Completed Research Projects


A Landscape Approach to Modeling Habitat Use and Monitoring Population Trends of the American Marten

Funding Source:
U.S.D.A. Forest Service
Utah Division of Wildlife Resources
Utah Wilderness Association

Project Location: Uinta Mountains of northeastern Utah

Project Duration: September 1990 - December 1994

Investigators:
Christina Hargis
John A. Bissonette

Technicians:
Peter Lortz
Rob Rood
The Ph.D. thesis was written as a series of three papers. The abstract for each are given below.

(1) THE BEHAVIOR OF LANDSCAPE METRICS COMMONLY USED IN THE STUDY OF HABITAT FRAGMENTATION

Abstract:

Landscape metrics have been developed for quantifying various aspects of landscape pattern, and several of these have been applied to studies of habitat fragmentation. A meaningful interpretation of landscape metrics is only possible when the limitations of each measure are fully understood, the range of attainable values is known, and the user is aware of potential shifts in the range of values due to characteristics of landscape patches. To examine the behavior of landscape metrics, I generated artificial landscapes that mimicked fragmentation processes while controlling the size, shape, and placement of disturbance patches. I created nine series of increasingly fragmented landscapes that differed from one another in the size or shape of patch used to represent disturbance and in the way disturbance was allowed to increase, and used these landscapes to investigate edge density, contagion, mean nearest neighbor distance, mean proximity index, perimeter-area fractal dimension, and mass fractal dimension. Most of the metrics provided unique information about an aspect of landscape pattern, but contagion and edge density were highly correlated, as were mass fractal dimension and the amount of disturbance. Correlations among metrics increased when disturbance growth was through buffered patches, because buffers restricted the proportion of disturbance on the landcape to less than 0.40, and many of the metrics were linearly-associated with increasing disturbance up to 0.40. None of the measures were able to distinguish landscape patterns of dispersed versus aggregated patches. The attainable values of all measures were altered by either patch size or shape, and in some cases, by both attributes. I summarize the utility of each metric in providing information about habitat fragmentation.

(2) THE INFLUENCE OF FOREST FRAGMENTATIONON AMERICAN MARTEN POPULATIONS

Abstract:

I investigated the effects of forest fragmentation on American marten (Martes americana) by evaluating differences in marten densities across 18 study sites that varied in landscape heterogeneity due to the combined effects of natural openings and timber harvest clearcuts. The 9-km2 sites were located in mature forests in the Uinta Mountains of northern Utah, and open areas occupied 2-42% of each site. I proposed three possible responses of marten populations to increases in fragmentation: 1) a linear negative decline based on habitat loss; 2) an initial positive response related to the added diversity and abundance of small mammals associated with clearcuts and meadows; and 3) an accelerated decline due to the combined effects of landscape pattern and habitat loss. I used ERDAS, a raster-based GIS, to classify the landscape images into 2-attribute maps representing forest and openings, and quantified fragmentation using five measures: the percent of each site in open areas, edge density, mean proximity index for open patches, mean nearest neighbor distance between open patches, and mass fractal dimension of the forest matrix. Within each site, I estimated marten abundance through live-trapping, estimated small mammal densities through snap-trapping, and collected data on forest structure. I found a significant negative correlation of marten captures to the combined effects of loss of forest habitat, as measured by the percent of each site in open areas and an increase in the proportional representation of snags in the overstory (P less than 0.05). The graphed relationship of marten captures and percent of site in openings suggested a possible threshold when open areas exceeded 20% of the landscape. Absence of marten occurred earlier than predicted by direct loss of habitat, implying additive effects of landscape pattern. Mean proximity index, which quantifies patch isolation based on the size and distance between open areas, was the strongest correlate with marten captures. Marten were not found in landscapes where patches were large and closely situated, and the combined effects of patch proximity and snag abundance explained more variation in marten captures than any other model examined (R2 = .59). I found no evidence of a positive response to low levels of fragmentation. Small mammal densities were significantly higher in clearcuts than in forests (P less than 0.01), but marten captures were not correlated with the increase in prey abundance or biomass associated with clearcuts. Conservation efforts for the marten must consider not only the structural aspects of mature forests, but the landscape pattern in which the forest occurs. I recommend that timber harvest blocks be aggregated to reduce forest edge and maintain maximum area in forest interior.

