiﬁed from 1999 subsamples. Orthocladinae comprised the largest percentage across
the four management conditions, followed by Tanypodinae and Chironominae (Table 4).

Mean richness, or the total number of taxa collected, was lowest in the old growth
management condition. It was signiﬁcantly lower than in the clear-cut (p<0.05) and
alder-dominated second growth (p<0.05) conditions in 1998 (Figure 3a). Mean richness
in the old growth condition also was signiﬁcantly lower than in the alder-dominated
second growth (p<0.05) in 1999 (Figure 3b).

Mean Shannon-Weiner diversity was greatest in both alder and conifer-dominated
second growth management conditions and signiﬁcantly less in clear-cut and old grth

during both years (Figure 4).

C u... 0,1'l’01'q ‘ I‘r 'r'. qr! :9H1 5.1:”! uqlef'ﬂ'l 0H H
In 1998, mean densities of invertebrates (6590 m’z) collected from large woody

debris and cobble habitats in clear-cut management conditions were signiﬁcantly greater

26

than mean densities in old grth (1423 mi!) (p<0.05) and conifer-dominated second
grth (988 m’z) (p<0.05). In addition, the old growth management condition had
signiﬁcantly lower mean densities (p<0.05) than the alder-dominated second growth
(3131 m'z) (Figure 5a). There were no signiﬁcant differences in mean densities of
invertebrates collected from large woody debris and cobble habitats among management
conditions in 1999. Mean densities of invertebrates collected from gravel habitats were
greatest in the clear-cut (416,667 m'3) management condition and least in the old growth
(93,364 m'3); however, these differences were only signiﬁcant in 1999 (p<0.05) (Figure
5b).

In 1998, mean dry mass of invertebrates collected from large woody debris and
cobble habitats was greatest in the clear-cut (291mg m'z) management condition and
signiﬁcantly greater than in the old growth (95mg m'z) (p<0.05) condition. In addition,
mean dry mass in- alder-dominated second growth (225mg m'z) also was signiﬁcantly
greater than in old growth (p<0.05) management (Figure 6a). In 1999, mean dry mass of
invertebrates collected from the large woody debris and cobble habitats was greatest in
alder-dominated second growth (644mg m'z) conditions and signiﬁcantly greater than in
the old growth (195mg m’z) (p<0.05) (Figure 6b). There were no signiﬁcant differences
in mean dry mass of invertebrates collected from gravel habitats among management
conditions in 1998. However, in 1999, mean dry mass of invertebrates collected from
gravel habitats in clear-cut (20,491mg m'3) management conditions was signiﬁcantly
greater than mean dry mass in old grth (6,043mg m'3) (p<0.05) and conifer-dominated

second growth (p<0.05). In addition, the old growth management condition had

27

Signiﬁcantly lower mean dry mass (p<0.05) than the alder-dominated (15,673mg m'3)

second growth (Figure 6c).

V ..r or. "an 0.0'OOOIII anor'uerqe‘u'r on”

The composition of macroinvertebrates changed functionally and taxonomically
relative to the old grth condition in 1998 (Figure 7). The old growth management
condition was characterized by a high percentage of predators (42%), followed by
scrapers (30%), collector-gatherers (19%), shredders (9%) and absence of collector-
filterers. The clear-cut condition had collector-gatherers as the dominant functional
group. Baetis mayﬂies and Ostracoda comprised the largest proportion of biomass of this
functional group in the clear-cut condition, whereas the mayﬂy Paraleptophlebia was
dominant in the old growth. The dominant scraper also changed taxonomically, from
Drunella mayﬂies in the clear-cut to Cinygmula mayﬂies in the old growth. In addition,
the clear-cuts had a greater proportion of collector-ﬁlterers, represented by the caddisﬂy
Dolophiloides, than the old growth. The alder and conifer-dominated second growth
condition changed slightly with respect to old growth condition. Scrapers and predators
were still the dominant functional groups present, but both second growth conditions had
relatively fewer collector-gatherers and more collector-ﬁlterers than the old growth.
Taxonomically, the dominant scraper in the conifer-dominated second growth was the
mayﬂy Ironodes, while Cinygmula spp. was dominant in both old growth and alder-
dominated second growth conditions.

