Table of Contents Executive Summary ............................................................................................................ 2 Introduction ......................................................................................................................... 3 Problem Statement .......................................................................................................... 3 Stakeholders .................................................................................................................... 3 Goals ............................................................................................................................... 4 Site Description ................................................................................................................... 5 Methods............................................................................................................................... 7 Research .............................................................................................................................. 8 Habitat Destruction, Alteration, and Fragmentation ....................................................... 8 Amphibian Movement Patterns and the Effects of Roads ............................................ 10 Global Amphibian Decline ........................................................................................... 20 Anthropogenic Causes of Amphibian Population Decline ........................................... 21 Consequences of Amphibian Loss ................................................................................ 25 Amphibians and Culverts .............................................................................................. 25 Culvert Replacement/Upgrade Costs ............................................................................ 27 Case Studies: Management Strategies ......................................................................... 29 Site Assessment Metric ................................................................................................. 32 Alternatives Table ............................................................................................................. 34 Alternatives Analysis and Discussion ............................................................................... 35 Recommendations ............................................................................................................. 38 Current Actions ............................................................................................................. 38 Further Research Needs ................................................................................................ 41 References ......................................................................................................................... 46 Page 1 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Executive Summary Purpose of project Talk about agency Talk about why grand isle and franklin Talk about evolution Need for future information This is designed to get the ball rolling until funding and knowledge catch up… Page 2 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Introduction Problem Statement Numerous species of amphibians found in Vermont wetlands have wide-ranging habitat requirements, leading to movement around and between wetlands and into upland habitats. This behavior coupled with the presence of roads through and near wetlands has led to high levels of amphibian mortality due to vehicle encounters. No safe passage currently exists for these delicate organisms to safely cross roads. Current actions in Vermont are sparse and mainly consist of using fencing to prevent access to roadways. This action is effective at reducing vehicle encounters but it will not work at any sites with species that need to access habitat on the other side of the road. Existing infrastructure contains thousands of road crossings such as culverts and bridges that are currently not heavily used by amphibians while crossing roads. A management plan is needed to utilize existing infrastructure to allow amphibians to safely cross the road while remaining economically feasible. In order to maximize the benefits of this plan a detailed site assessment metric is necessary. Stakeholders Vermont Agency of Natural Resources (VTRANS): The reason that amphibian road crossings may indeed become a reality in Vermont is due to the commitment of the staff of the Vermont Agency of Transportation (VTrans) Environmental Section. Currently, they are involved in several projects, such as a kestrel nest box program, a “frog fence” to prevent frog mortality on a busy state highway, mapping vegetation communities along I-89 in an attempt to reconfigure mowing schedules to least impact wildlife, as well as researching methods to implement more wildlife crossings on state highways and to reduce impacts of storm water discharges on water quality. It is their goal to make VTrans a proactive, environmentally conscious transportation agency that will become a model for other state transportation agencies. Page 3 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Biologists and ecologists: This problem is of special concern to scientists due to the numerous factors leading to widespread amphibian decline. These species are already very stressed by habitat loss, pollution, UV radiation, and numerous other challenges. Reptiles and amphibians are an important link in wetland food webs. Recreational people and sightseers: Many people relate the gentle chirping of spring peepers or the rough calls of bullfrogs to the feeling of being with nature. Catching frogs is a favorite activity for small children and some adults. Cyclists have complained about high numbers of frog road kill in the past as both disgusting and a potential safety hazard. Motorists: Hopefully few people feel good about running over snakes, turtles, or frogs. There is also a potential for accidents to occur while swerving to avoid these animals. The general public: The general public is the most important stakeholder due to the potential need of taxpayer funding for any construction projects related to addressing this problem. The general public may have cause for concern over mosquitoes and mosquito-borne diseases such as West-Nile due to the heavy feeding of amphibians on larval and adult mosquitoes. Political Representatives: Political representatives have a measurable impact on potential changes to be made. County representatives would be an integral part of any construction activities, site assessment, and funding arrangements. Volunteers: A Boy Scout troop or other civilian group could be a useful way to maintain silt fences or other impermanent structures. Civilian groups from other parts of the country have formed bucket brigades to move amphibians across the road. Goals Research causes and effects of amphibian decline on global and local scales to emphasize the importance of reducing mortality sources when feasible/possible Page 4 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Develop a metric for assessing and prioritizing sites for their suitability and need for road crossing modifications Determine “why the frog crossed the road”: Conduct research to determine why amphibians and reptiles move and come in contact with roadways Create a detailed list of short-term management actions for low-cost highreturn road crossing alternatives Create a detailed list of long-term management actions for permanent road crossing structures Present recommendations for VTRANS to address vehicular mortality in amphibian populations using crossing structures along Vermont-state roads Site Description Vermont is located in the northwest corner of New England and is bordered by Lake Champlain and New York state to the west; Quebec, Canada to the north; New Hampshire to the east; and Massachusetts to the south. It is oriented primarily along a north-south axis. Latitude is between 42º and 45º