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BENTHIC INVERTEBRATE COMMUNITY MEASURES
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RIVER DELTA, SOUTHCENTRAL ALASKA.

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BENTHIC INVERTEBRATE COMMUNITY MEASURES AMONG STREAM
CHANNEL TYPES OF THE COPPER RIVER DELTA, SOUTHCENTRAL
ALASKA.

By

Todd C. White

A THESIS

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

MASTER OF SCIENCE
Entomology

2009

ABSTRACT
BENTHIC INVERTEBRATE COMMUNITY MEASURES AMONG STREAM
CHANNEL TYPES OF THE COPPER RIVER DELTA, SOUTHCENTRAL
ALASKA.
By
Todd C. White
The Copper River Delta of Southcentral Alaska is the largest contiguous

wetland on the North American Pacific coast, and supports economically important
commercial and recreational fisheries for all five species of pacific salmon. Some
biological factors influencing salmon populations in the Copper River Delta have
been previously investigated, but little effort has been made to establish baseline
information on freshwater aquatic communities in the region. In an effort to provide
area managers with aquatic community measures for future comparisons, benthic
invertebrate community structure was contrasted among twelve streams representing
six stream channel types common to the area and important to salmonid
development. In general, invertebrate density, taxa richness, and diversity were
greatest in channel types designated as high potential for salmonids. Taxonomic
and functional feeding group measures show that macroinvertebrate communities of
Copper River Delta streams sampled are representative of early (<50 years) stages
of colonization after the major earthquake disturbance of 1964. Continued long

term monitoring of invertebrate populations is required to track changes in food

resources important to economically important fish species.

ACKNOWLEDGMENTS

I would like to thank Michigan State University’s Department of
Entomology for the opportunity to further my education, and my committee
members Dr. M., Kaufmann, Dr. D. Hayes, Dr. K. Cummins, and Dr. R.W. Merritt
who provided excellent guidance. I also thank Dr. M. Eric Benbow and Dr. Molly
McIntosh for their technical expertise. In addition, I’d like to thank Dr. M. Berg
and Dr. G. Lamberti for comments received on field sampling protocols. I would
also like to thank a number of contributors who funded my tuition and living
expenses, in part, while at Michigan State University: College of Agriculture and
Natural Resources, Department of Entomology, and the Department of Integrative
Studies. This research would not have been possible without the funding and
logistical support provided by the Pacific Northwest Research Station, USDA Forest
Service, Corvallis, OR, USA, and the Cordova Ranger District, Chugach National
Forest, Cordova, AK, USA. I would specifically like to thank Gordon Reeves,
Team Leader, Deyna Kuntzsch, Biologist, Dirk Lang, Biologist, Ken Hodges,
Biologist, Tim Joyce, Biologist, Samantha Greenwood, GIS/Ecology Specialist,
Technicians: Brian Nielsen, Sean Meade, Kirsti Jurica, Jonathon Kersh, and Rosa
McIntyre, and everyone else at the office in Cordova for making my time in Alaska
productive, life-changing, and bear attack free. Lastly, I would like to thank my
parents Duncan & Kathy, and my siblings Joshua, Nathan, and Amanda for their

continued support.

iii

TABLE OF CONTENTS

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

BENTHIC INVERTEBRATE COMMUNITY MEASURES AMONG STREAM
CHANNEL TYPES OF THE COPPER RIVER DELTA, SOUTHCENTRAL

ALASKA ....................................................................................... 1
Introduction .................................................................................... I
History and Management of the Copper River Delta ............................ 1
Summary of Copper River Delta Research Studies .............................. 2
Macroinvertebrate Community ..................................................... 5
Materials and Methods ........................................................................ 7
Study Area ............................................................................. 7
Sampling Sites by Channel Type Description .................................... 7
Benthic Macroinvertebrates ........................................................ 11
Analysis ............................................................................... 13
Results ......................................................................................... l4
Macroinvertebrate Richness & Diversity Among Stream Channel Types... l4
Macroinvertebrate Densities Among Stream Channel Types .................. 16
Functional Feeding Group Proportions Among Stream Channel Types. 17
Discussion .................................................................................... 20
Macroinvertebrate communities of the Copper River Delta. . . . . . . . 20
The Copper River Delta and other regions of Alaska .......................... 22
Conclusion .................................................................................... 25
Appendix 1 .................................................................................... 40
Literature Cited ............................................................................... 45

iv

LIST OF TABLES

CHAPTER 1

Table 1. Physical stream characteristics of study sites within the Copper River
Delta, Alaska, (modified from Paustian 1991) ........................................... 26

Table 2. Macroinvertebrate taxa collected from 12 streams during June-August
2005 and 2006 within the Copper River Delta, Southcentral Alaska.
* Chironomidae identified to subfamily ................................................... 27

Table 3. Selected macroinvertebrate indices for stream channel types within the
Copper River Delta, Southcentral Alaska ................................................. 32

Table 4. Mean relative abundance of subfamilies of the Chironomidae among
stream channel types of the Copper River Delta, Southcentral Alaska, 2005 &
2006 ............................................................................................ 33

Table 5. Dominant invertebrate taxa by functional feeding group among Copper
River Delta streams across channel type and year (2005-2006) ........................ 34

LIST OF FIGURES

CHAPTER I

Figure 1. Selected sampling sites, Copper River Delta, Chugach National Forest,
Southcentral Alaska .......................................................................... 35

Figure 2. Mean richness of invertebrates among streams across channel type over
entire study period (2005 and 2006). Means with different letters are significantly
different (Tukey HSD, p<0.05) ............................................................ 36

Figure 3. Mean Shannon-Weiner diversity of invertebrates among streams across
channel type over entire study period (2005 and 2006). Means with different letters
are significantly different (Tukey HSD, p<0.05) ......................................... 37

Figure 4. Mean densities of invertebrates among sampling sites across channel type
over entire study period (2005 and 2006). Means with different letters are
significantly different (Tukey HSD, p<0.05) ............................................. 38

