Appendix B
Spatial Scale of Road Effects on Ecological Conditions: Annotated Bibliography

Laurie Carr

TerraSystems Research

MAY 2003

INTRODUCTION

The following is an annotated bibliography of road effects on ecological conditions, with a special emphasis on spatial scale. Only studies that directly measured the effect of roads on the surrounding environment were included. References were organized into two main categories, abiotic and biotic consequences. Within abiotic consequences, the effects of roads on hydrology, geomorphology, natural disturbances, and the effects of road chemicals on ecosystems are included. The biotic consequences category is further divided into three subcategories, genetic consequences, plant and wildlife population consequences and ecosystem consequences. Within each subcategory the effects of roads on structure, function and composition are included.

Every aspect of roads has some interaction with the surrounding environment, from road construction to maintenance. However, this list focuses on the effects generated from the presence and use of the road itself. With the exception of culverts, the impacts of road structures such as bridges or roadside lamps are not included. The reciprocal effects of the environment on roads are also not included.



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Assessing and Managing the Ecological Impacts of Paved Roads Appendix B Spatial Scale of Road Effects on Ecological Conditions: Annotated Bibliography Laurie Carr TerraSystems Research MAY 2003 INTRODUCTION The following is an annotated bibliography of road effects on ecological conditions, with a special emphasis on spatial scale. Only studies that directly measured the effect of roads on the surrounding environment were included. References were organized into two main categories, abiotic and biotic consequences. Within abiotic consequences, the effects of roads on hydrology, geomorphology, natural disturbances, and the effects of road chemicals on ecosystems are included. The biotic consequences category is further divided into three subcategories, genetic consequences, plant and wildlife population consequences and ecosystem consequences. Within each subcategory the effects of roads on structure, function and composition are included. Every aspect of roads has some interaction with the surrounding environment, from road construction to maintenance. However, this list focuses on the effects generated from the presence and use of the road itself. With the exception of culverts, the impacts of road structures such as bridges or roadside lamps are not included. The reciprocal effects of the environment on roads are also not included.

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Assessing and Managing the Ecological Impacts of Paved Roads For all ecological effects, references are organized by the scale of the study. Three scales were used: single segment, intermediate—political/ecological, and national. Single segment refers to studies that examine the impacts of a single road on an ecosystem. In this case, several roads may be involved in the study; however, the results do not address the cumulative effects of these roads. For example, the effect of road pollution on insect populations living adjacent to a road, or the barrier effect of a road on a species' movement would be classified as “single segment.” Intermediate scale studies examine the cumulative effect of roads on a region. In other words, the combined effect of more than one road determines the results of the study. For example, the effect of several roads on one lake, range expansion by using roads as a dispersal corridor, or genetic isolation of a population surrounded by more than one road would be classified as intermediate. The boundary of a region is either politically (e.g., state of Florida, national park) or ecologically determined (e.g., a watershed, an animal’s home range). National scale studies are very rare and they cover the effects of roads over an entire country. The format for the bibliography is as follows: Sample Section Summary of Ecological Effects A paragraph is written here explaining the ecological effects for the section. Ecological effects are italicized in paragraph. Ecological effect: a. Single segment Reference 1 Reference 2 Etc. b. Intermediate—-political/ecological Reference 1 Reference 2 Etc. c. National Reference 1