(3) LANDSCAPE -LEVEL EFFECTS OF TIMBER HARVESTS ON SMALL MAMMAL POPULATIONS IN SPRUCE-FIR FORESTS

Abstract:

Small mammal communities are affected by changes in vegetation and habitat structure resulting from clearcut timber harvests, but the effects of multiple clearcut blocks at the landscape scale have not been studied. I conducted a small mammal survey within a series of 9 km2 landscapes containing varying proportions of forest and open areas to examine effects of landscape pattern on small mammal populations. The broad scale of this study also provided a large and geographically-disjunct set of clearcut blocks and forest stands for comparing small mammal communities between these habitats. I surveyed small mammal populations over a 7-week period between 22 July and 8 Sept, 1992, using 1,200 trap nights per study site, and captured 14 species of small mammals. The five most commonly-trapped species were the southern red-backed vole (Clethrionomys gapperi), deer mouse (Peromyscus maniculatus), Uinta chipmunk (Eutamias umbrinus), least chipmunk (Eutamias minimus), and shrew (Sorex spp). Capture rates of southern red-backed vole, the most frequently-captured species in mature forests, declined with loss of forest habitat, but the decline was greater than predicted from a null model of capture rates based on habitat loss alone. Landcape pattern appeared to serve as an additive effect contributing to lower captures of red-backed voles in landscapes with high edge density, little forest interior, and large, open areas in close proximity. In contrast, these landscape patterns were associated with increased capture rates of deer mice, which occurred in greatest numbers in clearcuts and in landscapes with high juxtaposition of open areas with forests. The Uinta chipmunk was the only other species captured in sufficient numbers to analyze statistically, and I found no change in capture rates with fragmentation. At the scale of individual cut blocks and habitat comparisons, I found greater species richness in mature forests, but higher overall capture rates in clearcut blocks. I concluded that land managers must be aware not only of the effects of individual cut blocks, but of the larger scale effects of cut blocks on landscape pattern and small mammals, as well as the potential influences on larger species for whom the small mammals serve as prey. I recommend aggregated rather than dispersed patterns of clearcuts so that a greater proportion of the landscape can be maintained in forest interior habitat for species that are sensitive to habitat area and edge.


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Mule Deer Highway Mortality in Northeastern Utah: An Analysis of Population-Level Impacts and a New Mitigative System


Funding Source:
U.S.Bureau of Reclamation
Utah Department of Transportation
Utah Division of Wildlife Resources

Project Location: Jordanelle Reservoir area, Utah

Project Duration: September 1993 - September 1996

Investigators:
John A. Bissonette
Mark Lehnert

Technicians: Christine Vogel
Gabe Bissonette
Karen Griffiths

Rerouting highways to accommodate construction of the Jordanelle Reservoir in northeastern Utah caused a dramatic increase in vehicle collisions with mule deer (Odocoileus hemionus). I evaluated the effectiveness of a new system of highway crosswalk structures installed to reduce deer losses and preserve seasonal migrations. In addition, I constructed computer simulation models to investigate how highway mortality has impacted the Jordanelle deer population.
The crosswalk system restricted deer crossings to specific, well-marked areas along highways where motorists could anticipate them. Subsequent to installation, mortality declined 42.3% and 36.8% along a four-lane and two-lane highway, respectively. I was unable to statistically demonstrate that observed mortality reductions were a direct result of the crosswalk system. The potential applicability of the structures, however, should not be dismissed. Reduced deer use of the highway right-of-way (ROW), the apparent maintenance of migratory behavior, and observations of animals crossing within crosswalk boundaries indicate the system warrants further testing. Lack of motorist response to crosswalk warning signs, the tendency for foraging deer to wander outside crosswalk boundaries, and the ineffectiveness of ROW escape gates contributed to most treatment area mortalities. I offer design modifications that address these shortcomings.

