AQUATIC MACROINVERTEBRATE and DIATOM SURVEYS in TETLIN NATIONAL WILDLIFE REFUGE, AK Final Report December 2012

AQUATIC MACROINVERTEBRATE AND DIATOM SURVEYS IN TETLIN NATIONAL WILDLIFE REFUGE, AK Final Report December 2012

Prepared by: Daniel Rinella, Daniel Bogan, and Rebecca Shaftel

Alaska Natural Heritage Program University of Alaska Anchorage Beatrice McDonald Hall 3211 Providence Dr. Anchorage, AK 99508

Tetlin Macroinvertebrate and Diatom Surveys

Table of Contents

INTRODUCTION 1

METHODS 1

Stream habitat measurements 3

Macroinvertebrate and diatom sampling 4

Sample processing 4

Data analysis 5

RESULTS AND DISCUSSION 6

CONCLUSIONS 16

REFERENCES 17

Tetlin Macroinvertebrate and Diatom Surveys

Tables

Table 1. Selected habitat characteristics for Desper, Gardiner, and Scottie Creeks...... 8

Table 2. Average taxa richness for macroinvertebrate and diatom quantitative and multi-habitat samples

over the four year sampling period. Quantitative samples were pooled to calculate richness...... 8

Table 4. Jaccard (persistence) and Bray-curtis (stability) distances for diatom communities from Desper,

Gardiner, and Scottie creeks based on three inter-annual comparisons from 2007 to 2010...... 15

Figures

Figure 1. Sampling sites in Tetlin National Wildlife Refuge...... 2

Figure 2. Photos of Desper Creek (A), Gardiner Creek (B), and Scottie Creek (C)...... 7

Figure 3. Hydrologic conditions at the time of sampling as indicated by stage and discharge readings for Desper, Gardiner, and Scottie creeks...... 9

Figure 4. Macroinvertebrate abundances from the quantitative samples for the six most frequently encountered taxa in Desper, Gardiner, and Scottie creeks from 2007 to 2010...... 11

Figure 5. Diatom abundances from the quantitative samples for the six most frequently encountered taxa in Desper, Gardiner, and Scottie creeks from 2007 to 2010...... 12

Figure 6. NMS ordination for macroinvertebrates at Desper, Gardiner, and Scottie creeks over four year sampling period...... 13

Figure 7. NMS ordination for diatoms at Desper, Gardiner, and Scottie creeks over four year sampling period...... 14

Appendices

Appendix 1. Macroinvertebrate taxa list

Appendix 2. Diatom taxa list

Tetlin Macroinvertebrate and Diatom Surveys

INTRODUCTION This report summarizes the results from macroinvertebrate and diatom surveys conducted in Desper, Gardiner, and Scottie creeks; lowland streams in Tetlin National Wildlife Refuge, interior Alaska. These three streams were sampled repeatedly during the summers of 2007 through 2010. The objectives of this study were to characterize the stream biological communities for streams in the Tetlin Refuge where data currently does not exist. Sites were sampled over several years in order to describe inter-annual changes in community composition.

Macroinvertebrates and diatoms have been used in biomonitoring to indicate the quality of the aquatic ecosystems in which they reside. Since they integrate attributes of their chemical and physical environments over their lifespans, taxa presence and abundance can be better predictors of past environmental conditions than snapshot water quality measurements. Potential disturbances to streams in this region include climate change, development along the Alaska Highway system, and the proposed Alaska natural gas pipeline.

In addition to describing macroinvertebrate and diatom community composition, the inter- annual persistence and stability of macroinvertebrate and diatom assemblages are important considerations for monitoring programs. Persistence measures the presence or absence of individual taxa over time, while stability includes both the presence/absence and the abundance of taxa over time. Aquatic habitats characterized by high community persistence and stability facilitate the detection of ecological changes over time. Low persistence and stability among macroinvertebrate communities in montane streams of Denali National Park, also in interior Alaska, was found to limit their applicability for ecological monitoring (Milner et al. 2006). Macroinvertebrate persistence and stability in the Denali streams was related to winter snow depth and channel stability (Milner et al. 2006), factors which may also affect streams in the Tetlin refuge. To our knowledge, no prior published studies have examined persistence and stability among diatom communities.

METHODS Sites were selected based on their accessibility via the Alaska Highway. Figure 1 shows the location of the three sampling sites within the Tetlin National Wildlife Refuge.

Tetlin Macroinvertebrate and Diatom Surveys

Figure 1. Sampling sites in Tetlin National Wildlife Refuge.

Tetlin Macroinvertebrate and Diatom Surveys

Stream habitat measurements

Stream habitat measurements were collected in the first year of sampling (2007) to characterize differences in channel morphology, riparian vegetation, and substrate composition between the three streams. We conducted habitat assessments using protocols from EPA’s Environmental Monitoring and Assessment Program (EMAP, Kaufman and Robinson 1998) to characterize in-stream and riparian habitat at each of 11 evenly spaced transects along the stream reach. Each stream reach was 150 m in length. Measurements collected at each transect included channel wetted width, bankfull width, bank height, bank angle, and bank undercut distance. We measured riparian canopy coverage with six densiometer readings along each transect. We recorded water depth, substrate size class, and embeddedness at five points along each transect. To characterize fish cover we estimated the extent of filamentous algae, macrophytes, big and small woody debris, live trees or roots, overhanging vegetation, undercut banks, and boulders at each transect using five areal cover classes (0%, <10%, 10-40%, 40-75%, or >75%). Between each of the 11 transects, we counted pieces of large woody debris within and above the bankfull channel according to several size classes. We characterized riparian vegetation cover separately for canopy, understory, and ground cover along the entire stream reach using five areal cover classes (0%, <10%, 10-40%, 40-75%, or >75%) and several types (deciduous, coniferous, mixed, or none). To characterize channel slope and sinuosity, we recorded the compass azimuth (aspect) and channel slope between each pair of transects.