In 1999, the functional feeding group characterization differed from 1998 (Figure

8). Collector-gatherers (48%) made up the largest proportion in the old growth condition,

28

followed by predators (30%), scrapers (18%), shredders (4%) and the absence of
collector-ﬁlterers. Due to large biomass of the predator stoneﬂy Sweltsa, they were the
dominant functional group in both clear-cut and conifer-dominated second growth
managements. As in 1998, the alder-dominated second growth had predators and scrapers
as the dominant functional groups. Taxonomically, the alder-dominated second growth
had the mayﬂies Paraleptophlebia and Baetis as the dominant collector-gatherers, just as
in the old growth condition, but different from the clear-cut and conifer-dominated

second growth conditions where Baetis spp. and Chironomidae were dominant.

\IE or ‘1‘”: ' 3' Hi 1"... '5; “or" ran." no...

Mean richness was greatest in the large woody debris habitat and least in the
cobble habitat. Mean richness on gravel was signiﬁcantly greater than on cobble habitats
(p<0.05) only in 1998, while mean richness on large woody debris was signiﬁcantly
greater (p<0.05) than on gravel or cobble habitats in both years (Figure 9).

Mean Shannon-Weiner diversity was greatest in large woody debris habitats
followed by gravel and cobble habitats on both years (Figure 10). Mean diversity on
gravel habitats was signiﬁcantly greater than on cobble habitat in 1998 (p<0.05) and 1999
(p<0.05). Mean diversity of invertebrates on large woody debris was signiﬁcantly greater

(p<0.05) than on cobble habitat both years.

V U: .01. '1‘0 -. ' "I i 1!. 3011-. A “1' r'-. I. -.I_ ._
Mean invertebrate densities and biomass were not comparable between gravel and

the other two habitats because of the differences in units; however, mean densities were

29

signiﬁcantly greater on large woody debris habitat (1998 - 5294m’2; 1999 - 1893m’2) than
on cobble (1998 - 723m‘2; 1999 - 507m'2) during both years (Figure 11). Mean dry mass
of invertebrates ﬁ'om large woody debris (302mg m'z) was greater than on the cobble

(83mg m'z) habitat; however, the difference was only signiﬁcant in 1998 (Figure 12).

VI ..r ._., "cw .._. ’ nun .nuv II'Q.I..1'..1

In 1998, the functional feeding groups on large woody debris included scrapers
(38%) (primarily Drunella and Cinygmula mayﬂies), collector-gatherers (23%) (Baetis
and Paraleptophlebia mayﬂies and Chironomidae), and shredders (18%) (stoneﬂies
Despaxia and Zapada). Cobble habitats had a high proportion of scrapers (37%)
(Cinygmula and Ironodes mayﬂies), and the caddisﬂy, Arctopsyche spp., as the dominant
collector-ﬁlterer (33%). In the gravel habitat, there were large proportions of predators
(43%) (primarily Sweltsa spp. and the crane ﬂy Dicranota), as well as scrapers (35%)
(Cinygma spp. and Cinygmula spp.) (Figure 13).

In 1999, functional groups included for the most part, similar taxa from the
previous year. On large woody debris there was a large proportion of scrapers (55%)
followed by collector-gatherers (18%) and shredders (18%) (crane ﬂy, T ipula). Cobble
habitats had scrapers (60%) and collector-gatherers (25%) (mayﬂy Epeorus) in greatest
proportions, while the gravel habitat had a large proportion of predators (5 5%) and
collector-gatherers (28%) (Baetis spp., Paraleptophlebia spp. and Chironomidae) (Figure

14).

30

DISCUSSION
1 l l C 1. .