Four years of field data revealed that highway mortality at Jordanelle was inversely density-dependent, removed between 5.6% and 17.4% of the population each year, and disproportionately impacted bucks. I incorporated this information into 3 competing simulation models in which highway losses operated in a strictly additive, partially compensatory, or strictly compensatory manner. The partial compensation model most closely tracked observed population dynamics, suggesting that highway losses were not completely offset by reductions in other mortality sources. Highway mortality apparently worsened a population crash initiated by severe winter conditions, and may be slowing the recovery. The disproportionate loss of bucks along roads altered sex ratios of simulated populations. Mitigative efforts should target road-kill reductions >60% to avoid population declines predicted by the partial compensation model. Annual variation in demographic parameters offset the impacts of highway mortality at high population levels. At low population levels, however, highway mortality was severe enough to drive declining population trends.


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Second Growth Forest as Potential Marten Habitat in Western Newfoundland: An Examination of Forest Habitat Structure and Microtine Abundance

Funding Source:
Western Newfoundland Model Forest
Newfoundland and Labrador Wildlife Division

Project Location: Western Newfoundland Model Forest

Project Duration: March 1993 - August 1995

Investigators:
Brian Sturtevant
John A. Bissonette

Technicians:
Keith Chaulk
Tina Newbury
William Dennis

The American marten (Martes americana) is associated with large tracts of relatively undisturbed, mature coniferous forests. I examined coarse woody debris (CWD) structure and small mammal abundance with respect to forest age and stem structure within second-growth forests, in comparison with old-growth stands in western Newfoundland. Results suggest that a critical change in marten habitat quality may occur at stand senescence, due to decreased tree competition, more complex subcanopy structure, and increased meadow vole (Microtus pennsylvanicus) abundance.
Analysis of stem structure within a chronosequence of 19 second-growth stands indicated high intertree competition, with dense canopy closure and active self-thinning, until stand senescence at 80 years. Old-growth stands were less dense, offering more canopy openings. CWD volume observed within the chronosequence demonstrated the typical U-shaped temporal relationship observed in other forested systems. Lowest CWD volumes were observed within semimature to mature second growth. Highest levels of both CWD volume and structural complexity corresponded with stand senescence and old growth.
Small mammals were sampled within immature, semimature, mature, and silviculturally overmature coniferous stands in western Newfoundland during 1993 and 1994. Meadow voles were most abundant within overmature stands (P less than 0.05). Vole abundance was negatively correlated with relative density (P less than 0.01) and positively correlated with CWD (P less than 0.01). I address the applicability of mainland marten habitat generalizations to the unique depauperate condition of Newfoundland.
Results suggest that the critical elements of marten foraging habitat currently are found within a senescent forest stand structure. Further review of Newfoundland harvest records indicated that anthropogenic disturbance transformed a historically heterogeneous forested landscape into a more contiguous, even-aged, second-growth environment. Using the stand density management diagram, I designed a silvicultural approach to marten habitat management that simulates the structure of older forests within younger stages of forest development.


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Biodiversity and Landscape Planning: Alternative Futures for the Region of Camp Pendleton, California


Funding Source:
DOD Strategic Environmental Research Development Plan
Department of Energy
Environmental Protection Agency

Project Location: Southern California

Project Duration: January 1995-September 1996

Investigators:
Carl Steinitz, Harvard University
Craig Johnson, USU
Michael Binford, Harvard University
Ross Keister, USDA FS
Paul Cote, Harvard University
David Mouat, EPA
Thomas C. Edwards, Jr., NBS, USU
Douglas Olsen, Harvard University
Stephen Irvin, Harvard University
Allan Shearer, Harvard University
Richard T. Forman, Harvard University
Richard Toth, USU
Robin Wills, Nature Conservancy
Scott Bassett, Utah State University

This study explores how urban growth and change in the rapidly developing area located between San Diego and Los Angeles might influence regional biodiversity. The study region is 90 km X 120 km rectangle that encompasses 5 major river drainage basins directly influencing Camp Pendleton. The research strategy is based on the hypothesis that the major stressors impacting biodiversity are related to increasing urbanization. As population increases and development spreads, habitat is lost due to grading, paving, ornamental landscaping, and other human activities. There are also indirect effects of development, such as changes in hydrologic and fire regimes which impact vegetation and, ultimately, biodiversity.