The U.S. Fish and Wildlife Service collects continuous hydrologic stage measurements at each of these creeks in order to estimate discharge, in addition to collecting periodic water quality samples throughout the year. For Scottie and Desper creeks, stage measurements were provided for each sampling date and for Gardiner Creek, preliminary discharge measurements were provided. These measurements will enable comparisons within each creek between the hydrologic conditions and the aquatic biological communities over the four year sampling period. A total of 15, 17, and 18 water quality samples were collected at Desper, Gardiner, and Scottie creeks from June 2006 to September 2009. The suite of constituents measured included hardness, pH, nutrients, dissolved organic carbon, specific conductance, numerous metals, and various inorganic salts. For this report, water quality parameters known to affect stream biological communities were summarized for the three streams to qualitatively describe differences in their chemistry.

Tetlin Macroinvertebrate and Diatom Surveys

Macroinvertebrate and diatom sampling

For both macroinvertebrates and diatoms we collected five quantitative subsamples from the benthic habitat in order to characterize variation in taxa abundance within a stream. Stream substrates were targeted for the quantitative subsamples so that the sampling area could be easily defined and measured, which is more difficult when sampling other habitats, such as streambanks or overhanging vegetation. Quantitative macroinvertebrate samples were collected using either a modified Surber sampler with 350-µm mesh in shallow water or an Eckman grab for deep water with fine substrates. Quantitative diatom samples were collected by scrubbing a fixed area (delineated by a bedrock sampler) on submerged woody snags or by skimming the top layer of streambed sediments (delineated by a sediment sampler).

We also collected semi-quantitative multi-habitat samples of macroinvertebrates and diatoms that we expected might provide a more comprehensive characterization of the biological communities. For both macroinvertebrates and diatoms, the multi-habitat sample was composed of 20 subsamples taken throughout the stream reach. All substrate types (i.e., sand/fine substrates, gravel/cobble substrates, undercut streambanks, woody snags, and macrophyte beds) were sampled in proportion to their abundance within the stream. We used a long-handled D-frame net (350-µm mesh) to collect the macroinvertebrate subsamples. Diatom subsamples were collected by scrubbing hard substrates or by skimming the top layer from soft sediments.

Sample processing

We preserved all macroinvertebrate samples in the field with ethanol and returned them to UAA’s Aquatic Ecology lab for processing. In the lab, we subsampled each multi-habitat macroinvertebrate sample to obtain a fixed count of 300 ±20% organisms to standardize the taxonomic effort across all sites. In addition, we conducted a five minute search through the remaining sample to select any large or rare taxa that may have been missed during subsampling. In the lab, we sorted all macroinvertebrates from each subsample under magnification. For the quantitative macroinvertebrate samples, all five subsamples were sorted in their entirety and all individuals identified. All insects were identified to genus or lowest practical taxonomic level, including Chironomidae, and non-insects to a higher taxonomic level (usually family or order) using standard taxonomic keys (Weiderholm 1983, Pennak 1989, Merritt and Cummins 1996, Wiggins 1996, Thorpe and Covich 2001, Stewart and Oswood 2006). 4

Tetlin Macroinvertebrate and Diatom Surveys

We homogenized each diatom subsample and cleared the samples with nitric acid for easier identification. We then neutralized the acid-digested aliquots by a succession of dilutions, concentrated the cleared diatom valves by allowing them to settle, and slide mounted the valves. From each subsample, we identified a fixed count of 600 valves to species or lowest practical taxonomic unit (Patrick and Reimer 1975, Krammer and Lange-Bertalot 1986-1991).

Data analysis

To determine the extent to which our quantitative samples adequately characterized the stream biological communities, we calculated the total taxa richness for each of the five subsamples for both macroinvertebrates and diatoms from each of the three streams and sampling dates. Subsamples were sorted in order of increasing richness and the difference in richness was calculated between the 3rd and 4th and the 4th and 5th subsamples to determine whether the amount of area sampled (as represented by the number of subsamples) was adequate to characterize the community. For each stream biological community, means and standard deviations were calculated for the increase in richness provided by the 4th and 5th subsamples (n = 60; 5 samples x 3 streams x 4 years).

In addition, to ensure that laboratory subsampling of the multi-habitat diatom samples did not underestimate the number of taxa present in the larger sample, we identified 1000 diatom valves (as opposed to the normal 600) from one sample and plotted the relationship between differing levels of taxonomic effort and taxa richness. The point at which the curve flattens indicates the minimum taxonomic effort required to characterize the diatom assemblage. For this analysis we used the sample from Scottie Creek in 2007 since it had the highest diatom richness.

The quantitative subsamples were used to explore the variation in abundance of individual macroinvertebrate and diatom taxa within a stream and over time. Mean abundances were plotted over time for each stream for the six most common (by frequency) macroinvertebrate and diatom taxa.

In order to explore differences in diatom and macroinvertebrate community composition between streams over time, we used non-metric multidimensional scaling (NMS). NMS is an ordination method helpful for reducing complex, multivariate endpoints into a smaller set of axes (usually 2 or 3 dimensions). Multi-habitat samples were used for the NMS ordinations because they represented taxa inhabiting all of the available stream habitats and thus were more reflective of the entire stream community than the quantitative samples. Prior to analysis, macroinvertebrate and diatom abundances

Tetlin Macroinvertebrate and Diatom Surveys

were log transformed [log10(x+1)] to lessen the influence of abundant taxa. We used Bray-Curtis distances for the macroinvertebrate and diatom NMS ordinations. Successional vectors were added to the ordination to indicate the direction and magnitude of change in community composition for each stream over time.

We quantified community persistence and compositional stability based on differences in community composition for successive years in each stream. Following methods described in earlier studies (Townsend et al. 1987, Milner et al. 2006), we quantified persistence using between-year Jaccard’s dissimilarity (J), a distance measure based on the presence or absence of individual taxa. We quantified compositional stability using between-year Bray-Curtis dissimilarity (BC), a distance measure that based on the abundances of individual taxa. For both J and BC, potential values range from 1 (no similarity, low persistence or stability) to 0 (identical samples, high persistence or stability). Both quantitative and multi-habitat samples were used to calculate persistence and stability in order to determine if the field methodology affected measurements of persistence and stability over time. All analyses were performed in the R statistical platform (version 2.14.1; R Development Core Team, Vienna, Austria) using the vegan library (Oksanen et al. 2011).