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Assessing and Managing the Ecological Impacts of Paved Roads Reference 2 Etc. ABIOTIC CONSEQUENCES OF THE ECOLOGICAL EFFECT OF ROADS Hydrology and Geomorphology Summary of Ecological Effects Roads have an impervious surface that collects and reroutes precipitation along its length or along roadside ditches. The temporary addition of waterflows from the road network affects flooding, groundwater supplies and channel morphology of stream networks. For logging roads specifically, precipitation and vehicular use results in sediment production. Sediment is carried into the watershed by wind or water and often contains chemicals. Roads that transect waterbodies cause changes in waterflows by restricting circulation. Roads can also become barriers to surface drainage when they are elevated compared to the landscape. Stream Networks a. Single segment Almost all streams and intermittent channels crossed by a highway were channelized. Distance from 30 m to 400 m upslope and from 30 m to 500 m downslope of the road. Wetland drainage effects extended outward from the road for distances varying from 50 m to 500 m (Forman and Deblinger 2000). b. Intermediate—political/ecological When hard surface (road, buildings, parking areas) reaches 30-40% of the area about 30% of precipitation water becomes surface runoff, at 80-90% more than 55% of water becomes runoff. As a result, groundwater supplies may not be fully recharged, streams tend to degrade, and flooding often increases (Schueler 1995). Once hard-surface coverage exceeds 25%, streams in area tend to be degraded, as characterized by unstable channel morphology, polluted water, and highly altered or impoverished fish communities. In Seattle (USA) region, a road density of 5 km/km2 (8 mi/mi2) corre-

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Assessing and Managing the Ecological Impacts of Paved Roads sponded to a hard-surface cover of about 20% in the watershed. Hard-surface model suggests stream networks are “impacted” at levels of hard surface as low as 10% coverage. (Center for Watershed Protection 1998). Results from a conceptual model of interactions between road networks and stream networks show that road networks appear to affect floods and debris flows, thus modifying disturbance patch dynamics in stream and riparian networks in mountain landscapes (Jones et al. 2000). Road effects on stream networks result in changes in watershed processes far removed from the site. Roads can increase drainage density by increasing waterflow from an impervious surface, and diverting subsurface water to surface and roadside ditches. This can result in floods, and alter aquatic and riparian ecological conditions, including fish populations in the lower parts of the stream systems (Eaglin and Hubert 1993). Stormwater flows along roads or ditches often effectively create new segments connected to the natural stream network. In a mountain forestry case approximately 57% of the road network functioned as an extension of the stream network, thereby increasing drainage density by 21% to 50% (Wemple, et al. 1996). Water from roadside ditches may be routed to streams, thus effectively increasing the density of stream channels in a watershed. A model predicted increases in the mean annual flood due to forest roads ranged from 2.2% to 9.5% (La March and Lettenmaier 2001). c. National No citations Sediment Production a. Single segment Inventories of almost 500 km of forest roads in several catchments indicate that untreated roads produced 1500 to 4700 m3 of sediment per kilometer of road length (Madej 2001). b. Intermediate—political/ecological Lake Tahoe basin receives chemically laden sediment-bearing road runoff from a highly disturbed, road-laced watershed. Erosion along unpaved timber-harvesting roads caused by vehicular passage leads to sediment transported by wind or water (Zeigler et al. 2001).

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Assessing and Managing the Ecological Impacts of Paved Roads The greatest accumulation of fine sediments in streambeds was associated with logging road areas that exceeded 2.5% of the total basin area. Total road lengths of 2.5 km/km2 of watershed basin produced sediment > 4 times natural rate (Cedarholm et al. 1981) A study of a forest road network in the western Cascade Range, Oregon found that debris slides from mobilized road fills were the dominant process of sediment production from roads. Overall, this study indicated that the nature of geomorphic processes influenced by roads is strongly conditioned by road location and construction practices, basin geology and storm characteristics (Wemple 2001). c. National No citations Changes in Waterflow a. Single segment Restricted tidal flow in salt marsh of northern Massachusetts resulted in difference in salinity and hence vegetation of each side of culverts. Invasion of freshwater common reed (Phragmites communis) occurred in the side of low salinity (Massachusetts Executive Office of Environmental Affairs 1995). A causeway bisecting the Great Salt Lake in Utah altered circulation, resulting in changes in salt concentration between the two sides of the lake (Loving et al. 2000). A major effect of permanent roads in the artic is the blockage of surface drainage during spring snowmelt, thereby allowing water to accumulate beside a road in the relatively flat terrain. This can result in flooded areas or impoundments and sometimes road washouts (Walker et al. 1987). b. Intermediate—political/ecological Four causeways transect an estuary of Florida's Tampa Bay. This has altered circulation of the bay resulting in changes in contaminant and sediment transportation (Goodwin 1987). c. National No citations