Figure 5. Percentage of functional feeding groups, based on abundance, present
among sampling sites across channel types over entire study period (2005 and
2006) ........................................................................................... 39

vi

CHAPTER 1
BENTHIC INVERTEBRATE COMMUNITY MEASURES AMONG STREAM

CHANNEL TYPES OF THE COPPER RIVER DELTA, SOUTHCENTRAL
ALASKA

INTRODUCTION
History and Management of the Copper River Delta

The Copper River Delta, Alaska is the largest contiguous coastal wetland on
the Pacific coast of North America (Thilenius 1990). Fed by one of the world’s
largest river systems, the Delta extends 75 miles along the Gulf of Alaska southeast of
Anchorage, and encompasses 700,000 acres of constantly changing river channels,
marshland, tidal flats, and sloughs (Christensen 2000). The Delta is located in one of
the moSt seismically active regions of the world (Thomas et al 1991), experiencing
major earthquakes every 600-1000 years; the most recent event occurring in 1964.
The 1964 earthquake measured 8.6 on the Richter scale, and lifted the Delta an
average of 6.7 feet above previous levels (Hansen and Eckel 1971). The associated
upheaval resulted in the establishment of 1.5 kilometers of new marshland on
previously un-vegetated tidal flats. The rapid transformation of the formerly open
marshland zone into shrubland was dominated initially by willow (Salix spp.), and
more recently alder (Alnus spp.) (Thilenius 1995). The dynamic tectonic environment
of the Delta results in alternate expansion of dry-land vegetation types which are
eventually drowned and buried by sediment transport from upriver. Thus, Delta plant
and animal communities constantly undergo regular and perpetual cycles of renewal

and succession (Christensen 2000).

The Copper River Delta is located entirely within the Chugach National Forest
under the management of the Forest Service, US. Department of Agriculture (USDA).
The Delta is unique among USDA Forest Service systems in that it is the only such
system mandated (Alaska National Interest Land Conservation Act 1980) with a
priority to manage and conduct research to support the conservation of fish, wildlife,
and their associated habitat. The Delta currently supports the most productive
commercial fishery in Southcentral Alaska with annual spawning migrations of five
species of Pacific salmon, thus sustaining a lucrative commercial fishing industry,
human subsistence use, and a diverse native wildlife fauna (Christensen 2000).
Current USDA Forest Service management assessments (Kruger 1995) in the Delta
region have identified a critical need for baseline data on ecosystem processes and
conditions in order to monitor and document physical and biological changes over
time. In addition, understanding the natural processes that influence the Delta’s
salmon populations has been designated as a research area of high priority because of
the ecological and economic importance of the region’s fisheries (Kruger 1995).
Summary of Copper River Delta Research Studies

To date, only preliminary studies concerning the importance of freshwater
habitats have been conducted within the Copper River Delta. The majority of the
research questions posed have concentrated on the effects of tectonic uplift on plant
communities (Thilenius 1995), the role of marine—derived nutrients in salmon ecology
(Hicks et al 2005; Lang et al 2005), and the genetic relationships of some salmonid
populations (Saiget et al 2007; Williams et al 2007). Only two studies concerning

freshwater aquatic macroinvertebrates have been published since the earthquake of

1964, and none prior to, illustrating the need for additional research and baseline data
in this area.

Thilenius (1995) investigated the effects of the 1964 uplift event on plant
communities within the COpper River Delta. Pre-uplift aerial photographs were
compared to vegetation sampling and aerial photography conducted in 1974 and 1979.
Results showed that vegetation within uplifted areas of the Delta were undergoing a
rapid successional sequence from grass/sedge-dominated communities to communities
dominated by woody shrubs (alder/willow). The sequence of succession Observed
during the study period was typical to the Delta region above the uplift affected area.

Hicks et al (2005) studied the occurrence of marine-derived nutrients in Delta
freshwater-riparian food webs. Seasonal sampling of stable isotopes showed that
juvenile coho salmon, threespine sticklebacks, and aquatic macroinvertebrates in Delta
beaver ponds were enriched with marine nitrogen and carbon, and that artificial
enrichment with salmon carcasses increased the marine-derived N and C values of
juvenile coho salmon. Aquatic vascular plants were found to be enriched with marine
N only, and riparian vegetation showed no marine-derived enrichment.

Lang et al (2005) investigated the influence of fall-spawning adult coho
salmon on the growth and production of juvenile coho salmon in Delta beaver ponds.
They compared beaver ponds with natural spawning, ponds without spawning, and
ponds without spawning but artificially enriched with salmon eggs and carcasses.
Results were variable in the study, with increased growth in some pond-spawning
adults and improved conditions of juvenile salmon. Enrichment of ponds without

natural Spawning significantly increased the growth and condition of juvenile coho

salmon, but the results provided little evidence that the Observed short-term growth
benefits led to greater overwinter survival/outmigration.

Spawning and movement of coastal cutthroat trout on the Delta was
investigated by Saiget et al (2007). Movements of coastal cutthroat trout were
monitored for two years using radio telemetry and tag-recapture. Similar sized,
morphologically indistinct individuals displayed anadromous and/or potamodromous
migrations seemingly at random. Timing of spawning stream entry and post-spawning
movements were highly variable among individual trout and spawning was found to
be concentrated within the upper reaches of streams.

Natural hybridization of rainbow and coastal cutthroat trout was investigated
by Williams et al (2007). Mitochondrial DNA molecular genetic methods were used
to identify the presence of rainbow/cutthroat (cuttbow) hybrids at eleven sites on the
Delta. Hybridization of cutthroat and rainbow trout populations across study sites
varied from 0% to 58% of fish sampled. NO Significant correlation was found between
stream channel process groups and number of hybrid fish sampled. Backcrossed
hybrid individuals were found indicating that at least some cuttbow hybrids on the
Delta were reproductively viable.