RESULTS AND DISCUSSION Habitat measurements collected in 2007 showed that all three stream beds had low gradients and were composed entirely of fine sediments, although they differed in their channel and riparian habitat characteristics. Gardiner Creek had the deepest channel incision, the highest canopy coverage, the highest amount of woody debris, and low aquatic macrophyte cover (Figure 2, Table 1). Desper Creek had the least channel incision, the least canopy coverage, and the highest macrophyte cover. Scottie Creek was found to be intermediate with regards to the sampled habitat characteristics.

Tetlin Macroinvertebrate and Diatom Surveys

Total taxa richness calculations for the quantitative and multi-habitat samples are shown in Table 2 for macroinvertebrates and diatoms. Both sampling methods captured similar taxa richness for the macroinvertebrate community in Gardiner and Scottie creeks. In Desper Creek, the multi-habitat sampling method increased the total richness by an average of nine taxa over the four year sampling period, which may be due to a higher diversity of habitats in Desper Creek. Total taxa richness was consistently higher in the quantitative samples for diatoms due to the larger laboratory effort expended on the quantitative samples versus the multi-habitat samples (3000 valves counted versus 600 valves counted). The fixed count of 1000 valves on the Scottie Creek sample from 2007 indicated that the 600 valve fixed count was adequate to characterize the diatom community as few taxa (<5) were added by counting the additional 400 valves. A total of 100 unique macroinvertebrate taxa and 198 unique diatom taxa were identified from the quantitative and multi-habitat samples.

Figure 2. Photos of Desper Creek (top two panels), Gardiner Creek (middle two panels), and Scottie Creek (bottom two panels).

Tetlin Macroinvertebrate and Diatom Surveys

Table 1. Selected habitat characteristics for Desper, Gardiner, and Scottie Creeks.

Habitat variables Desper Gardiner Scottie Channel habitat characteristics (mean ± sd) Canopy cover (%) 12.6 + 22.2 49.8 + 36.4 24.3 + 34.3 Channel slope (%) 1.2 + 0.5 1.2 + 0.5 1.3 + 0.6 Fine substrate (%) 100 100 100 Wetted width (m) 13.3 + 1.8 10.2 + 1.3 20.1 + 7.4 Bankfull width (m) 17.4 + 3.1 11.2 + 1.8 22.5 + 8.0 Bankfull height (m) 0.5 + 0.2 1.0 + 0.2 1.4 + 0.2 Incised height (m) 1.5 + 0.3 4.0 + 0.5 2.7 + 0.2 Fish cover types (%) Macrophytes 47 2 5 Woody debris 7 12 8 Brush 12 17 7 Overhanging vegetation 21 7 5 Undercut bank 0 5 3 Large woody debris (total per stream reach) 0.1 – 0.3 m diameter 62 136 48 0.3 – 0.6 m diameter 8 27 28 0.6 – 0.8 m diameter 0 0 1

Table 2. Average taxa richness for macroinvertebrate and diatom quantitative and multi-habitat samples over the four year sampling period. Quantitative samples were pooled to calculate richness.

Community Stream Sample Type Mean Std. Dev. Macroinvertebrates Desper Quantitative 19.3 5.1 Macroinvertebrates Desper Multi-habitat 28.5 7.0 Macroinvertebrates Gardiner Quantitative 22.0 8.8 Macroinvertebrates Gardiner Multi-habitat 22.5 9.4 Macroinvertebrates Scottie Quantitative 18.5 6.2 Macroinvertebrates Scottie Multi-habitat 17.8 7.4 Diatoms Desper Quantitative 86.8 8.1 Diatoms Desper Multi-habitat 44.0 9.2 Diatoms Gardiner Quantitative 75.5 12.4 Diatoms Gardiner Multi-habitat 42.5 7.5 Diatoms Scottie Quantitative 90.0 9.6 Diatoms Scottie Multi-habitat 57.0 13.1

Tetlin Macroinvertebrate and Diatom Surveys

Hydrologic conditions recorded by USFWS on the sampling date for each of the three streams are presented in Figure 3. Preliminary discharge readings for Gardiner Creek indicate that hydrologic conditions during 2009 and 2010 were much wetter than during 2007 and 2008, possibly due to spring melt as sampling was conducted one to two weeks earlier in those years. The stage readings for Scottie Creek also indicate higher water elevations in 2009 and 2010, while Desper Creek had similar water elevations throughout the four year sampling period.

18 400 16 350 14 300

12 250 10 200 8 150 6 Desper stage feet/second)

Stage (feet) Stage 100 4 Scottie stage (cubicDischarge

2 Gardiner discharge50 0 0 2007 2008 2009 2010

Figure 3. Hydrologic conditions at the time of sampling as indicated by stage and discharge readings for Desper, Gardiner, and Scottie creeks.

Trends in abundance for the most common macroinvertebrate taxa show large variability both across the three streams and over time (Figure 3). Scottie Creek tended to have the largest abundances of macroinvertebrate taxa during the sampling conducted in 2010, except for baetid mayflies, which peaked in 2008. Gardiner Creek had an explosion of chironomids (non-biting midges) in 2010, which was likely due to the depths sampled that year. In 2007 and 2008, all of the replicate quantitative 9

Tetlin Macroinvertebrate and Diatom Surveys samples were collected in depths less than or equal to one foot deep. In 2009, replicate samples were collected at depths from 3 to 5 feet while in 2010, sample depths ranged from 5 to 16.5 feet (mean = 9.6 feet). Sphaerid clams and ceratopogonids (biting midges) were most abundant in Desper and Scottie creeks as compared to Gardiner Creek. Diatom abundances tended to vary less across streams, across replicates within each stream, and over time than the macroinvertebrate abundances (Figure 5). Variability in the diatom data represents relative abundances rather than the absolute abundances measured for macroinvertebrates, which reduces the scale of differences both across and within streams.

Tetlin Macroinvertebrate and Diatom Surveys

Figure 4. Macroinvertebrate abundances from the quantitative samples for the six most frequently encountered taxa in Desper, Gardiner, and Scottie creeks from 2007 to 2010.