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Assessing and Managing the Ecological Impacts of Paved Roads Chemical Characteristics Summary of Ecological Effects Major sources of roadside pollution are vehicles, roads and bridges, and dry and wet (dust and rain) atmospheric deposition. Less frequent sources are accidental spills of oil, gasoline, and industrial chemicals. The majority of roadside chemicals come from vehicles (83%), 22% come from sanding and de-icing agents, 17% from roadbed and road surface wear, and 13% from herbicide and pesticide use. These figures do not include heavy metals and other chemicals that leach from bridges into streams and other water bodies (Federal Highway Administration 1996, Kobringer 1984). Vehicular chemical pollutants include mineral nutrients (e.g. nitrogen and phosphorus), heavy metals (e.g. zinc and lead) and organic compounds (petroleum products). Of all the heavy metals, lead is the most studied ecologically. Lead was removed from gasoline in the mid-80’s in North America, and lead levels in plants and animals are now relatively low (Forman, et al. 2003). The actual levels of heavy metals other than lead (Cadmium, copper, zinc, nickel, mercury and chromium) along roads, and their ecological effects remains poorly understood (Forman, et al. 2003). De-icing salt (sodium chloride) is applied on roads for snow and ice. Literature on the contamination of surface water and groundwater from road-salt is voluminous. Only a few examples are presented in this table. Mineral Nutrients a. Single segment The largest source of phosphorus entering Lake Chocorua in New Hampshire was runoff from a multilane highway that passed near the eastern shoreline (Schloss 2002). Chemical nitrogen (from vehicle exhaust) enrichment of roadside soil favours few dominant flowering plants at the expense of more sensitive conifers, ferns, mosses, fungi, algae, and lichens in heathlands. This effect is higher along multilane highways compared to smaller roads (Angold 1997). Nitrogen (nitrogen oxides from traffic) caused increased growth in plant species in healthland communities in southern England.

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Assessing and Managing the Ecological Impacts of Paved Roads Consequently species composition changed. Effects were measured up to 200 m (656 ft) from road (Angold 1992). b. Intermediate—political/ecological No citations c. National No citations Heavy Metals a. Single segment Increased levels of toxic heavy metals have been found up to 50 or 100 m (165 to 330 ft) of highways in the air, soil and plants (Ministry of Transport 1994b). Manganese concentrations in the soil along interstate highways in Utah are 100 times higher than historic levels. Roadside aquatic plants were sensitive bio-indicators of manganese contamination (Lytle et al. 1995). Some plants may have enhanced root growth as a result of soil contamination from roadside dust carrying trace metals (Wong et al. 1984). Combustion gases may cause reductions in species richness in arthropods, some groups flourished in the environment polluted by combustion gases. (Przybylski 1979). Overall lead levels were low in insects but high in earthworms, especially near highways (Marino et al. 1992). Early studies suggest that the abundance and diversity of invertebrates do not decline with increasing amount of metal pollution in roadside habitats (Muskett and Jones 1980). Lead contamination of insects near a highway in Kansas was more than 3 times level of those far from a road. Meadowlarks did not follow this trend reflecting little exposure to lead contaminated insects (Udevitz et al. 1980). Lead concentrations in little brown bats, short-tailed shrews, and meadow voles adjacent to a highway are high enough to cause mortality in domestic animals. (Clark 1979). Lead in roadside median strips of a highway was not considered a threat to adult ground-foraging songbirds (Grue et al. 1986).

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Assessing and Managing the Ecological Impacts of Paved Roads b. Intermediate—political/ecological No citations c. National No citations Organic a. Single segment Green treefrog (Hyla cinerea) tadpole growth and metamorphosis was negatively associated with presence of petroleum contamination of freshwater, such as would occur from runoff (Mahaney 1994). b. Intermediate—political/ecological No citations c. National No citations De-icing Salt a. Single segment Road-salt (CaCl) on an unpaved forest road inhibited crossing of the road by salamanders. (Demaynadier and Hunter 1995). Survivorship of salamander species was lower in roadside pools that were heavily contaminated by de-icing salts. (Turtle 2000). A highway that crosses the eastern portion of the Hubbard Brook Valley watershed has increased sodium and chloride concentrations in Mirror Lake (Likens et al. 1977). The gradual build-up of salt in soil and lowered moisture conditions can make it more difficult to maintain natural vegetation along roads (Thompson and Rutter 1986). Chloride concentrations in streams downstream from a salted highway were 31 times that upstream from the highway (Demers and Richard 1990.).