A computer-based GIS was developed for the region. Analytical models use the data in the GIS to evaluate the complex dynamic processes at work in the study area and to provide a medium for modeling different future scenarios. Regional change is simulated via six alternative projections of development to the year 2010 and to subsequent "build-out." Biodiversity is assessed in three ways: a landscape ecological model, ten selected species models, and a species richness model. The alternative scenarios may be used by regional stakeholders to assess the desirability of various policies affecting future biodiversity to the area.

The project received the American Planning Associations 1996 Outstanding Achievement Award for Unique Contribution. A book contract is currently being negotiated, and various manuscripts are in progress.


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What Triggers Outbreaks of Avian Botulism?


Funding Source:
Utah Division of Wildlife Resources
U.S. Fish and Wildlife Service

Project Location: Great Salt Lake Wetlands

Project Duration: June 1994 - June 1996

Investigator: John A. Kadlec

Technician: Karen Lindsey

Goals and Objectives:
In the past, outbreaks of avian botulism were often observed to be associated with re-flooding of mud-flats exposed by receding water levels in late summer. This project will evaluate the environmental consequence of this phenomenon in a 2-year controlled switch-back experiment using two impoundments at the Ogden Bay Waterfowl Management Area. The first year's data, on preliminary examination, show a large increase in sulphur as the reflooding occurs. Presumably, this reflects an abrupt change in microbiological activity in the sediment, which might favor enhanced growth of the botulism organism, (Clostridium botulinum).

The second year's data confirmed an increase in soluable sulphur, most likely in the form of sulfate, at reflooding of the impoundment. In addition, pH decreases as water levels were lowered. The decreased pH most likely reflects the observed changes in sulfate concentration, mediated by the presumed changes in microbial activity. Although the micro-organisms reliant on sulphate are negatively affected by the drawdowns, the relationship between sulphur bacteria and C. botulinum is poorly understood.


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Spawning and Recruitment of Salmonids in Strawberry Reservoir


Funding Source:
Utah Division of Wildlife Resources
Utah Reclamation, Conservation, and Mitigation Commission

Project Location: Strawberry Valley, Utah

Project Duration: March 1994 - January 1997

Investigators:
Curtis Knight
Rick Orme
David A. Beauchamp

Technicians:
Jane Chen
Jeffrey Godfrey
Corey Hendricks
Janene Graff
Fredrik Norrsell
Eric Asarian

Goals and Objectives:
Our objective was to estimate the potential for Bear Lake cutthroat trout and kokanee salmon to support self-sustaining, naturally reproducing populations in Strawberry Reservoir and its tributaries.

We estimated the timing, distribution, and abundance of spawning cutthroat trout and resultant production of fry in tributaries to Strawberry Reservoir in 1994 and 1995. Spawners first entered streams during the first week of May in 1994, but the run did not accelerate until around 20 May. In 1995, the spawning run accelerated around the first week in June. Earlier detection of spawners was hampered by high and turbid stream flows. The beginning of the spawning run in both years corresponded to decreasing stream discharges, mean daily stream temperatures of around 10øC, and approximately 45 days after ice off. The majority of the spawning activity took place in Indian Creek, Trout Creek, and Strawberry River during both years. In 1995, increased stream flows allowed spawners to enter numerous smaller tributary streams. An increased use of higher stream reaches was noted in 1995, likely a response to higher stream flows. Indian Creek was the most productive stream with an estimated 2,808 spawners in 1995 and 3,444 spawners in 1994. Spawners in the Strawberry River were not estimated in both years due to prolonged high flows and turbid water; however, we were able to count redds in 1994 (366). The number of spawners in Trout Creek were estimated at 605 for 1995 and 388 for 1994. Sex ratios varied between years in Indian Creek and Trout Creek with females representing 61% of the spawners in 1994 and only 22% in 1995. In addition, annual trend gill net data, peak spawners, sex ratios, and estimated spawner abundance were examined for usefulness as future predictive tools for fry production.