Tetlin Macroinvertebrate and Diatom Surveys

Figure 5. Diatom abundances from the quantitative samples for the six most frequently encountered taxa in Desper, Gardiner, and Scottie creeks from 2007 to 2010.

Tetlin Macroinvertebrate and Diatom Surveys

The macroinverertebrate NMS ordination indicated that the community composition in Desper Creek was different from the communities found in Scottie and Gardiner Creeks, which had similar composition over the four year sampling period (Figure 6). Community composition for all three streams tended to increase along the first axis of the NMS ordination over the four year sampling period, indicating they may have been responding to similar changes in their environments over time. The diatom NMS ordination indicated that the three streams differed in their community composition over the four year sampling period and stability was highest at Scottie Creek (Figure 7). Both Desper and Scottie creeks tended to increase along the second axis of the NMS ordination over the four year sampling period, while Gardiner Creek tended to shift negatively along the second axis from 2007 to 2009, but in a similar direction as the other two creeks from 2009 to 2010.

Figure 6. NMS ordination for macroinvertebrates at Desper, Gardiner, and Scottie creeks over four year sampling period.

Tetlin Macroinvertebrate and Diatom Surveys

Figure 7. NMS ordination for diatoms at Desper, Gardiner, and Scottie creeks over four year sampling period.

The quantitative macroinvertebrate samples had higher (closer to 0) community persistence (presence of taxa in the community over time) than the multi-habitat samples for both Desper and Scottie creeks, but were comparable between the two sampling methods for Gardiner Creek (Table 3). Stability was higher in Gardiner and Scottie creeks in the multi-habitat samples (presence and abundance of taxa in the community over time), whereas in Desper Creek, it was higher in the quantitative sample. Generally, the amount of variation (as measured by the standard deviation) was higher for community stability than for persistence in the macroinvertebrate communities sampled over time. For diatoms, the quantitative samples consistently had higher persistence and stability (closer to 0) than the multi-habitat samples (Table 4). In addition, the diatom communities were more persistent and stable and had lower variation across the sampling period than the macroinvertebrate communities.

Tetlin Macroinvertebrate and Diatom Surveys

Table 3. Jaccard (persistence) and Bray-curtis (stability) distances for macroinvertebrate communities from Desper, Gardiner, and Scottie creeks based on three inter-annual comparisons from 2007 to 2010. Distance Measure Stream Sample Type Mean SD Jaccard Desper multi-habitat 0.72 0.05 Jaccard Desper quantitative 0.52 0.09 Jaccard Gardiner multi-habitat 0.70 0.03 Jaccard Gardiner quantitative 0.72 0.12 Jaccard Scottie multi-habitat 0.82 0.04 Jaccard Scottie quantitative 0.59 0.07 Bray-Curtis Desper multi-habitat 0.75 0.11 Bray-Curtis Desper quantitative 0.52 0.15 Bray-Curtis Gardiner multi-habitat 0.66 0.18 Bray-Curtis Gardiner quantitative 0.76 0.15 Bray-Curtis Scottie multi-habitat 0.62 0.13 Bray-Curtis Scottie quantitative 0.78 0.10

Table 4. Jaccard (persistence) and Bray-curtis (stability) distances for diatom communities from Desper, Gardiner, and Scottie creeks based on three inter-annual comparisons from 2007 to 2010. Distance Measure Stream Sample Type Mean SD Jaccard Desper multi-habitat 0.66 0.02 Jaccard Desper quantitative 0.46 0.03 Jaccard Gardiner multi-habitat 0.55 0.02 Jaccard Gardiner quantitative 0.46 0.03 Jaccard Scottie multi-habitat 0.53 0.08 Jaccard Scottie quantitative 0.48 0.10 Bray-Curtis Desper multi-habitat 0.44 0.07 Bray-Curtis Desper quantitative 0.31 0.02 Bray-Curtis Gardiner multi-habitat 0.40 0.03 Bray-Curtis Gardiner quantitative 0.34 0.07 Bray-Curtis Scottie multi-habitat 0.36 0.08 Bray-Curtis Scottie quantitative 0.33 0.12

Tetlin Macroinvertebrate and Diatom Surveys

CONCLUSIONS Two sampling methodologies were used to sample the stream biological communities in this study. The quantitative samples captured almost twice the number of taxa and provided consistently higher measurements of persistence and stability for the diatoms in the three streams over time. Quantitative sampling requires a much higher level of effort processing samples in the laboratory due to the number of diatom valves identified, but may be necessary for long-term monitoring programs. For the macroinvertebrate communities, neither sampling methodology provided consistently better measurements of taxa richness, persistence, or stability. It is not clear why the quantitative macroinvertebrate samples did not do a more consistent job capturing the taxa richness, persistence, and stability of these communities since they also required a much higher laboratory effort (mean of 1500 individuals identified compared to 300 organisms in the multi-habitat fixed count).

Our data indicate that Tetlin macroinvertebrate communities have very low persistence and stability, even lower than the range observed in streams of Denali National Park (Milner et al. 2006) or alpine streams of the French Pyrenees (Brown et al. 2006). Diatom communities were more persistent and stable than the macroinvertebrate communities, but still just within the ranges of macroinvertebrate communities reported for streams of Denali National Park. This high level of inter- annual variability undoubtedly limits the ability of Tetlin stream communities to detect ecological changes over time. One potential (and admittedly speculative) reason for the lower inter-annual variability among diatoms as compared to macroinvertebrates may be their relatively rapid regeneration rates following spring ice break up, which is undoubtedly a major source of disturbance in these subarctic streams. Given these considerations, it appears that diatoms will give greater resolution for detecting ecological changes in streams in the Tetlin National Wildlife Refuge and potentially elsewhere in interior Alaska.

Tetlin Macroinvertebrate and Diatom Surveys

REFERENCES Brown, L.E., A.M. Milner, and D.M. Hannah. 2006. Stability and persistence of alpine stream macroinvertebrate communities and the role of physicochemical habitat variables. Hydrobiologia 560: 159-173.