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Assessing and Managing the Ecological Impacts of Paved Roads b. Intermediate—political/ecological De-icing salt used in the Rochester, New York, area dissolved and entered Lake Ontario via storm water drainage. This caused increased salinity of Irondequoit Bay to the extent that it prevented vertical mixing of the bay water in spring (Bubeck et al. 1971). Road salting caused changes in salt concentration of a small meromictic lake. This weakened the lake's meromictic (chemical-dependent) stability and had implications for primary productivity of the lake (Kjensmo 1997). c. National No citations Natural Disturbance Summary of Ecological Effects Logging and mountain roads are susceptible for creating landslides due to unstable soil, steep slopes and high road densities (Havlick 2002). Roads can increase water discharge rates in a watershed increasing the potential for landslides. Landslides a. Single segment No citations b. Intermediate—political/ecological Eighty-eight percent of landslides in Boise and Clearwater national forests in Idaho were road related. Most landslides on Idaho’s South Fork of the Salmon River were also road related (Megahan 1980). After the 1964 flood in the Pacific Northwest, landslide frequency due to forest roads was up to 30 times the rates in unmanaged forested areas (Swanson and Dryness 1975). c. National No citations

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Assessing and Managing the Ecological Impacts of Paved Roads GENETIC CONSEQUENCES OF THE ECOLOGICAL EFFECT OF ROADS Structure Summary of Ecological Effects Roads can act as a barrier to movement through road-kill and behavioralioural road avoidance. As barriers, roads subdivide continuous populations and reduce gene flow between sub-populations. The result is isolated and smaller populations. Genetic consequences for larger species with smaller population sizes or for endangered species with low dispersal abilities are thought to be greater (Gerlach and Musolf 2000) but have not been investigated. Barrier to Movement a. Single segment A recent highway, at least 25 yrs old, was shown to have an effect on the genetic sub-structuring of bank vole (Clethrionomys glareolus) populations due to reduced gene flow. Small population size, a country road, and a railway did not appear to affect genetic structure (Gerlach and Musolf 2000). b. Intermediate—-political/ecological Common frog (Rana temporaria) population surrounded by roads, a highway and a railway had reduced average amount of heterozygosity (genetic variation) and genetic polymorphism (diversity of forms) (Reh and Seitz 1990). c. National No citations Function Summary of Ecological Effects Isolated populations have a lower chance of survival without the demographic and genetic input of immigrants, and of recolonization after

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Assessing and Managing the Ecological Impacts of Paved Roads extinction (Lande 1988). Low gene flow in isolated populations has the negative result of increased inbreeding (weak offspring) and decreased fecundity therefore increasing the probability of extinction (Hartl et al. 1992 as cited in Gerlach and Musolf 2000). Little is known on the long-term ecological effects of roads on mammalian speciation through isolation (Baker 1998). Isolated Populations a. Single segment No citations b. Intermediate—-political/ecological No citations c. National No citations Composition Summary of Ecological Effects Roads have a filtering effect, in that different species or segments of a population may interact distinctively in relation to roads. This can result in natural selection of specific genes. Filtering Effect a. Single segment Selection for early flowering and salt tolerance has developed in populations of Anthoxanthum odoratum L., growing along a roadside and in adjacent pastures, in less than 40 years (Kiang 1982). Genetic immunity to car pollution has selected for micropopulations of Tenebrionidae that have lived along roadsides for many generations. Beetle larvae that did not posses the immunity perished at instar II due to pollution exposure (Minoranskii and Kuzina 1984).