The intertidal benthic resources of silt-clay substrates at outflows of two Delta
rivers were studied by Powers et a1. (2002). The benthic community was
characterized by low species diversity, and was dominated by tellinid bivalves,
polychaete worms, and corophid arnphipods. Temporal and spatial changes in benthic
community abundances and densities were found to correspond to differences in tidal

inundation and sediment temperature.

Powers et al. (2006) also studied the distribution of the invasive bivalve Mya

arenaria on intertidal flats of the Copper River Delta. Abundance of M. arenaria was

found to be greatest (4000/m2) in areas of higher salinity and water clarity. Density

and growth of M. arenaria in tidal flats of the Delta were observed to be similar to
values reported for the White Sea (Russia), an area located at a similar latitude.

Mason (1991) conducted a 2 week interdisciplinary survey of aquatic habitats
on the Copper River Delta including geomorphology, limnology, and wetland plant
ecology. In this study, samples were qualitative and concentrated among wetlands
that experience periodic flooding. Mason (1991) found that abundance and diversity
of phytoplankton, macrophytes, macroinvertebrates, and fish in interbasin habitats
(beaver ponds) was higher than in main channel (glacial) habitats . Marsh (tidal)
habitats were characterized by dominance of euryhaline crustacea while pond - (non-
tidal) habitats were dominated by Cladocera and Chironomidae. Wetland ponds also
were shown to have the highest diversity of macroinvertebrates of all habitats
sampled.
Macroinvertebrate community

Macroinvertebrate communities play an important role within food webs of
many aquatic ecosystems by providing a link between organic matter inputs, primary
producers, and higher trophic levels (i.e. fish) (Allan and Castillo 2007). Aquatic
macroinvertebrates influence nutrient cycles, decomposition rates, and translocation of
materials within stream habitats (Hauer and Resh 2006), and also serve as indicators
of stream integrity and water quality (Wallace & Webster 1996). The feeding habits

of benthic macroinvertebrates convert both autochthonous (in-stream) primary

production by macrophytes and periphyton, and allochthonous (terrestrial) inputs of
organic matter into insect biomass directly available to fish in aquatic habitats
(Cummins and Klug 1979). In the Pacific Northwest, macroinvertebrates serve not
only as a direct form of nutrition for many fish species, but also facilitate indirect
transport of marine derived nutrients to higher trOphic levels of coastal food webs
(Hicks et al. 2005). Inputs of macroinvertebrates into the rearing habitats Of juvenile
fish species can be of particular importance, especially in the case of juvenile
salmonid fishes that undergo a migration event before attaining adulthood. Fisheries
research has shown that juvenile salmonids that attain the greatest size prior to
migration have the highest probability of returning as reproductive adults (Smith and
Griffin 1994; Bilton et al. 1982), making the quantity and quality of prey items
consumed during rearing Of considerable importance.

The objectives of this study were to characterize and contrast benthic
macroinvertebrate communities across stream channel types common to Alaska
temperate streams of the Copper River Delta. Baseline data generated from this study
will be used by fisheries managers of the USDA Forest Service to make future
management decisions concerning commercial and recreational fishing as well as land

use on the Copper River Delta.

MATERIALS AND METHODS
Study Area

The study was conducted within 12 streams of the Copper River Delta,
Southcentral Alaska (Figure 1). Weather patterns of the Delta are similar to those seen
in Southeast Alaska, with approximately 380 inches of rain per year with greatest
discharges most common in the fall months. Low water periods typically occur in late
spring and mid- summer (Meyer et al. 2001). The Delta is separated into east and
west ranges by the mainstream and braided channels of the Copper River (Figure l).
The area is hydrologically complex, and characterized by networks of beaver ponds
interconnected by stream channels (Hicks et a1 2005). Twelve study sites representing
six US Forest Service designated channel types were selected using aerial
photographs, US Forest Service GIS data, channel morphology, presence/absence of
salmonid rearing habitat, and access. Physical characteristics of each channel type are
summarized in Table l. Benthic macroinvertebrate sampling was conducted on a
monthly basis during the period of June—August, 2005 and 2006.

Sampling Sites by Channel Type Description: (from Paustian et al. 1992)
Estuarine Channel Type: Silt Substrate Estuarine Channel or Slough (ESI)

Silt Substrate Estuarine streams are associated with shallow embayments along
coastal forelands and large glacial river deltas. The ES] channels are characterized by
stream gradient 0-0.5%, incision depth 0-4 m (13 ft), bankfull width < 20m (66 ft),
and dominant substrate of silt/clay and sand. Riparian vegetation is dominated by
non-forested grass and sedge communities with some alder/willow. Silt Substrate

Estuarine streams are depositional channels with low energy due to nearly flat

gradients. Water flow and depth is strongly influenced by tidal stage. Suspended
glacial silt load is generally high. Little, if any, spawning occurs in ES] streams due
to fine sediments dominating the substrates. Available rearing area is high with pools
showing good depth for overwintering, but habitat is generally underutilized due to
suspended sediment load. One study area was classified as ESl: Alaganik River in the
Alaganik river system.

Estuarine Channel Type: Large Estuarine flannel (E84)

Large Estuarine Channels on the Delta are associated with moderate to large
drainage basins of inland bays and inlets. The ES4 channels are characterized by
stream gradient 5 2%, incision depth of < 5 m (16.5 ft), bankfull width > 10 m (33 ft),
substrate dominated by gravel and cobble. Riparian vegetation is dominated by grass
and sedge communities. Large estuarine streams are depositional channels subject to
tidal influences. Low stream energy, gravel and sand bars, and large woody debris
typify ES4 channel types on the Delta. High quality gravel substrate is frequented by
pink and churn salmon in high densities during spawning. Rearing habitat is minimal
with salmon fry only temporarily remaining in the system before moving seaward.
One study site was classified as ES4 channel type: Hartney Creek in the Hartney
Range system.