Kaufman, P.R. and E.G. Robinson. 1998. Physical habitat assessment. Pages 77-118 in J.M. Lazorchak, D.J. Klemm, and D.V. Peck (editors). Environmental Monitoring and Assessment Program – surface waters: field operations and methods for measuring the ecological condition in wadeable streams. EPA/620/R-94/004F. US Environmental Protection Agency, Washington, DC.

Krammer, K. and H. Lange-Bertalot. 1986 – 1991. Susswasserflora von Mitteleuropa. Band 2. Parts 1-4. Bacillariophyceae. Gustav Fischer Verlag. Stuttgart. New York.

Merritt, R.W., and K.W. Cummins (editors). 1996. An introduction to the aquatic insects of North America. 3rd edition. Kendall/Hunt Publishing Company, Dubuque, Iowa.

Milner, A.M., S.C. Conn, and L.E. Brown. 2006. Persistence and stability of macroinvertebrate communities in streams of Denali National Park, Alaska: implications for biological monitoring. Freshwater Biology 51: 373–387.

Patrick, R. and C.W. Reimer. 1975. The Diatoms of the United States. Vol. 2, Part 1. Monograph No. 13. Academy of Natural Sciences, Philadelphia, Pennsylvania.

Rinella, D.J., and D.L. Bogan. 2007. Quality assurance plan for aquatic macroinvertebrate and diatom surveys in Alaska refuges. Report prepared for U.S. Fish and Wildlife Service.

Thorpe, J.H. and A.P. Covich. 2001. Ecology and classification of North American freshwater invertebrates. 2nd edition. Academic Press.

Townsend, C.R., A.G. Hildrew, and K. Schofield. 1978. Persistence of stream invertebrate communities in relation to environmental variability. Journal of Animal Ecology 56: 597–613.

Willacker, J., D. Rinella, and D. Bogan. 2008. Aquatic macroinvertebrate and diatom surveys in Tetlin National Wildlife Refuge, AK. Progress report, 2008. Report prepared for U.S. Fish and Wildlife Service.

Tetlin Macroinvertebrate and Diatom Surveys

Appendix 1. Complete macroinvertebrate taxa list, by percent composition in each multihabitat sample. *Denotes occurrence in the replicate samples but not the multihabitat sample; “0.0” denotes occurrence in the 5-minute search but not in the 300-organism subsample. Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010 Aeshna 0.0 0.0

Agabus 0.7 0.9 0.0

Agrypnia 0.4

Amphipoda 10.4 5.5 10.1 3.3 0.4 0.3 0.0 0.0

Baetis 1.8 32.9 46.1 51.6 14.7 34.7 53.4 80.3 61.0 51.9 94.0 60.5 Brachycentrus * 1.4 * *

Caenis * *

Ceraclea * * 0.3

Ceratopogonidae * 6.7 * * 0.6 0.4 * * 2.1 * 1.8

Chelifera *

Chironomidae 4.7 32.9 22.5 31.0 62.5 21.1 19.8 4.9 19.9 11.3 1.1 17.5 Cinygmula 0.3 0.7

Cladocera 55.0 0.3 0.9 * 0.4 3.5 *

Clinocera *

Coenagrionidae 0.4 2.0

Collembola 0.9 * * 0.4 *

Copepoda 10.4 * * * 0.7 *

Corixidae 1.4 0.4 0.0 0.0 2.5 0.3 0.3

Dicranota 0.3 * *

Diptera 0.3 0.6 * *

Dixella 4.3 1.1

Dolichopodidae * * * 0.3 0.0

Dytiscidae 0.4 0.6 * 0.4 0.3

Dytiscus 0.3

Empididae 0.0 1.8

Eocosmoecus 0.0 0.3

Ephemerellidae 0.7

Ephemeroptera * *

Gastropoda 2.9 1.2 1.5 2.1 0.9 0.0 0.4 * * 0.0

Glyphopsyche 0.0 *

Gyrinus *

Halesochilataylori * 0.0

Haliplidae 0.3 0.3

Hexatoma *

Hirudinea 0.4 2.9 * 0.3 0.3 * * 3.1

Hydracarina 1.4 1.2 * 2.4 * * 1.9 * * *

Hydridae * *

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010 Hydrophilidae 0.4 *

Hydroporus 0.7 0.3

Hygrobatidae 0.3 *

Lebertia * 4.5 * * 0.4 * * *

Lestes 0.0

Limnephilidae 1.1 0.6 1.5 * 1.2

Limnephilus 1.1 0.6

Limnophila 0.6

Malenka 0.3 0.0

Manayunkia * * 0.4 * * *

Metretopus * 0.4 0.8

Molannodes * *

Muscidae 0.3 * *

Nematoda * * * * *

Nematomorpha * *

Nemouridae *

Oligochaeta 1.4 2.3 * 0.3 2.1 0.0 1.6 * * 0.0 0.6 2.2 Oreodytes 0.7 0.4 * * *

Oribatei * * * *

Ostracoda * * * * * 1.9 * * * 9.6

Paraleptophlebia * 0.3 2.6 1.8 * 0.8 0.3

Plecoptera 0.3

Polycentropodidae 2.1

Procloeon 1.7 12.0 1.5 * * 0.3

Prosimulium 0.3 * 0.4 1.2 *

Psychoglypha 0.0 0.4 0.4 0.8

Serratella * 2.1 0.8 0.4 1.4 1.4 3.1 1.3

Simuliidae 0.9 7.4 0.8 6.4 *

Simulium 5.0 0.4 2.4 8.1 33.7 18.6 1.1 31.3

Siphlonurus 0.7 0.0 1.2 6.4 0.0 2.2

Sperchon *

Sphaeriidae * 1.2 * * * * * * * *

Staphylinidae 0.0 0.6 0.4

Stratiomyidae 0.3

Tabanidae 0.4 0.0 * * * * * * * *

Tipula 0.0 0.0

Tipulidae 0.3 * * 0.4 * *

Trichoclinocera 0.3 *

Trichoptera * * * 0.4

Turbellaria * 19

Tetlin Macroinvertebrate and Diatom Surveys

Appendix 2. Complete diatom taxa list, by percent composition in each multihabitat sample. *Denotes occurrence in the replicate samples but not the multihabitat sample.