The results of this study showed that taxonomic and functional differences in
macroinvertebrate composition occurred between harvested sites and old growth
management conditions. The primary reasons for these changes may have been due to
changes in the availability of food resources (e.g. algal AF DM, labile allochthonous
inputs and ﬁne particulates) in the harvested conditions. Anderson (1992) found that
through the emergence of aquatic insects, streams in old growth management conditions
had a greater number of taxa and evenness than from recent clear-cut and second grth
deciduous riparian management conditions. In contrast, richness of macroinvertebrates in
our old growth conditions was lower than in clear-cut and alder-dominated second
growth conditions. Old grth conditions were lacking three taxa (Micrasema spp.,
Goeracea spp. and Prosimulium spp.) that were present in the other management
conditions. In addition, two dipterans (Thaumaleidae and Limonia spp.) were present
only in the clear-cut condition, and Neaphylax spp., Chyranda spp., Rithrogena spp. and
Ptilodactylidae were present only in alder-dominated second growth. Vegetation in the
old growth consisted primarily of more refractory coniferous allochthonous material,
while all other management conditions have sources of more labile deciduous
allochthonous inputs (i.e. red alder and salmonberry) that may be used more readily as a
food source by the caddisﬂy shredders Micrasema and Chyranda and the crane ﬂy
shredder, Limonia. In addition, mean algal AF DM was greatest in the clear-cut
management condition, which was similar to the ﬁndings of Murphy et al. (1986) who

reported their clear-cut streams averaged 130% greater periphyton AF DM than in

31

buffered and old growth streams, which they attributed to an increase in amount of light
reaching the stream. Increased algal AF DM in the clear-cut condition may have resulted
in a suitable food source for the caddisﬂy scraper, Goeracea, and the dipteran scraper,
Thaumaleidae. Lastly, the presence of the black ﬂy (Prosimulium) in managements other
than the old growth suggested that the harvest resulted in greater ﬁne particulates
entering the stream through bank erosion, or lateral inputs from runoff and resuspension
as Anderson and Sedell (1979) described. Webster et al. (1990) also showed that
disturbed streams exported signiﬁcantly more particulate organic matter than reference
watersheds. My study was conducted upstream of logging roads in order to avoid any
sedimentation effects caused by road construction, so ﬁne particulates to the stream
would have likely arisen from the harvest itself. However, our results also may have
differed from those of Anderson (1992) because of greater taxonomic resolution in his
study.

Although diversity of macroinvertebrates has been reported to be lower in
harvested streams (N ewbold et al. 1980), our results showed mean diversity was not
lower in clear-cut streams than in old growth management streams. In addition to lower
macroinvertebrate diversity, streams within harvested areas have generally been found to
have greater macroinvertebrate densities throughout the lower United States (Hawkins et
al. 1982; Silsbee and Larson 1983), in the Paciﬁc Northwest (Murphy et al. 1981), and
Alaska (Duncan and Brusven 1985b). However, Hartman et al. (1996) described negative
impacts of harvesting on macroinvertebrate densities in Carnation Creek, a locality near
Prince of Wales. In this study, large numbers of Chironomidae in the old growth

condition and large numbers of midges and Baetis spp. in the clear-cut condition, as well

32

as fewer taxa, were primarily responsible for lower diversity. The increase in densities
after a harvest was largely due to large numbers of Baetis spp., Chironomidae and the
mayﬂy Drunella in clear-cut conditions. Similarly, Wallace and Gurtz (1986) reported
increased numbers of Baetis spp. in their harvested streams and attributed it to increases
in autochthonous production. The increase in macroinvertebrate densities in our alder-
dominated second growth conditions were due to large numbers of Zapada spp. and
Micrasema spp., both of which are shredders and presumably using allochthonous inputs
from the red alder riparian vegetation. Culp and Davies (1985) also found higher
macroinvertebrate abundances in substrate patches with alder detritus as compared to
hemlock detritus. I found that headwater streams of the Maybeso Experimental Forest
and adjacent Harris River watershed had greater invertebrate densities following a
harvest, with the clear-cut managements having the greatest densities. Second growth
conditions had intermediate densities, and old grth had the lowest densities of
invertebrates.