Indian was the most productive stream in both years, producing an estimated 561,047 fry in 1994 and 436,264 fry in 1995. Trout Creek produced 157,896 fry in 1994 and 169,632 fry in 1995. We estimated 108,536 fry produced in the Strawberry River in 1994 and because high and turbid stream flows were present for the majority of the spawning run in 1995, we were unable to estimate fry production. These estimates should be considered a minimum for the whole Strawberry Basin, as smaller streams, especially during years of high runoff such as 1995, provided additional spawning habitat for cutthroat trout. Cutthroat trout fry undertook two different life history strategies; fry either migrated to the reservoir within days of emerging from the gravel or fry remained in the stream for 1-2 years. In 1994, an estimated 21% of fry produced in Indian Creek emigrated to the reservoir and 25% emigrated in Trout Creek. In 1995, emigration rates were lower (3.5% for Indian Creek and 11.4% in Trout Creek), presumably because higher stream flows provided more rearing habitat for fry. In 1995, mean lengths and weights of fry sampled from outmigrant fry traps were consistent (26mm and 0.10 g) throughout the fry outmigration from 8 August to 3 September, suggesting that fry spent little time in the stream before emigrating. Outmigrant fry traps were fished until 19 October 1995; however, very few fry migrated from 3 September to 19 October.

Population estimates of young-of-year cutthroat trout remaining in the stream indicated that mortality rates were extremely high (99.8% in 1994, 98% in 1995) during their first growing season (August-November). Thereafter, mortality rates were low, even during winter, and most losses from the streams were attributed to outmigration to the reservoir. Juveniles remained in the streams for 1-2 years with few fish remaining past three years.

Kokanee spawning and recruitment patterns were examined starting in fall 1994 through winter 1996 to determine if kokanees could become self-sustaining in the Strawberry Basin. We documented run timing and distribution of tributary spawners, and estimated spawner abundance, fecundity rates, total egg deposition, and egg-to-fry survival rates in the major spawning tributaries to Strawberry Reservoir. Spawners were more abundant and more broadly distributed in 1995 than in 1994. In 1995, we estimated that 17,838 kokanees (8,889 females and 8,949 males) spawned in Indian Creek, 10,308 (4,741 females and 5,567 males) in Trout Creek, and 2,995 (1,529 females and 1,467 males) spawned in Strawberry River. Two size modes of both sexes spawned in 1994, but only the larger mode was evident in 1995. Fecundity averaged 1,400 eggs/female and mean egg retention was 294 (SD = 419) eggs/female. We estimated that 9,859,175 kokanee eggs were deposited in Indian Creek, plus 5,366,436 eggs in Trout Creek, and 1,725,485 eggs in Strawberry River; this represented a nearly 5-fold increase over the estimated egg deposition in 1994. Our current estimates of egg-to-fry survival are 6.5% in 1994 and 8.0% in 1995. These survival rates would translate into 1,356,000 emergent fry among the three major tributaries in spring 1996, compared to 144,000 emergent fry in spring 1995. The implications of these recruitment rates for kokanee management in Strawberry Reservoir will be evaluated.