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010 AULACOSEIRALES Melosira varians Agardh 2.0 1.3 * 2.0 16.0 11.3 3.2 11.2 2.3 8.8 1.2 1.3

Aulacoseira ambigua (Grunow) 0.3 6.2 4.8 2.8 0.3 1.7 * 1.5 2.2 1.7 * 2.2 Simonsen

Aulacoseira italica (Ehrenberg) 3.3 1.0 * * * Simonsen

Aulacoseira subarctica (Müller) 0.3 2.0 * 1.0 * Haworth

Aulacoseira valida (Grunow) 0.3 * * 0.4 * Krammer THALASSIOSIRALES Cyclotella bodanica fo. lemanica * (Muller in Schroter; Muller in Chodat) Bachmann Cyclotella meneghiniana Kützing 0.3 * * 1.0 * * 0.2 0.5 0.7 0.4 0.3

* Cyclotella michiganiana Skvortzow 0.3 * * Cyclotella ocellata Pantosek Discostella pseudostelligera * (Hustedt) Houk et Klee

Discostella stelligera (Hustedt) Houk * 0.2 * * et Klee TABELLARIALES

15.5 10.7 5.7 18.2 2.3 11.1 9.7 13.2 5.0 4.2 6.5 5.3 Tabellaria flocculosa (Roth) Kützing

* * 0.3 * * Tetracyclus glans (Ehrenberg) Mills FRAGILARIALES * * Asterionella formosa Hassal Diatoma mesodon (Ehrenberg) * * * * Kützing * * 0.7 0.2 * 1.1 Diatoma moniliformis Kützing 0.7 0.3 * 0.2 * 0.3 0.6 0.7 * 1.5 Diatoma tenuis Agardh * * * 0.3 0.3 2.0 3.5 4.1 0.4 1.5 1.2 4.3 Meridion circulare (Greville) Agardh

Fragilaria capucina var. rumpens 5.7 1.8 (Kützing) Lange-Bertalot Fragilaria capucina Desmazières 11.7 2.8 5.5 14.2 1.3 2.7 3.5 4.4 6.0 11.1 3.4 3.3

Fragilaria neoproducta Lange- * Bertalot 20

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010

Fragilaria vaucheriae (Kützing) * Petersen

Fragilariforma bicapitata (Mayer) * 0.3 0.3 * * * 0.2 Williams et Round

* 0.3 * * * * Hannaea arcus (Ehrenberg) Patrick

Pseudostaurosira brevistriata * * * (Grunow) Williams et Round

Pseudostaurosira parasitica (Smith) * 0.3 * 0.3 * * * Morales

Pseudostaurosira parasitica var. 0.3 * subconstricta (Grunow) Morales

Stauroforma exiguiformis (Lange- * * Bertalot) Flower, Jones et Round

Staurosira construens var. binodis 1.2 (Ehrenberg) Hamilton Staurosira construens Ehrenberg * 14.8 8.8 8.2 0.2 * 0.7 9.3 8.2 12.1 18.8

Staurosira construens var. venter * (Ehrenberg) Hamilton

Staurosirella leptostauron * * * * * * * (Ehrenberg) Williams et Round

Staurosirella pinnata (Ehrenberg) 2.5 5.2 3.8 * 0.3 0.3 * 0.5 14.5 11.5 12.6 7.7 Williams et Round Synedra ulna (Nitzsch) Ehrenberg 4.7 1.3 3.0 5.3 2.0 4.3 5.0 7.2 4.5 5.3 3.1 2.0 EUNOTIALES 0.5 Eunotia alpina Kützing 6.8 3.3 3.2 1.3 0.7 0.3 2.8 2.6 2.6 2.3 1.0 2.8 Eunotia bilunaris (Ehrenberg) Mills

Eunotia bilunaris var. mucophila * 0.3 * * * 3.0 0.3 0.4 0.3 * Lange-Bertalot et Nörpel 0.3 Eunotia crista-galli Cleve 0.2 * Eunotia diodon Ehrenberg Eunotia exigua (Brébisson ex * 0.2 0.8 Kützing) Rabenhorst

Eunotia implicata Nörpel, Lange- 1.2 Bertalot et Alles Eunotia incisa Smith ex Gregory 16.2 3.3 5.2 1.5 * 1.3 2.7 2.5 0.2 * 1.5 * * * * 0.7 1.5 0.3 0.3 0.7 * Eunotia minor (Kützing) Grunow * * Eunotia monodon Ehrenberg Eunotia muscicola var. tridentula 0.3 Nörpel et Lange-Bertalot * Eunotia paludosa Grunow Eunotia pectinalis (Müller) 1.3 * 0.2 Rabenhorst Eunotia praerupta Ehrenberg 0.2 1.7 1.5 2.0 * 0.7 1.5 2.2 0.4 1.8 1.2 1.5 21

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010 * 0.3 Eunotia serra Ehrenberg Eunotia soleirolii (Kützing) * * 0.3 * 0.2 * Rabenhorst ACHNANTHALES

Achnanthes biasolettiana (Kützing) 6.3 3.3 2.0 1.0 * * 1.4 0.8 0.7 * * Grunow * Achnanthes curtissima Carter Achnanthes impexiformis Lange- * Bertalot 0.7 * * 0.2 * * Achnanthes nodosa Cleve Achnanthidium minutissimum 2.3 2.5 9.0 5.8 8.2 8.3 9.2 8.8 5.8 3.2 1.5 2.2 (Kützing) Czarnecki Cocconeis placentula Ehrenberg 17.0 11.2 9.8 11.2 0.7 2.3 6.0 4.2 5.6 6.8 7.4 14.0

Eucocconeis alpestris (Brun) Lange- * * Bertalot

* 0.8 * * Eucocconeis flexella (Kützing) Cleve

Eucocconeis laevis (Østrup) Lange- * 0.7 0.3 * * 1.3 2.2 0.5 0.4 0.7 * 0.5 Bertalot

Karayevia laterostrata (Hantzsch) * Bukhtiyarova

Planothidium frequentissimum * * 2.0 * * 0.3 * 2.3 (Lange-Bertalot) Lange-Bertalot