Floodplain Channel Type: Narrow Low Gradient Flood Plain Channel (FP)

Floodplain streams on the Delta are characterized by stream gradient 5 2%,
incision depth 5 2m (6.5 ft), bankfull width < 10m (33 ft), and dominant substrate of
sand to small rubble. Riparian vegetation is dominated by Sitka spruce, western

hemlock, and alder communities. The FP3 streams function as temporary sediment

deposition systems dominated by deposits of sand and fine gravel with frequent large
woody debris accumulations. Available spawning habitat for coho salmon is high. If
located next to accessible lakes, F P3 channels provide excellent spawning habitat for
sockeye salmon. Good average depth, woody debris occurrence, and beaver dams
provide good overwintering and rearing habitat for juvenile salmonids. Two study
sites were classified as F P3 streams: the Little Martin River in the Martin River
system of the East Delta, and BlackHole Creek of the Alaganik system on the West
Delta.

Glacial Ouput Channel Type: Moderate Width Glacial Ch_annel (GO)

Moderate width glacial output channel types occur in the mid to upper valley
position in glacial watersheds. The G04 streams are characterized by stream gradient
of 2-6%, incision depth 5 4m (13 ft), variable bankfull width, mean = 31 m (103 ft),
and dominant substrate of coarse gravel to small boulder. Riparian vegetation is
dominated by non-forested alder and willow shrub communities. Glacial output
streams are moderate energy streams that transport large sediment loads. Available
salmonid spawning and rearing habitat is low due to high glacial silt suspended load
and regular flushing events. One study site was classified as a G04 stream: Power
Creek in the Upper Eyak system.

Glacial Ougmt Channel Type: Large Braided Glacial Outwash Channel (G03)

G03 channels occur in broad, glacial valley bottoms or outwash plains in very
large glacial drainage basins. G03 streams are characterized by gradient < 3%,
incision depth _<_ 2 m (6.5 ft), variable bankfull width from 60-300m (200 — 1000 ft),

with a dominant substrate of coarse gravel to large cobble. The riparian vegetation is

dominated by nonforested Sitka alder and willow shrub communities. G03 channels
function as sediment deposition systems with extremely large sediment loads. G03
channels typically provide salmonids with migration routes to spawmng areas in clear
water tributaries. Spawning habitat is limited by fine sediments, and coho salmon rear
in low numbers in these channels. One study site was classified as a G03 stream:
Ibeck creek in the Lower Eyak River system.

Moderate Gradient Contained Channel Type: Narrow Sfihallow Contained Channel
LMQ)

Moderate gradient contained channel types occur in middle to upper valley
positions of glacially scoured lowland landforms. MCl channels are characterized by
stream gradient of l-6%, incision depth of < 4 m (13 ft), bankfull width < 10m (33 ft),
and dominant substrate of cobble and bedrock. Riparian vegetation is dominated by
mixed conifers. The MCI streams function as sediment transport systems with
moderate energy, and instream storage of fine sediments is minor. Salmonid
spawning habitat is limited, but coho salmon and Dolly Varden char use pools for
summer rearing. Two study sites were chosen from this area and classified as Narrow
Shallow Contained Channel (MCI): Upper Pipeline Creek in the Alaganik River
system and 1971 Pond in the 18 Mile system, both of the West Delta.

Palustrine Ch_annel Type: Narrow Placid Flow Channel (PAI)

Narrow placid flow channels on the Delta are characterized by stream gradient
less than 2%, incision depth of less than or equal to 2m (6.5 ft), bankfull width less
than 10m (33ft), and dominant substrate of organic silt to fine gravel. Riparian

vegetation communities are dominated by non-forested sedge, Sphagnum bog, and

10

Sitka alder. Sediment retention is very high in PA] channels so fish spawning
potential is low. Deep, pooled water and cover from overhanging streambank
vegetation provide good rearing habitat for coho salmon and Dolly Varden. Two
study sites were classified as narrow placid flow channels: the Middle Branch of the
18 Mile system, and Salmon Creek of the Alaganik system.
Palustrine Channel Type: Beaver Ponds (PAS)

Beaver pond habitats on the Delta are characterized by a stream gradient of 0-
1%, incision depths less than or equal to 2 m (6.5 ft), bankfull width normally greater
than 10m (33 ft), and dominant substrate of sand and organic silt. Riparian vegetation
is dominated by nonforested communities of sedge and Sphagnum bog with some
Sitka alder (A lnus sinuate) and willow (Salix spp). PAS channels function as sediment
sinks and buffer flows from extreme runoff events. Salmonid spawning potential is
low due to prevalence of fine sediments. PA5 channels provide the necessary depth
and woody debris cover for overwintering and growth making them good rean'ng
habitat for juvenile coho salmon and Dolly Varden char. Two of the sampling sites in
this PAS channel were beaver ponds: Pipeline #4 in the Alaganik River system, and a
second pond in an adjacent system known as Goose Meadows.
Benthic Macroinvertebrates

To estimate abundance and composition Of macroinvertebrate communities,
quantitative samples were collected with a 0.1m2 Hess sampler (Merritt et al. 2008).
During each sampling period, I collected three replicate samples from each stream
reach. Hess samples were taken at random within the first riffle area encountered

upstream Of access point 2 10m2 in area and S 0.3 m depth. The sampler was placed

11

in a shallow, fast-flowing section of stream, and the enclosed substrate was agitated
for ~1 minute allowing the disturbed macroinvertebrates to be washed into the
collection bag at the end of the sampler. Samples were passed through a 250-micron
sieve and transferred into a 250 ml Whirl-Pak©, preserved in 70% ethanol, and
returned to the lab for sorting & identification under a dissecting scope. All
invertebrates were picked from each sample, counted, and identified to the lowest
possible taxonomic level, mostly generic (except Chironomidae) using Merritt et al.
(2008). Chironomidae were identified to subfamily.