Mean biomass also was greater in harvested areas, particularly in the alder-
dominated second growth and recently clear-cut management conditions. This greater
invertebrate biomass in harvested areas may be the result of a greater amount of nutrient
availability through increased litter processing rates (Stone and Wallace 1998),
particularly alder litter which is processed quicker than conifer needles (Sedell et al.
1975). Stone and Wallace (1998) suggested that these processes could lead to increased
production of macroinvertebrates in harvested streams.

Changes in the functional feeding group composition of streams have been shown

to be the result of changes or differences in food availability (Vannote et al. 1980). It is

33

probable that differences in macroinvertebrates between years may have resulted from
different seasonal sampling times. Overall, larger proportions of scrapers were present in
July than in June. Similarly, Duncan and Brusven (1985b) also found a tendency for
increased scrapers from spring to summer in their studies in Prince of Wales Island.
Large proportions of predators across management conditions may have resulted from an
under representation in the non-predator fauna due to low sampling efﬁciency, since all
available habitats (ﬁne woody debris, root wads, mosses and bedrock) were not sampled
in this study. Another possible reason for the presence of a large proportion of predators
is a prey base with a rapid turnover in generation time. For example, large numbers of
Harpactacoida copepods, with a presumably short generation times, were collected from
all management conditions, primarily from large woody debris and gravel habitats where
the predator Sweltsa were more commonly found.

Streams with alder-dominated second growth riparian vegetation had a functional
and taxonomic similarity to old growth management conditions. However, alder-
dominated second grth also had an abundant, richer and more diverse fauna.
Therefore, streams in alder-dominated second growth conditions were potentially
contributing a greater amount and variety of benthic invertebrates to larger ﬁsh-bearing

streams.

IIJnstremHalzitats
In the event of a disturbance such as timber harvest, Gurtz and Wallace (1984)
concluded that invertebrates responded by increasing their abundance on physically

larger substrates that required more energy to move. In my study, the physical stability,

34

structural complexity, and ability to retain organic matter resources for invertebrate
consumption may have all been important. Invertebrate richness and diversity were
greatest in the large woody debris followed by gravel and cobble substrates. This could
have been due to a combination of physical stability and structural complexity of the
substrates. Large woody debris is physically larger than either cobble or gravel substrates,
and has greater structural complexity than cobble substrates, possibly allowing for a more
stable habitat, greater variety of food resources and refugia for invertebrates. Wallace et
al. (1995) showed an increase of coarse and ﬁne particulate matter accumulation after log
additions to a stream in North Carolina. In addition, large numbers of taxa have been
found in association with wood as shown by Dudley and Anderson (1982) who recorded
56 taxa closely associated with woody debris and another 129 species as facultatively
associated.

Gravel substrates also were areas of high numbers and biomass of invertebrates in
my study streams. Gravel is more structurally complex than cobble. Cobble is generally
smooth surfaced while the gravel habitat has interstitial spaces for refuge and detritus
accumulation. Culp and Davies (1985) showed the importance of interstitial detritus in
gravel substrates in determining the distribution of macroinvertebrates, with greater
numbers of invertebrates associated with either high or low levels of red alder detritus
than hemlock.

All functional groups were represented in each of the three habitats under study.
Scrapers made up a large proportion in each of the habitats; however, each habitat had a
dominant functional group associated with it. Large woody debris had the highest relative

proportion of shredders, reﬂecting on its ability to retain coarse particulates (i.e. leaf litter

35

and detritus) (Bilby and Likens 1980). Cobble substrates had the highest relative
proportion of collector-ﬁlterers suggesting their importance to this group even in the
presence of stable large woody debris substrates. Gravel habitats had the highest
proportion of predators, likely due to the presence of high abundances of Harpactacoida
copepods and chironomid as a prey base.