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Assessing and Managing the Ecological Impacts of Paved Roads Belford, D. A., and W. R. Gould. 1989. An Evaluation of Trout Passage through Six Highway Culverts in Montana. North American Journal of Fisheries Management 9:437-45. Bellamy, P. E., R. F. Shore, D. Ardeshir, J. R. Treweek, and T. H. Sparks. 2000. Road Verges as Habitat for Small Mammals in Britain. Mammal Review 30:131-39. Bennet. 1991. Roads, Roadsides and Wildlife Conservation: A Review. Pages 99-118 in Saunders and Hobbs, editors. Nature Conservation 2: The Role of Corridors. Surrey Beatty and Sons. Bhattacharya, M., R. B. Primack, and J. Gerwein. 2003. Are Roads and Railroads Barriers to Bumblebee Movement in a Temperate Suburban Conservation Area? Biological Conservation 109:37-45. Braun, S., and W. Fluckiger. 1984. Increased Population of the Aphis Aphis Pomi at a Motorway. Part 2—the Effect of Drought and Deicing Salt. Environmental Pollution (Series A) 36:261-70. Brody, A., and M. Pelton. 1989. Effects of Roads on Black Bear Movements in Western North Carolina. Wildlife Society Bulletin 17:5-10. Bubeck, R. C., W. H. Diment, B. L. Deck, A. L. Baldwin, and S. D. Lipton. 1971. Runoff of Deicing Salt: Effect on Irondequoit Bay, Rochester, New York. Science 172:1128-32. Burnett, S. E. 1992. Effects of a Rainforest Road on Movements of Small Mammals: Mechanisms and Implications. Wildlife Research 19:94-104. Carbaugh, B., J. P. Vaughan, E. D. Bellis, and H. B. Graves. 1975. Distribution and Activity of White-Tailed Deer Along an Interstate Highway. Journal of Wildlife Management 39:570-81. Carr, L. W., and L. Fahrig. 2001. Effect of Road Traffic on Two Amphibian Species of Differing Vagility. Conservation Biology 15:1071-78. Cedarholm, C. J., L. M. Reid, and E. O. Salo. 1981. Cumulative Effects of Logging Road Sediment on Salmonid Populations in the Clearwater River, Jefferson County, Washington. Pages 38-74 in Washington Water Research Council. 1981. Proceedings from the Conference on Salmon-Spawning Gravel: A Renewable Resource in the Pacific Northwest. Washington State University, Washington Water Research Centre, Report 39, Pullman, Washington. Center for Watershed Protection. 1998. Rapid Watershed Planning Handbook: A Comprehensive Guide for Managing Urbanizing Watersheds. Center for Watershed Protection, Elliot City, Md. Chapin, F. S., and G. R. Shaver. 1981. Changes in Soil Properties and Vegetation Following Disturbance of Alaskan Arctic Tundra. Journal of Applied Ecology 18:605-17. Clark, D. R. 1979. Lead Concentrations: Bats vs. Terrestrial Small Mammals Collected near a Major Highway. Environmental Science and Technology 13:338-40.