Invertebrate density, (mean total number of organisms per square meter), was
estimated from abundance and surface area calculations for the Hess sampler, and
converted to number per m2. Richness was measured as total number of taxa present,
and diversity was measured using the Shannon-Weiner diversity index (- Hauer and
Resh 2006). Macroinvertebrates were designated a functional feeding group status
(shredders, filtering-collectors, gathering-collectors, scrapers, and predators)
according to Merritt et al. (2008). The following metrics were calculated in order to
classify stream channels taxonomically: mean percent EPT taxa (no. of
Ephemeropetera, Plecoptera, and Trichoptera / total # of organisms) , mean percent
Diptera (# of Diptera / total # of organisms), and mean percent non-insect taxa, (# of

non-insect taxa / total # of organisms).

12

Analysis

The main objective of this study was to assess differences in the overall
macroinvertebrate community among streams and stream channel types. As a result,
variation among months was not a focus of this paper and all monthly samples were
combined to calculate overall average macroinvertebrate values for each year (n=12
samples per stream per year). Two-way ANOVAs (IMP 8.0, 2008) were generated
contrasting community measures among sampling sites, between sampling years, and
site by year interactions (or = 0.05). In addition, multiple comparison tests using
Tukeys Honest Significant Difference (HSD) Tests were made among streams (CL =
0.05). All data were tested for normality and (log + I) transformed where necessary to
meet statistical test assumptions. We found no significant differences due to year and
no significant interaction effects in all our analyses; thus to simplify and summarize
differences due to only stream type, samples from both years were combined (n=18),

averaged and listed in all data tables and figures.

13

RESULTS
Macroinvertebrate Richness and Diversity Among Stream Channel Types

Over 29,000 macroinvertebrates representing 52 distinct taxa were collected
from streams of the Copper River Delta (Table 2). A total of 20 taxa were collected
from the insect orders Ephemeroptera, Plecoptera, Trichoptera, and Diptera (Table 2).
In addition, 5 subfamilies of Chironomidae were identified (Table 2). The chironomid
subfamily Orthocladiinae comprised the largest percentage of chironomid abundance
across the 12 streams, followed by the Diamesinae and the Tanytarsinae (Table 4).
Twelve additional insect taxa and fifteen non—insect taxa were also collected from
Copper River Delta streams during 2005 & 2006 (Table 2).

Results showed a significant difference in taxa richness among streams (F =
67.108, df = 11, p = < .0001), however no difference among years (F = 0.725, df = l,
p = 0.396), or an interaction effect (F = 1.283, df = 11, p = 0.236). Mean richness
(total number of taxa collected) was lowest (2-6 taxa) in estuarine (ES) and glacial
output (GO) stream channels. Richness was significantly lower (Tukey HSD, p<0.05)
than higher ranked stream channels with the exception of Salmon Creek (PAl) that
was not significantly different from Hartney Creek (ES) (Figure 2). Mean richness
was intermediate (7-12 taxa) in moderate gradient contained (MC), placid flow (PAl ),
and floodplain (FP) channel types. Within this group, all sites were significantly
greater (Tukey HSD,p<0.05) than lower ranked sites with the exception of Salmon
Creek (PAl) which was not significantly greater than Hartney Creek (ES) (Figure 2).
Mean richness was greatest (1 3-17 taxa) in beaver pond channel types (PAS). Mean

richness at both PAS sites, Goose Meadows and Pipeline # 4, was significantly greater

14

than the low-ranked channel types, but neither were significantly greater than all
intermediate channel types (Figure 2).

I found a significant difference in diversity among streams (F = 44.943, df =
11, p = <.0001); however, no difference among years (F =l.164, df = l, p = 0.2821)
and no interaction effect (F = 0.576, df = 11, p = 0.8470). Mean Shannon-Weiner
diversity was lowest (0.79 — 1.18) in all estuarine, glacial output, and one PAl
channel, Salmon Creek (Figure 3). These sites were significantly lower than higher
ranked sites with the exception of Hartney Creek (ES) and Ibeck Creek (G0) which
were not significantly different from Blackhole Creek (FP). Diversity was
intermediate (1.37-1.93) in all placid flow, floodplain, and in one moderate gradient
contained channel, Pipeline Up. These sites were all significantly greater than (Tukey
HSD, p<0.05) lower ranked sites with the exception of Blackhole Creek (FP) (Figure
3). Shannon-Weiner diversity was greatest (>20) in beaver pond channel types
(PAS). Both Goose Meadows (PAS) and Pipeline # 4 (PAS) were significantly greater
(p<0.05) than the lowest ranked sites, but neither were significantly greater than all
intermediate ranked sites (Figure 3).

Mean percent EPT (Table 3) was low (_<_ 5%) in all beaver pond channels
(PAS), and estuarine channels (ES), intermediate (5 — 10%) in one placid flow channel
(PAl) (Salmon Creek), one glacial output channel (GO) (Power Creek), and one
floodplain charmel (F P) (Blackhole Creek). Mean percent EPT was high (> 10%) in
all moderate gradient channels (MC), one placid flow channel (PAl) (18 Middle), one
glacial output channel (GO) (Ibeck Creek), and one floodplain channel (FP) (Little

Martin) (Table 3).

15

Mean percent Diptera (Table 3) was high (> 60%) in all placid flow channels
(PAl), moderate gradient channels (MC), glacial output channels (GO), floodplain
channels (FP), and one estuarine channel (ES) (Alaganik). Mean percent Diptera was
intermediate for both beaver pond channels (PAS) and low for one estuarine channel
(ES) (Hartney Creek) (5%) (Table 3).