Instream habitat quantiﬁcation revealed that the sections of streams examined
were comprised primarily of cobble and gravel substrates and the proportion of large
woody debris present and available for macroinvertebrate colonization in old growth
management conditions averaged 12.1% of the total habitat available. The proportion of
large woody debris in clear-cut management conditions also was small (3.3%). Very few
large pieces of wood remained in these streams after the harvesting event and the
majority of the wood that was available was small and appeared to be slash from the
harvesting event itself. Alder-dominated second growth conditions also had a small
proportion of large woody debris (5.4%). Bilby and Ward (1991) compared woody debris
inputs to streams from old growth, clear-cut and second growth forests and concluded
that large woody debris inputs from second growth forests with red alder riparian stands
were minimal. Surprisingly, conifer-dominated second growth conditions had the greatest
proportion of large woody debris (20.9%); however, most of the large woody debris was
largerin diameter than that of the current forest, suggesting that the origin of the wood
was not from the second growth stand and was in the streams before the harvest or as a
direct result of the harvest.

In addition to the functional importance of large woody debris in: 1) channel and

pool formation (Keller and Swanson 1979); 2) the retention of organic matter (Bilby

36

1981); and 3) serving as a habitat for invertebrate colonization (Gurtz and Wallace 1984),
it is important biologically as a food resource for invertebrates (Anderson et a1. 1979;
Kaufman and King 1987). Large woody debris contributed signiﬁcantly to the richness,

diversity, and abundance of macroinvertebrates in Southeast Alaskan headwater streams.

CONCLUSION

The results of this study suggested that forest succession after a timber harvest
affected macroinvertebrate community structure in the upland forests of Southeast
Alaska, as a result of changes in food availability relative to the old growth condition.
First, canopy removal has led to increases in sunlight penetration to the streambed and
consequently to higher autochthonous food resources which results in greater biomass
and densities of scraper and collector-gatherer invertebrates in clear-cut management
conditions. Secondly, in the alder-dominated second growth condition, provision of more
labile allochthonous organic matter (i.e. red alder) has increased the abundance and
number of different shredder invertebrates. Lastly, timber harvest has led to the presence
of collector-ﬁlterer organisms in all harvested conditions. Although clear-cut conditions
had the highest densities of invertebrates, alder-dominated second grth conditions had
high densities of invertebrates in addition to a richer and more diverse fauna. Thus,
management of upland forests in Southeast Alaska should provision for red alder riparian
vegetation to maximize macroinvertebrate diversity and abundance.

The evaluation of instream habitats showed that large woody debris and gravel
substrates were important to the contribution of high densities and biomass of

invertebrates to upland Southeast Alaskan headwater streams. In addition, large woody

37

debris also contributed to high taxa richness and diversity. Maintenance of both gravel
and large woody debris substrates within these streams is advantageous for large numbers
of benthic invertebrates that could potentially beneﬁt the diet of downstream
economically important ﬁsh communities.

There is a need for an alternative method to clear-cut harvesting of upland forests
of Prince of Wales. The alternative must result in an abundant, richer and more diverse
community of benthic invertebrates. Results of this study suggest that this could be
achieved by Opening up a portion of the canopy, having red alder along the riparian
margin, and maintaining sources of wood to the streams. I would suggest selectively
cutting a proportion of the riparian vegetation to potentially allow for: 1) greater sunlight
penetration to stimulate autochthonous production; while 2) maintaining wood
recruitment to the stream; and 3) planting of red alder saplings along the riparian margin
for labile sources of allochthonous organic matter. This management strategy should
positively inﬂuence invertebrate richness, diversity and abundance, without the loss of

instream habitat.

38

 

Table 1. Average daily maximum, minimum and differences in streamwater

temperature (°C) 1999.