Planothidium haynaldii * * * 4.5 5.0 6.2 0.4 0.4 * * (Schaarschmidt) Lange-Bertalot

* * * * * 1.6 0.7 1.0 1.0 2.5 Planothidium lanceolatum (Brébisson ex Kützing) Lange-Bertalot

Planothidium oestrupii (Cleve-Euler) * * Round et Bukhtiyarova

Psammothidium bioretii (Germain) * * * * Bukhtiyarova et Round

Psammothidium rossii (Hustedt) 0.3 Bukhtiyarova et Round

Psammothidium subatomoides 0.3 * * * * * * 0.2 * * (Hustedt) Bukhtiyarova et Round

Rossithidium pusillum (Grunow) 3.3 2.0 * * * 0.8 0.5 Round et Bukhtiyarova NAVICULALES * * * Navicula constans Hustedt 0.7 0.3 2.2 4.5 0.7 * 3.4 1.2 1.7 0.6 11.2 Navicula cryptocephala Kützing Navicula cryptotenella Lange- 1.7 3.0 7.7 11.2 9.5 7.2 0.8 10.6 6.3 11.6 Bertalot * 0.3 * 10.5 3.2 3.8 0.3 0.3 0.4 0.5 Navicula gregaria Donkin

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010

Navicula lanceolata (Agardh) 0.3 1.3 1.0 * * * 0.4 0.7 * Ehrenberg * Navicula laterostrata Hustedt 0.3 * 0.3 0.3 Navicula menisculus Schumann * * Navicula pseudosilicula Hustedt 0.7 0.3 0.9 * Navicula radiosa Kützing Navicula rhynchocephala Kützing 0.3 1.0 1.0 0.5 0.7 3.7 1.5 2.5 0.6 0.3 0.2 * * * Navicula spp. 0.8 * Navicula tridentula Krasske Navicula viridula (Kützing) Kützing * emend. Van Heurck

Aneumastus tusculus (Ehrenberg) * Mann et Stickle

Cavinula cocconeiformis (Gregory ex * * Greville) Mann et Stickle

Cavinula lapidosa (Krasske) Lange- * * * Bertalot

0.3 Cavinula pseudoscutiformis (Grunow ex Schmidt) Mann et Stickle

* * 1.7 * * * Chamaepinnularia soehrensis (Krasske) Lange-Bertalot et Krammer

* * * * Craticula cuspidata (Kützing) Mann

Diadesmis contenta (Grunow ex Van 0.3 * 0.3 * 0.3 0.7 5.5 1.1 0.2 0.3 0.4 0.5 Heurck) Mann

Fallacia pygmaea (Kützing) Stickle et * * * * Mann

Geissleria ignota (Krasske) Lange- * Bertalot et Metzeltin Hippodonta capitata (Ehrenberg) Lange-Bertalot, Metzeltin et * 1.5 4.3 1.8 4.2 6.5 0.3 0.4 1.4 2.0 2.2 2.2 Witkowski Hippodonta hungarica (Grunow) * * Lange-Bertalot, Metzeltin et Witkowski * * Luticola cohnii (Hilse) Mann Luticola goeppertiana (Bleisch) * 0.3 * 0.3 * * * 0.3 0.2 * 0.2 * Mann * * * * * Luticola mutica (Kützing) Mann Microcostatus krasskei (Hustedt) * * * * 1.5 0.4 * Johansen et Sray Parlibellus crucicula (Smith) * * Witkowski, Lange-Bertalot et Metzeltin 23

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010

* 0.3 * * * 0.4 0.5 0.2 Placoneis elginensis (Gregory) Cox

Placoneis placentula (Ehrenberg) * * Hienzerling

Sellaphora bacillum (Ehrenberg) * Mann

Sellaphora laevissima (Kützing) * 0.3 * 0.2 Mann

Sellaphora pupula (Kützing) 1.0 * 0.5 * 1.7 2.2 1.3 0.2 1.7 0.6 0.3 Meresckowsky

Sellaphora seminulum (Grunow) * * 0.7 * Mann

Amphipleura pellucida (Kützing) * * * 0.2 1.0 1.3 * Kützing

Anomoeoneis sphaerophora * (Kützing) Pfitzer 0.5 * Brachysira brebissonii Ross * Caloneis bacillum (Grunow) Cleve

Caloneis schumanniana (Grunow) * 0.8 * * * Cleve

* 1.0 1.8 0.8 * * * 0.5 0.4 0.3 0.2 0.7 Caloneis silicula (Ehrenberg) Cleve * * Diploneis boldtiana Cleve * 0.7 0.3 0.7 0.3 1.0 * 0.8 0.3 * 0.7 Diploneis elliptica (Kützing) Cleve

* * Diploneis finnica (Ehrenberg) Cleve

Diploneis oblongella (Naegeli ex * * * Kützing) Ross * 0.3 Diploneis peterseni Hustedt Frustulia krammeri Lange-Bertalot * 0.5 * * 0.4 * * et Metzeltin

* * 2.8 * * * * * Frustulia vulgaris (Thwaites) De Toni

Gyrosigma acuminatum (Kützing) * 0.1 * 0.2 * Rabenhorst

* * Neidium affine (Ehrenberg) Pfitzer

Neidium ampliatum (Ehrenberg) 1.0 0.5 1.0 * Krammer

Neidium binodis (Ehrenberg) * Hustedt

Neidium bisulcatum (Lagerstedt) * * * 0.3 1.0 * 0.4 0.4 0.3 * * Cleve Pinnularia borealis Ehrenberg * * * 0.7 0.3 * 0.7 0.2 0.6 0.3 0.4 0.3

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010 * Pinnularia brandelii Cleve Pinnularia divergentissima (Grunow) * Cleve * * * * 0.5 Pinnularia gibba Ehrenberg Pinnularia hemiptera (Kützing) * * Rabenhorst