Mean percent non-insect taxa, (Table 3) was high in both beaver pond channels
(PAS), and one estuarine channel (ES) Hartney Creek, intermediate in all floodplain .-
(FP), moderate gradient channels (MC), one estuarine channel (ES) Alaganik, and one
placid flow channel (PAl) 18 Middle. Mean percent non-insect taxa was low in both
glacial output channels (GO), and one placid flow channel (PAI) Salmon Creek
Macroinvertebrate Densities Among Stream Channel Types

A statistically significant difference in macroinvertebrate density among
streams (F = 27.783, df = 11, p = <.0001) was identified; however, no difference
among years (F = 0.139, df = 1, p = 0.7094) and no interaction effect (F = 0.411, df =
11, p = 0.9497) was observed. Mean densities of macroinvertebrates were lowest (100
— 999 individuals/m'z) among all glacial output streams, one estuarine stream,
Alaganik Slough (ES), and one moderate gradient contained stream, Pipeline Up (MC)
(Figure 4). Within this group the mean density of Ibeck Creek (G0) was significantly

lower (Tukey HSD, p<0.05) than the other sites. Mean densities were intermediate

(1000-1999 individuals/ma) in all PAl , beaver pond (PAS), one floodplain site,

Blackhole Creek (F P), and one moderate gradient contained site, 1971 Pond (MC).
Multiple comparison tests indicated that intermediate ranked streams were

significantly greater. than lower ranked sites with the exception of 1971 Pond (MC)

16

(Figure 4). Mean density was greatest (>2000 individuals/ma) in one estuarine site,
Hartney Creek (ES), and one floodplain site, Little Martin (F P). Mean density of both
of these sites was significantly greater (Tukey HSD, p<0.05) than all low ranked
streams, but only Hartney Creek (ES) was significantly greater than all intermediate
ranked streams (Figure 4).
Functional Feeding Group Proportions Among Stream Channel Types

Functional feeding groups and their proportions varied both across and within
stream channel types (Figure 5; Table 5). Estuarine channel types (ES) were
characterized by relatively few taxa in large numbers. Alaganik Slough (AG) was
dominated by collector-gatherers (86%) (Figure 5) of the chironomid subfamily
Orthocladiinae (Table 5), with some scrapers (12%) (Chironomidae: Diamesinae), and
few collector-filterers (1%), shredders (<1%), and predators (<l%). Hartney Creek
(HC) was characterized by a high percentage of shredders (61%) (euryhaline
Amphipoda and Isopoda), collector-gatherers (38%) (Oligochaete worms), few
scrapers (1%), and the absence of collector-filterers and predators (Figure 5). The
functional communities of floodplain channel types, Blackhole Creek (BH) and Little
Martin (LM), were composed of similar macroinvertebrate assemblages and included
collector-gatherers (BH = 57%; LM=44%) (Chironomidae: Orthocladiinae, - Baetis
mayflies and Oligochaete worms), scrapers (BH = 23%; LM = 26%) (Chironomidae:
Diamesinae, planorbid snails and Cinygmula mayflies), collector-filterers (BH = 5%;
LM = 16%) (Simulium black flies and Chironomidae: Tanytarsinae), predators (~ 10%
both sites) (Chironomidae: Tanypodinae, Ceratopogonidae and Haploperla and

Isoperla stoneflies), and shredders (~ 5% both sites) (Zapada stoneflies and

17

Ecclisomyia caddisflies) (Figure 5; Table 5)). The glacial output channel types, Ibeck
Creek (IC) and Power Creek (PC), were dominated by collector-gatherers (IC = 47%;
PC = 74%) (Chironomidae: Orthocladiinae), followed by scrapers (IC = 39%; PC =
21%) (Chironomidae: Diamesinae), predators (IC = 12%; PC = 3%) (Haploperla and
Isoperla stoneflies), and few filterers (~ 2%), and shredders (~ 1%) (Figure 5; Table
5). Moderate gradient contained sites, 1971 Pond (19P) and Pipeline Up (PU), were
characterized by similar proportions Of shredders (7%) (Lepidostoma, Onocosmoecus,
Psychoglypha caddisflies, and Zapada stoneflies), scrapers (15 % — l7 %)
(Chironomidae: Diamesinae and Planorbid snails), and predators (10 % - 13 %)
(Chironomidae: Tanypodinae and Ceratopogonid biting midges). Moderate gradient
sites differed in the percentages Of collector-gatherers (19P = 57%; PU = 31%)
(Chironomidae: Chironominae, Orthocladiinae, and Paraleptophlebia mayflies) and
collector-filterers (19P = 8%; PU = 35%) (Simulium blackflies, Chironomidae:
Tanytarsinae, and sphaeriid fingernail clams) (Figure 5; Table 5). Placid flow channel
types, 18 Middle (18M) and Salmon Creek (SC), differed in percentages of collector-
gatherers (18M = 51%; SC = 34%) (Chironomidae: Orthocladiinae and Baetis
mayflies), scrapers (18M = 12%; SC = 60%) (Chironomidae: Diamesinae), and
collector-filterers (18M = 26%; SC = 1%) (Simulium blackflies and Chironomidae:
Tanytarsinae). Placid flow channel types showed similar percentages of predators
(4% at both sites) (Chironomidae: Tanypodinae, — Haploperla and Isoperla stoneflies),
and shredders (18M = 7%; SC = 1%) (Zapada stoneflies and Ecclisomyia and
Onocosmoecus caddisflies) (Figure 5; Table 5). Beaver pond channel types (Figure 5;

Table 5)), Goose Meadows (GM) and Pipeline #4 (P4), were very similar in

18

taxonomic assemblages of all functional feeding groups including collector-gatherers
(28%-30%) (Copepoda, Hydrachnida, and Chironomidae: Orthocladiinae), collector-
filterers (46% - 47%) (Cladocera, Sphaeriidae, and Chironomidae: Tanytarsinae),
shredders (12% -l6%) (Ostracoda, Amphipoda, Haliplus beetles, and Onocosmoecus
caddisflies), scrapers (5% - 8%) (Chironomidae: Diamesinae; planorbid and valvatid
snails), and predators (4% at both sites) (Chironomidae: Tanypodinae;

Ceratopogonidae; Dytiscidae; Aeshnidae)

19

DISCUSSION
Macroinvertebrate communities of the Copper River Delta.