 

 

 

Old Growth Clear-cut Alder SG Conifer SG
Ave daily max 10.44 9.96 9.13 7.71
Ave daily min 10.03 9.03 8.38 7.35
Ave max minus min 0.41 0.93 0.75 0.36

 

Table 2. Percentage of instream large woody debris, cobble and gravel habitats within
management conditions. Do not equal 100 percent in cases where ﬁne woody debris or

bedrock substrates were present.

 

 

Management

condition large woody debris cobble gravel
Old Growth 12.1 46.3 40
Clear-cut 3.3 53.8 32.1
Alder SG 5.4 22.1 67.5
Conifer SG 20.9 24.1 34.2

 

39

Table 3. Checklist of taxa collected from upland forest headwater streams

of Maybeso Experimental Forest and adjacent Harris River watershed

Prince of Wales, Southeast Alaska.

 

Ephemeroptera
Baetidae
Baetis
Heptageniidae
C inygma
C ynigmula
Epeorus
Ironodes
Rithrogena
Ephemerellidae
Drunella
Leptophlebiidae

Paraleptophlebia

Plecoptera
Nemouridae
Zapada

Visoka

Leuctridae
Despaxia
Chloroperlidae

Sweltsa

Trichoptera
Philopotamidae
Dolophiloides
Hydropsychidae

Arctopsyche

40

Ryacophilidae
Rhyacophila
Brachycentridae
Micrasema
Limnephilidae
Cryptochia
Chiranda
Moselyana
Pseudostenophylax
Psychoglypha
Goeridae
Goeracea
Uenoidae
Neophylax
Glossossomatidae

Anagapetus

Table 4 (cont)

 

Coleoptera

Ptilodactylidae

Diptera
Thaumaleidae
Ceratopogonidae

Atrichopogon
Probezzia
Chironomidae
Chironominae
Orthocladinae
Tanypodinae
Dixidae

Dixa

Psychodidae
Pericoma
Simuliidae
Prosimulium
Tipulidae
Dicranota
Limonia
Hexatoma
Pedicia
T ipula
Empididae
Chelifera
Clinocera

Oreogeton

Non-insects
Turbellaria
Annelida

Oligochaeta
Copepoda

Harpactacoida
Ostracoda

Hydracarina

 

41

Table 4. Percent distribution of Chironomidae subfamilies collected from large woody

debris, cobble and gravel habitats in each management condition.

 

 

 

 

Habitat Taxa Old Growth Clear-cut Alder SG Conifer SG
Large Woody Orthocladinae 59.3 91 85.6 83.4
Debris Tanypodinae 33 .3 6. 1 13.1 10.9
Chironominae 7.5 2.9 l .3 5.7
Cobble Orthocladinae 100 100 100 94
Tanypodinae 0 O 0 2.7
Chironominae 0 0 O 3.4
Gravel Orthocladinae 100 86.2 80.2 95.9
Tanypodinae 0 2.8 4 0
Chironominae 0 1 1 . 1 15.8 4. l

 

42

Alaska

 

IS

 

t" ‘9‘:
Maybeso
Experimental
Forest,
Prince of Wales
Island

 

 

 

 

Figure 1. Maybeso Experimental Forest, Prince of Wales Island, Southeast Alaska

43

Mean algal AFDM (mg / cm2 i SE)

0.08
0.07
0.06
0.05
0.04
0.03
0.02
0.01

 

Old Growth Clear-cut Alder SG Conifer SG

Figure 2. Mean algal AF DM among management conditions 1999. Means with

different letters are signiﬁcantly different (p<0.05).

44

Mean richness (number of taxa 3: SE)

 

Old Growth Clear-cut Alder SG Conifer SG

Figure 3. Mean richness of invertebrates among management conditions from (a)

1998 and (b) 1999. Means with different letters are signiﬁcantly different (p<0.05)

45

Mean Shannon-Weiner diversity 5: SE

 

Old Growth Clear-cut Alder SG Conifer SG

Figure 4. Mean Shannon-Weiner diversity of invertebrates among management
conditions from (a) 1998 and (b) 1999. Means with different letters are

signiﬁcantly different (p<0.05).