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Carrying Capacity of Strawberry Reservoir Salmonids

Funding Source:
Utah Division of Wildlife Resources
Central Utah Project

Project Location: Strawberry Valley, Utah

Project Duration: March 1994 - January 1996

Investigators:
James R. Ruzycki
Casey Baldwin
David A. Beauchamp

Technicians:
Jeff Godfrey
Jason Van Tassell
Vance Mumford

Goals and Objectives:
The primary management goal of Strawberry Reservoir was to provide a quality salmonid fishery sustained by the natural reproduction of cutthroat trout and kokanee salmon. To provide a self-sustaining fishery in a reservoir, balance must be maintained among the various biotic and abiotic components of the aquatic environment. To achieve a balance and to provide maximum fishery potential, fishery stocks should be maintained near but also below carrying capacity. Estimating the carrying capacity is therefore an important objective of the current management plan. During 1994, we initiated a cooperative effort with the Utah Division of Wildlife Resources (UDWR) to determine the carrying capacity of Strawberry Reservoir. Work completed during this initial year indicated that the reservoir's fishery was significantly below capacity. In 1995 we refined our carrying capacity estimates and provided initial information for examination of the food web processes which may be limiting recruitment. These refinements include more accurate, quantified diet histories and more representative sampling of the temporal and spatial distribution of salmonids. The consumption demand of the salmonid community was estimated using a bioenergetics modeling approach. Temporal and spatial distributions of salmonids were estimated with a combination of netting and hydroacoustic techniques.


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Great Basin Gap Analysis

Funding Source:
U.S. Fish and Wildlife Service

Project Location: Logan, Utah

Project Duration: October 1990 - September 1996

Investigators:
Thomas C. Edwards, Jr.
Allan Falconer
Collin Homer
Scott Bassett

Technicians:
Janene Shupe
Mary Jane Crandall
Mathew McCune

Progress to Date:
All data for the Utah component of the Great Basin Gap analysis have been synthesized and are available in digital format for use by management agencies. Several manuscripts have been submitted. All objectives except for the final report have been met.


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Flaming Gorge Reservoir Fisheries Assessments

Funding source:
Utah Division of Wildlife Resources

Project Location: Flaming Gorge Reservoir, Utah-Wyoming

Project Duration: January - December 1996

Investigators:
Michael Canning
Chris Luecke

Technicians:
Phaedra Budy
Todd Seamons

Goals and Objectives:

  1. Test spatially-explicit predator-prey model for lake trout and planktivorous fishes in Flaming Gorge Reservoir.
  2. Assess the genetic strains of the two spawning stocks of kokanee in the reservoir, and use these genetic differences to assess dispersion of individuals of these two groups.
  3. Continue development of food web model and test of effects of variation in lake trout abundance and how chlorophyll concentration affects survival, growth and angler harvest of lake trout, kokanee, and rainbow trout.
  4. Compare results of the food web model to results of fishery simulation model developed and used by University of British Columbia fishery biologists.
  5. Continue monitoring program for forage fishes using hydroacoustic surveys in August of 1995.
  6. Continue monthly monitoring of limnological characteristics of the reservoir during the May through September period.

Progress to Date:
A spatially-explicit foraging and bioenergetics model of lake trout growth potential was developed for use in Flaming Gorge Reservoir, Utah. Abundance of forage fish available for lake trout consumption was estimated from hydroacoustic and trawling surveys. These surveys indicated that the abundance of forage fish biomass declined by 40% from 1989-1995. Species dominance shifted from Utah chub to kokanee during this period. Simulation models indicated that the potential growth rate of lake trout should decline by 20% because of changes in the abundance and spatial distribution
of forage fishes. Annual sampling of lake trout in Flaming Gorge Reservoir indicated that the growth rate of marked individuals and the condition factor of individuals captured in annual gill netting surveys declined from 1989-1995. Information on forage fish abundance, lake trout growth, and lake trout density was used to formulate harvest regulations for the lake trout fishery.

A combination of foraging and bioenergetics models are used to assess the potential growth rate of other salmonid fishes given environmental conditions of temperature, water velocity, oxygen concentration, and food availability. Models were developed from literature values on the foraging and physiological relationships of individual species. Model validation was conducted by growing fish in
field enclosures. Three examples highlight the use of such models for lake management. Simulations for kokanee in Flaming gorge Reservoir indicated that fish assess tradeoffs between zooplankton food availability and temperature in choosing habitats. Model simulations were used to assess stocking strategies for salmonids. Finally, growth rate potential for lake trout was estimated to assess the effects of forage fish reductions on the production potential of this highly prized trophy fishery.

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