The results of this study showed that taxonomic and functional differences in
macroinvertebrate composition occurred among stream channel types of the Copper
River Delta. Studies that compared streams of Glacier Bay, Southeast Alaska indicated
that community development was mainly influenced by abiotic factors, especially
water temperature and channel stability (Milner 1987; Sidle and Milner 1989). In
general, invertebrate densities, richness, and diversity were lowest in Delta streams
and channel types characterized by high gradient and stable substrate (glacial output
channels), or low gradient and unstable substrate (estuarine channels). Invertebrate
densities, richness, and diversity were highest in Delta streams and channel types
characterized by low gradient and stable organic substrates (beaver pond and
floodplain channels) (Figure 1). The Copper River Delta pulse study (Bryant 1991)
qualitatively compared diversity of beaver ponds with main channel habitats with
similar results. Estuarine and glacial output stream channel types were characterized
by low densities, low taxa richness, low diversity, and few functional feeding groups
of freshwater macroinvertebrates. Estuarine stream channel types (ES), Alaganik
Slough and Hartney Creek, were characterized by the dominance of relatively few taxa
in large numbers. Habitat diversity at Alaganik Slough was dominated by unstable
sand/silt restricting the invertebrate community to collector-gatherers of the
chironomid subfamily Orthocladiinae. Hartney Creek was immediately adjacent to the
estuary at Hartney Bay, and underwent tidal inundation daily. Exposure to saltwater

allowed the colonization of the substrate by euryhaline arnphipod and isopod

20

shredders, and effectively eliminated freshwater macroinvertebrates from the stream
channel. Glacial output stream channels (GO), Ibeck Creek and Power Creek, were
characterized by high glacial suspended sediment loads and contained the lowest
densities, richness, and diversity of all stream sites examined. Both of these stream
channel types were dominated by chironomid collector-gatherers (Orthocladiinae) and
scrapers (Diamesinae). Moderate gradient contained stream channel types (MC),
Pipeline Up and 1971 Pond, were characterized by low invertebrate density, high taxa
richness, high invertebrate diversity, and presence of all five functional feeding
groups. Accumulations of organic debris provided habitat for Paraleptophlebia
(Leptophlebiidae) mayfly gatherers and caddisfly, Lepidostoma (Lepidostomatidae),
shredders which were not found at other stream channel types (Table 5). Placid flow
stream channels (PAl), Middle 18 and Salmon Creek, were characterized by similar
densities and taxa richness, but differing diversities of functional feeding groups. The
Middle 18 stream habitat consisted of gravel riffles with an associated fauna
consisting of Orthocladiinae (Chironomidae) and Baetis collector-gatherers, Simulium
collector-filterers, chironomid (Diamesinae) scrapers, limnephilid caddisfly shredders
(Ecclisomyia and Onocosmoecus), and predatory chironomid midges (Tanypodinae).
The dominant habitat of Salmon creek also consisted of gravel riffles, but this site was
dominated by chironomid scrapers (Diamesinae) and collector-gatherers
(Orthocladiinae). Salmon Creek was located in close proximity to the mouth of
McKinley Lake which could have acted as a source of chironomid influx into the
surrounding aquatic habitat. Milner et al 2000, found that percent Chironomidae was

significantly greater downstream of lakes in stream habitats in Glacier Bay, Southeast

21

Alaska. Floodplain stream channels (FP), Blackhole Creek and Little Martin River,
were characterized by high invertebrate densities, taxa richness, diversity, and the
presence of all five functional feeding groups. Habitat at both floodplain stream sites
was similar and dominated by gravel riffles. The fauna of both floodplain streams was
similar with the exception of the scraper and shredder communities which differed in
taxa (Table 5). Beaver pond stream channel types (PAS), Goose Meadows and
Pipeline #4, were characterized by high invertebrate densities, taxa richness, diversity,
and the presence of all five functional feeding groups. Habitat at both beaver pond
sites was similar and dominated by organic silt and sand substrates. The fauna of both
beaver pond sites was similar and was dominated by fauna more indicative of lentic
habitats (Table 5). In addition, beaver ponds were found to contain invertebrate taxa
not normally found in other stream channel types including predatory diving beetles
(Dytiscidae), immature dragonflies (Aeshnidae), hydroptilid caddisflies (Oxyethira),
and the dipteran Pericoma (Psychodidae). This may have been due to a more lentic
environment conducive to colonization by the above taxa.
The Copper River Delta and other regions of Alaska.

Invertebrate densities and diversity in Copper River Delta streams were found
to be lower than those reported for other streams and rivers in Alaska (Milner et al.

2000, Hernandez et al. 2005). With one exception, Copper River Delta streams were

found to have mean density values less than 2500 m-z; whereas, macroinvertebrate
densities for gravel riffle habitats ranged from 3750 to 5700 m'2 for streams of Glacier

Bay, Alaska (Milner et al 2000), and from 3000 to 6500 m-2 for streams on Prince of

22

Wales Island, Alaska (Hernandez et a1 2005). The exception, the estuarine stream
channel, Hartney Creek, was immediately adjacent to Hartney Bay estuary, and
received daily tidal inundation which allowed euryhaline arnphipods and isopods to
colonize the channel, thus artificially increasing the total invertebrate density for the
site. Macroinvertebrate diversity also was lower in Copper River Delta streams (S2
taxa) as compared to Glacier Bay streams where 128 taxa were recorded (Milner et al.
2000)