46

9000
8000
7000
6000
5000
4000
3000

Mean density (# / m2 :1: SE)

2000
1 000

600000

500000

400000

300000

200000

Mean density (# / m3 i SE)

1 00000

 

Old Growth Clear-cut Alder SG Conifer SG

Figure 5. Mean densities of invertebrates among management conditions from (a)
large woody debris and cobble habitats 7-15 July 1998; and (b) gravel habitats 11-

14 June 1999. Means with different letters are signiﬁcantly different (p <0.05).

47

400

300
200
E5
V)
,F 100
a
Di
3; 0
g 1000
e
j; 800
S
O
:2 600
400
200
0
,4 30000
[-1.1
V)
;H 25000
E
1; 20000
3%
a 15000
N
E
43 10000
5 5000
2

 

Old Growth Clear-cut Alder SG Conifer SG

Figure 6. Mean dry mass of invertebrates among management conditions from (a)
1998 large woody debris and cobble habitats; (b) 1999 large woody debris and
cobble habitats; and (c) 1999 gravel habitats. Means with different letters are

signiﬁcantly different (p<0.05).

48

Old Growth Clear-cut

   

Alder Second Conifer Second
Growth Growth

   

EDEE-

Shredders Gatherers Filterers Scrapers Predators

Figure 7. Percentage of functional feeding groups, based on biomass, present

among management conditions 7-15 July 1998.

49

Old Growth Clear-cut

   

Alder Second Conifer Second
Growth Growth

   

ED

Shredders Gatherers Filterers Scrapers Predators

 

Figure 8. Percentage of functional feeding groups, based on biomass, present

among management conditions 11-14 June 1999.

50

Mean richness (number of taxa :1: SE)

 

Large woody debris Cobble Gravel

Figure 9. Mean richness of invertebrates among habitats from (a) 7-15 July 1998
and (b) 11-14 June 1999. Means with different letters are signiﬁcantly different

(p<0.05).

51

Mean Shannon-Weiner diversity :t SE

 

Large woody debr's Cobble Gravel

Figure 10. Mean Shannon-Weiner diversity among habitats from (a) 7-15 July
1998 and (b) 11-14 June 1999. Means with different letters are signiﬁcantly

different (p<0.05).

52

7000

6000

5000

4000

3000

2000

Mean Density (# / m2 i SE)

1000

1998

 

 

 

 

Large woody debris

Cobble

 

l 999

Figure 11. Mean densities of benthic invertebrates on large woody debris and cobble

habitats across all management conditions. 7-15 July 1998 and 11-14 June 1999.

53

400

350

300

250

200

150

100

Mean dry mass (mg / m2 3: SE)

50

 

Large woody debris Cobble

Figure 12. Mean dry mass of invertebrates among large woody debris and cobble
habitats. 7-15 July 1998. Means with different letters are signiﬁcantly different

(p<0.05)

54

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56

LITERATURE CITED

57

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Duncan, W.F.A. and Brusven, M.A. 1985a. Energy dynamics of three low-order
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61

APPENDIX

62

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.: 2001-05
Title of thesis or dissertation (or other research projects):
BENTHIC INVERTEBRATE COMMUNITY STRUCTURE AS AFFECTED BY FOREST

SUCCESSION AFTER CLEAR-CUT LOGGING ON PRINCE OF WALES ISLAND, SOUTHEAST
ALASKA

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

Entomology Museum, Michigan State University (MSU)

Other Museums:

Investigator’s Name(s) (typed)
OSV HERNANDEZ

 

 

Date 20. IV 2001

*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.

63

Appendix 1.1

Voucher Specimen Data

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Voucher Specimen Data

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Appendix 1.1

Voucher Specimen Data

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Page 3 of 5

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66

Appendix 1.1

Voucher Specimen Data

ges

__ _Pa

Page 4 of 5

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67

Appendix 1.1

Voucher Specimen Data

5 of __5__ Pages

 

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68

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