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Assessing and Managing the Ecological Impacts of Paved Roads Clark, W. D., and J. R. Karr. 1979. Effects of Highways on Red-Winged Black-bird and Horned Lark Populations. Wilson Bulletin 91:143-45. Clarke, G. P., P. C. L. White, and S. Harris. 1998. Effects of Roads on Badger Meles Meles Populations in South-West England. Biological Conservation 86:117-24. Cole, E. K., M. D. Pope, and R. G. Anthony. 1997. Effects of Road Management on Movement and Survival of Roosevelt Elk. Journal of Wildlife Managemen. 61:1115-26. Curatolo, J. A., and S. M. Murphy. 1986. The Effects of Pipelines, Roads and Traffic on the Movements of Caribou, Rangifer Tarandus. Canadian Field Naturalist 100:218-24. Demaynadier, P. G., and M. L. Hunter. 1995. The Relationship between Forest Management and Amphibian Ecology: A Review of the North American Literature. Environmental Review 3:230-61. Demaynadier, P. G., and M. L. Hunter. 2000. Road Effects on Amphibian Movements in a Forested Landscape. Natural Areas Journal 20:56-65. Demers, C. L., and W. S. Richard. 1990. Effects of Road Deicing Salt on Chloride Levels in Four Adirondack Streams. Water, Air, Soil Pollution 49:369-73. Demers, M. N. 1993. Roadside Ditches as Corridors for Range Expansion of the Western Harvester Ant (Pogonomyrmex Occidentalis Cresson). Landscape Ecology 8:93-102. Develey, P. F., and P. C. Stouffer. 2001. Effects of Roads on Movements by Understory Birds in Mixed-Species Flocks in Central Amazonian Brazil. Conservation Biology 15:1416 Eaglin, G. S., and W. A. Hubert. 1993. Effects of Logging and Roads on Substrate and Trout in Streams of the Medicine Bow National Forest, Wyoming. North American Journal of Fisheries Management 13:844-46. Fahrig, L., J. H. Pedlar, S. E. Pope, P. D. Taylor, and J. F. Wegner. 1995. Effect of Road Traffic on Amphibian Density. Biological Conservation 74: 177-82. Federal Highway Administration. 1996. Evaluation and Management of Highway Runoff Water Quality. FHWA-PD-96-032, U.S. Department of transportation, Washington, D.C. Ferreras, P., J. J. Aldama, J. F. Beltran, and M. Delibes. 1992. Rates and Causes of Mortality in a Fragmented Population of Iberian Lynx—Felis Pardina Temminck, 1824. Biological Conservation 61:197-202. Ferris, C. R. 1979. Effects of Interstate 95 on Breeding Birds in Northern Maine. Journal of Wildlife Management 43:421-27. Findlay, C. S., and J. Bourdages. 2000. Response Time of Wetland Biodiversity to Road Construction on Adjacent Lands. Conservation Biology 14:86-94. Findlay, C. S., and J. Houlahan. 1997. Anthropogenic Correlates of Species Richness in Southeastern Ontario Wetlands. Conservation Biology 11:1000-09.

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Assessing and Managing the Ecological Impacts of Paved Roads Foppen, R., and R. Reijnen. 1994. The Effects of Car Traffic on Breeding Bird Populations in Woodland. II. Breeding Dispersal of Male Willow Warblers (Phylloscopus Trochilus) in Relation to the Proximity of a Highway. Journal of Applied Ecology 31:95-101. Forman, R. T. T., and R. D. Deblinger. 2000. The Ecological Road-Effect Zone of a Massachusetts (U.S.A.) Suburban Highway. Conservation Biology 14:36 Forman, R. T. T., B. Reineking, and A. M. Hersperger. 2002. Road Traffic and Nearby Grassland Bird Patterns in a Suburbanizing Landscape. Environmental Management 29:782-800. Forman, R. T. T., and et al. 2003. Road Ecology: Science and Solutions. Island Press, Washington, D.C. Forysa, E. A., C. R. Allen, and D. P. Wojcikc. 2002. Influence of the Proximity and Amount of Human Development and Roads on the Occurrence of the Red Imported Fire Ant in the Lower Florida Keys. Biological Conservation 108:27-33. Fowle, S. C. 1990. The Painted Turtle in the Mission Valley of Western Montana. Master’s. University of Montana, Missoula. Fuller, T. 1989. Population Dynamics of Wolves in North-Central Minnesota. Wildlife Monographs 105:1-41. Garland, T. J., and W. G. Bradley. 1984. Effects of Highway on Mojave Desert Rodent Populations. American Midland Naturalist 111:47-56. Gerlach, G., and K. Musolf. 2000. Fragmentation of Landscape as a Cause for Genetic Subdivision in Bank Voles. Conservation Biology 14:1066-74. Getz, L. L., F. R. Cole, and D. L. Gates. 1978. Interstate Roadsides as Dispersal Routes for Microtus Pennsylvanicus. Journal of Mammalogy 59:208-12. Gibbs, J. P. 1998. Amphibian Movements in Response to Forest Edges, Roads, and Streambeds in Southern New England. Journal of Wildlife Management 62:584-89. Gibeau, M. L. 2000. A Conservation Biology Approach to Management of Grizzly Bears in Banff National Park, Alberta. Ph.D. University of Calgary, Alberta. Gilbert, O. L. 1989. The Ecology of Urban Habitats. Chapman and Hall Ltd. New York. Goldsmith, C. D., and P. F. Scanlon. 1977. Lead Levels in Small Mammals and Selected Invertebrates Associated with Highways of Different Traffic Densities. Bulletin of Environmental Contamination and Toxicology 17:311-16. Goodwin, C. R. 1987. Tidal-Flow, Circulation, and Flushing Changes Caused by Dredge and Fill in Tampa Bay, Florida. U.S.G.S. Water-Supply Paper 2282, U.S. Geological Survey, Washington, D.C. Goosem, M. 2000. Effects of Tropical Rainforest Roads on Small Mammals: Edge Changes in Community Composition. Wildlife Research 27:151-63.