Invertebrate communities of stream channels of the Copper River Delta were
dominated by taxa indicative of new or disturbed systems in Alaska. Mackay (1992)
suggested that the first macroinvertebrate colonizers of new or disturbed systems
would typically be Baetidae and Leptophlebiidae (Ephemeroptera), Simuliidae,
Orthocladiinae (Chironomidae), and Hydropsychidae (Trichoptera). Taxonomically,
most streams of the Copper River Delta are characterized by dominance of the
chironomid subfamilies Orthocladiinae and/or Diamesinae. Milner et a1 (2000),
during a colonization and development study of aquatic habitats representing a time
span of 200 years in Glacier Bay, Alaska, found that Orthocladiinae and Diamesinae
were the dominant groups of Chironomidae in young (5 50 years) streams with lakes
and non-lake systems respectively. The genus Baetis (Baetidae) dominated the
ephemeropteran taxa collected from most Copper River Delta stream channels with
the exception of moderate gradient channels which were dominated by
Paraleptophlebia (Leptophlebiidae). The genus Cinygmula (Heptageniidae) was
collected from only one stream, the floodplain channel Blackhole Creek, and the

genus Epeorus (Heptageniidae) was collected from only one stream, the glacial output

23

channel Power Creek. The common genera of several southeast Alaskan streams
(i.e.,Drunella (Ephemerellidae), Rhithrogena (Heptageniidae), Ameletus (Ameletidae
(Lessard and Merritt 2006, Lessard et al. 2009), were not collected from Copper River
Delta streams. Milner et al (2000) found that among the Ephemeroptera genera found
in Glacier Bay streams, only Baetis was collected from streams less than 50 years old;
C inygmula was added to the fauna in streams between 50 and 100 years; Drunella,
Epeorus, and Rhithrogena between 100 and 150 years; and Ameletus was found in the

Oldest post-glacial streams examined (150 - 200 years).

24

' CONCLUSION

The results of this study suggested that macroinvertebrate communities vary in
density, taxonomic diversity, and functional feeding group diversity among stream
channel types of the Copper River Delta, southcentral Alaska. Main channel habitats
were found to be characterized by high densities of a few invertebrate taxa
representing selective functional groups. Densities were reduced, but taxonomic and
functional feeding group diversity was greater in interbasin channel types.
Macroinvertebrate taxa found in streams of the Copper River Delta are also indicative
of those found after recent disturbance (< 50 years).

The Copper River Delta is a system driven by recent geologic disturbance.
Understanding how disturbance affects aquatic macroinvertebrate communities will
allow managers to make informed decisions regarding resource use. There is a need
for long term monitoring of macroinvertebrate communities on the Copper River Delta
in order to understand the effects Of time and succession upon this important aquatic

ecosystem.

25

Table 1. Physical stream characteristics of study sites within the Copper River
Delta, Alaska, (modified from Paustian I991).

 

 

Stream Channel Gradient Bankfull Substrate Salmonid
Type (%) Width Type Potential
(m)

Alaganik ES 0.5 8 silt/sand Low

Hartney ES 1 23 gravel/cobble Low

BlackHOle

Creek F P 1 6 sand/ gravel High

Little Martin FP 1 6 sand/gravel High

Power Creek GO 4 31 cobble/boulder Low

Ibeck Creek GO 2 65 gravel/cobble Low

Upper Pipeline MC 3 6 cobble/bedrock Moderate
organic

l97l Pond MC 1 4.6 silt/gravel Moderate

Salmon Creek PA] 1 6 sand/ gravel Moderate
organic

Middle 18 PA] 1 4.6 silt/gravel Moderate

Pipeline #4 PAS 1 > 10 m organic silt/sand High

Goose

Meadows PAS l > 10 m organic silt/sand High

 

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Table 5. Dominant invertebrate taxa by functional feeding group among Copper River
Delta streams across channel type and year (2005-2006).

 

 

Stream Channel Collector Collector Scrapers Shredders Predators
Type Gatherers Filterers
Alaganik ES Orthocladiinae * Diamesinae * *
Hartney ES Oligochaeta * * Amphipoda *
Power Crk GO Orthoc ladiinae * Diamesinae * Haploperla
Isoperla
Ibeck Crk GO Orthoc ladiime * Diamesinae * Haploperla
Isoperla
Pipeline Up MC Orthocladiinae Simulium Diamesinae Lepidostoma Tanypodime
Paraleptophlebia Planorbidae Zapada Ceratopogon
1971 Pond MC Chironomime Tanytarsime Diamesinae Onocosmoecus Tanypodinae
Paraleptophlebia Planorbidae Psychogbpha Ceratopogon
BlackHole FP Orthoc ladiinae Simulium Diamesinae Ecclisomyia Tanypodinae
Baetis Tanytarsinae C inygmula Ceratopogon
Lt Nhrtin FP Orthocladiinae Sirmdium Diamesinae Zapada quloperla
Baetis Tanytarsinae Planorbidae Isoperla
Salmon Crk PAl Orthocladiinae Tanytarsime Diamesinae Zapada I-lqvloperla
Baetis Isoperla
Middle 18 PA] Orthocladiinae Simulium Diamesinae Ecclisonyia Tanypodinae
Baetis Onocosmoecus
Pipeline #4 PAS Orthocladiinae Cladocera Diamesinae Ostracoda Tanypodinae
Copepoda Sphaeriidae Planorbidae Haliplus Ceratopogvn
Goose PAS chocladiinae Cladocera Diamesinae Ostracoda Tanypodime
Copepoda Splneriidae Planorbidae Haliplus Ceratopogon

 

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APPENDIX

40

Appendix 1
Record of Deposition of Voucher Specimens*

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

Voucher No.: 2009-07

 

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

BENTHIC COMMUNITY MEASURES AMONG STREAM CHANNEL TYPES OF
THE COPPER RIVER DELTA, SOUTHCENTRAL 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)

TODD C. WHITE

 

 

Date 15. XII 2009

 

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

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

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

Research project files.

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

41

Appendix 1.1

Voucher Specimen Data

3 Pages

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Page 1

23

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Appendix 1.1

Voucher Specimen Data

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43

Appendix 1.1

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23

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44

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45

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