Pinnularia mesolepta (Ehrenberg) * * * * * * 0.2 * 0.8 * Smith

Pinnularia microstauron (Ehrenberg) * 0.5 * 1.0 * * 0.3 0.6 0.5 0.6 * Cleve

Pinnularia nodosa (Ehrenberg) 0.7 0.3 * * 0.8 0.3 Smith * Pinnularia obscura Krasske * 0.2 Pinnularia spp. * 0.2 Pinnularia streptoraphe Cleve Pinnularia subcapitata Gregory 0.3 * 0.3 2.0 * 0.4 0.7 0.2 0.3

Pinnularia viridis (Nitzsch) * * * Ehrenberg

Pleurosigma angulatum (Quekett) 0.3 0.3 Smith Stauroneis anceps Ehrenberg * 0.7 * * 0.7 0.3 0.3 0.3 0.7 * 0.8 0.7

Stauroneis phoenicenteron (Nitzsch) 0.3 0.3 * * * * 0.3 0.7 0.2 * Ehrenberg * 0.3 0.7 * * * * * * 0.3 Stauroneis smithii Grunow THALASSIOPHYSALES

Amphora copulata (Kützing) * 0.3 1.0 0.3 * * 0.4 0.8 1.3 * 3.0 Schoeman et Archibald * 0.3 1.0 * * 0.7 0.5 0.2 1.0 0.3 * Amphora inariensis Krammer * Amphora ovalis (Kützing) Kützing Amphora pediculus (Kützing) 0.3 0.3 * * * 1.0 * * * 0.4 Grunow CYMBELLALES * * 0.3 * * Cymbella caespitosa Brun Cymbella cistula (Ehrenberg) * * * * 0.3 Kirchner

Cymbella gracilis (Ehrenberg) * 0.3 * * * * * * * * * Kützing

Cymbella naviculiformis Auerswald 0.3 0.3 0.7 * * * 0.7 0.6 0.3 0.4 0.3 ex Héribaud * Cymbella obscura Krasske * * 0.2 * Cymbella subaequalis Grunow * * * Cymbella subcuspidata Krammer * Cymbella tynii Krammer

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010

Encyonema gaeumanii (Meister) 0.2 Krammer

* * * 1.7 * 2.3 3.2 0.8 1.0 0.7 Encyonema minutum (Hilse) Mann

Encyonema reichardtii (Krammer) 0.3 Mann

Encyonema silesiacum (Bleisch) * 0.7 2.0 1.0 0.7 1.7 * 4.8 3.3 0.3 1.9 1.8 Mann

Reimeria sinuata (Gregory) Kociolek * * 1.0 0.9 0.2 * * et Stoermer

Gomphonema acuminatum * * 0.3 0.5 0.2 0.2 * Ehrenberg

* * 1.0 * * * 0.3 Gomphonema brebissonii Kützing

0.3 * * Gomphonema clavatum Ehrenberg

Gomphonema gracile Ehrenberg 0.7 * emend Van Heurck * 0.2 Gomphonema insigne Gregory 0.3 Gomphonema lagerheimii Cleve 0.3 * * 0.7 0.4 * Gomphonema micropus Kützing Gomphonema minutum (Agardh) * Agardh

Gomphonema olivaceum (Lyngbye) 0.3 * * 0.3 Kützing

* Gomphonema olivaceoides Hustedt

Gomphonema parvulum (Kützing) * * * * 1.7 2.0 1.0 * 1.8 0.7 0.6 1.0 Kützing

0.3 1.3 * 1.5 * 0.3 0.7 * 0.5 * Gomphonema sarcophagus Gregory * * * Gomphonema spp. 0.3 * 0.2 0.3 Gomphonema truncatum Ehrenberg

Rhoicosphenia abbreviata (Agardh) * * * * * Lange-Bertalot BACILLARIALES * * * * * Denticula kuetzingii Grunow Hantzschia amphioxys (Ehrenberg) * * * 0.3 1.0 * * * 0.6 0.7 1.6 1.5 Grunow

4.2 0.3 Nitzschia acicularis (Kützing) Smith * 0.3 * 1.0 0.7 0.6 * Nitzschia amphibia Grunow 0.3 * * Nitzschia archibaldii Lange-Bertalot

Tetlin Macroinvertebrate and Diatom Surveys

Desper Creek Gardiner Creek Scottie Creek Taxon 2007 2008 2009 2010 2007 2008 2009 2010 2007 2008 2009 2010

* 0.7 * * 2.0 2.0 * 1.7 0.6 0.5 0.2 * Nitzschia dissipata (Kützing) Grunow

Nitzschia filiformis (Smith) Van 0.3 0.3 * 0.3 0.7 * * * Heurck

Nitzschia frustulum (Kützing) 0.4 * Grunow

Nitzschia linearis (Agardh ex Smith) 1.8 * * * * 0.3 * 0.9 * * 0.2 * Smith Nitzschia palea (Kützing) Smith 2.2 4.0 2.0 * 7.7 3.3 4.2 1.6 1.9 1.3 7.5 1.0 0.3 Nitzschia pellucida Grunow Nitzschia perminuta (Grunow) * 0.3 * * 0.3 * 2.3 0.4 0.4 0.7 0.4 * Peragallo

Nitzschia recta Hantzsch ex 0.3 1.0 0.7 0.2 0.3 * * Rabenhorst 0.3 Nitzschia spp. Nitzschia tubicola Grunow in Cleve * 0.7 * * * * * 0.3 et Grunow RHOPALODIALES

Epithemia adnata (Kützing) 0.8 * 0.3 0.3 * * 0.6 0.4 * 0.7 0.8 Brébisson * Epithemia spp. Rhopalodia gibba (Ehrenberg) * 0.3 2.2 * * * Müller * * Rhopalodia spp. SURIRELLALES

Cymatopleura solea (Brébisson) * * * 0.6 0.2 Smith * * Surirella amphioxys Smith Surirella angusta Kützing 0.7 * * 0.3 0.7 * * 0.4 * 0.4 *

Surirella brebissonii Krammer et * * 0.3 * * * Lange-Bertalot * * Surirella elegans Ehrenberg * * Surirella linearis Smith 1.0 * * Surirella minuta Brébisson * Surirella robusta Ehrenberg Surirella spiralis Kützing *