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Assessing and Managing the Ecological Impacts of Paved Roads Goosem, M. 2001. Effects of Tropical Rainforest Roads on Small Mammals: Inhibition of Crossing Movements. Wildlife Research 28:351-64. Greenberg, C. H., S. H. Crownover, and D. R. Gordon. 1997. Roadside Soils: A Corridor for Invasion of Xeric Scrub by Nonindigenous Plants. Natural Areas Journal 17:99-109. Grue, C. E., D. J. Hoffman, W. N. Beyer, and L. P. Franson. 1986. Lead Concentrations and Reproductive Success in European Starlings (Sturnus Vulgaris) Nesting within Highway Roadside Verges. Environmental Pollution (Series A) 42:157-82. Haskell, D. G. 2000. Effects of Forest Roads on Macroinvertebrate Soil Fauna of the Southern Appalachian Mountains. Conservation Biology 14:57. Havlick, D. G. 2002. No Place Distant. Island Press, Washington, D.C. Hels, T., and E. Buchwald. 2001. The Effect of Road Kills on Amphibian Populations. Biological Conservation 99:331-40. Huey, L. M. 1941. Mammalian Invasion Via the Highway. Journal of Mammalogy 22:383-85. Huijser, M. P., and P. J. M. Bergers. 2000. The Effect of Roads and Traffic on Hedgehog (Erinaceus Europaeus) Populations. Biological Conservation 95:111-16. Jaeger, J. A. G., and L. Fahrig. 2001. Modeling the Effects of Road Network Patterns on Population Persistence: Relative Importance of Traffic Mortality and ‘Fence Effect.’ Pages 298-312 in G. Evink et al., eds. Proceedings of the International Conference on Ecology and Transportation. Centre for Transportation and the Environment, North Carolina State University, Raleigh, North Carolina USA. Jensen, W. F., T. K. Fuller, and W. L. Robinson. 1986. Wolf (Canis Lupus), Distribution on the Ontario-Michigan Border near Sault Ste. Marie. Canadian Field Naturalist 100:363-66. Jones, J. A., F. J. Swanson, B. C. Wemple, and K. U. Snyder. 2000. Effects of Roads on Hydrology, Geomorphology, and Disturbance Patches in Stream Networks. Conservation Biology 14:76-85. Jones, M. E. 2000. Road Upgrade, Road Mortality and Remedial Measures: Impacts on a Population of Eastern Quolls and Tasmanian Devils. Wildlife Research 27:289-96. Kasworm, W. F., and T. Manley. 1990. Road and Trail Influences on Grizzly and Black Bears in Northwest Montana. International Conference on Bear Research and Managemet 8:79-84. Keller, V. E. 1991. The Effect of Disturbance from Roads on the Distribution of Feeding Sites of Geese (Anser Brachyrhynchus, A. Anser), Wintering in North-East Scotland. Ardea 79:229-32. Kiang, Y. T. 1982. Local Differentiation of Anthoxanthum Odoratum L. Populations on Roadsides. American Midland Naturalist 107:340-50.

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