north, and longitude is between 70º and 73º west (Perkins Museum of Geology website). Total land area is approximately 9,250 square miles (U.S. Census Bureau website). Vermont topography varies throughout the state, which has led to the state being divided into six physiographic regions: the Green Mountains, the Taconic Mountains, the Valley of Vermont, the Champlain Lowlands, the Vermont Piedmont, and the Northeast Highlands. The Green Mountains run the entire north-south length of Vermont, down approximately the middle of the state, and contain the highest peaks in Vermont. The Taconic Mountains are located in southwest Vermont and are lower, rounder, and older than the Green Mountains. The Valley of Vermont is a flat, narrow strip of land that runs between the Green Mountains to the west and the Taconic Mountains to the east. It is essentially an extension of the Champlain Lowlands, which is the area including Lake Champlain and its surrounding, fertile, relatively flat basin. The Champlain lowlands are bordered to the west by the Green Mountains, to the east by the Adirondack Mountains, Page 5 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont and to the south by the Taconic Mountains. Due to the moderating effects of Lake Champlain and the protection offered by the mountain ranges on three sides, this area experiences the mildest climate in the state. The Vermont Piedmont runs the north-south length of the state to the east of the Green Mountains, comprising the foothills of the Green Mountains and characterized by rolling hills, wide valleys, and many lakes. Finally, the Northeast Highlands are located in the northeast corner of Vermont and are actually an extension of the White Mountains of New Hampshire, making them quite different from any other region in Vermont. Here, the climate is second in severity to only the highest peaks of the Green Mountains (Johnson 1998). Generally, the climate of Vermont is considered humid continental-cool summer, which is characterized by average temperature of the coldest month (January) < 32º F and average temperature of the warmest month < 72º F. In Vermont, the temperature is influenced more by altitude in the summer, by latitude in the winter, and varies with topography statewide. The average length of the growing season also differs depending on topography, from approximately 90 days at higher elevations and in small, low valleys to 150 days near Lake Champlain. Lake Champlain has a considerable buffering effect on the climate of the Champlain Valley (Meeks 1986). Precipitation patterns mirror those of temperature, again varying with topography. Average annual precipitation ranges from approximately 32” to 52”. In general, higher elevations (Green Mountains) receive more precipitation, as do the southern part of the state and the Northeast Highlands. Vermont is also one of the cloudiest places in North America; the amount of solar energy that falls on Vermont is nearly the smallest amount in the United States (Meeks 1986). Dominant vegetation can be approximated into four primary zones. Northern hardwoods (birch, beech, and maple) dominate below 2,000 feet, except in Northeast Vermont. The central Champlain Valley is characterized by oak-hickory forest. Above 2,000 feet and in the Northeast Highlands, the primary vegetation communities consist of spruce-fir. Finally, on the few peaks close to 4,000 feet, there are scattered spots of alpine vegetation. In many of the lower elevations, such as in Franklin and Addison Counties, dairy farms still comprise a large portion of the landscape, so these areas may be less forested than other locations in the state (Meeks 1986). Page 6 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Vermont remains one of the most rural states in the nation, with an estimated 2001 population estimate of 613,090 people. In 2000, the average number of persons per square mile was 65.8. Population is most dense in the Greater Burlington/Chittenden County area, and also in the Rutland, Brattleboro, and Montpelier areas (U.S. Census Bureau website). The 9, 250 square mile land area is traversed by a total of approximately 15,009 miles of roads (14,305 miles of state public roads and 704 miles of the National Highway System). In 2002, the grand total of urban and rural annual vehicle miles of travel was 9,573.8 (VTrans website). Although it is difficult to summarize traffic statistics, for there is too much variance between different roadways, a generalization that may be made is that traffic decreases in intensity from Interstate highways; down through Class I, II, III, and IV state highways; to town highways. However, this is only a generalization, for town roads in more urbanized locations may experience greater traffic intensity than state highways in more rural locations. Historical Context Methods Research: We conducted an extensive literature review to determine the context, extent, and factors related to amphibian mortality with roads and other causes. Through email we communicated with people in Transportation Industry such as Nelson Hoffman, Chris Slesar, and Chad Allen from VTRANS. Information was also gathered via email with people in Construction Agencies such as Robert Flint of the Florida Department of Transportation to determine costs of various efforts. Email with biologists such as Scott Jackson from University of Massachusetts was used to gather information on amphibian requirements for culverts. Site Assessment: We designed a site assessment metric to prioritize sites based on several parameters. GIS data layers containing these parameters were downloaded Vermont Center for Geographic Information (VCGI); we utilized all of the applicable data currently available in digital format for Grand Isle and Franklin Counties. The metric was developed to include all culverts found within 400 meters of wetlands. This distance was chosen due to it being the approximate maximum distance traveled by amphibians that migrate to and from wetland habitats. Within these culverts we then prioritized sites based on traffic volume, Page 7 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont viable upland habitat, and proximity to ideal amphibian habitat (wetlands). A traffic volume of 6000 vehicles per day was selected to represent an estimated mortality rate of >90% for amphibians crossing the road (Hels and Buchwald 2001). We then narrowed the ranges on the existing parameters to identify high priority sites. The high priority sites were chosen based on the presence of wetland habitat within 100 meters of a culvert (this includes the maximum distance traveled between habitats for almost all Vermont amphibians), and a road with a traffic volume > 6000 vehicles per day. A detailed GIS map of all highpriority sites was created including all data relevant to the metric. Recommendations: The recommendations were compiled through analysis and interpretation of the findings and from the results of the metric. Based on the parameters needed for a detailed statewide site assessment metric we also compiled a list of missing information. Research The following section of this report will include all of our research findings utilized in creating recommendations for future management actions. An extensive literature review was conducted to find both the factors and causes behind the problem as well as potential management alternatives. Vehicle encounters are a significant source of mortality for certain amphibian species due to movement patters and the increasing presence of roads and vehicles near and through amphibian habitat. The need to reduce this source of mortality is greatly strengthened by global population declines evident in amphibian populations across the planet. These population declines are thought to be the result of anthropogenic degradation of the global environment. This section of the report will also present the research findings that formed the basis for our management recommendations for future actions of the Vermont Agency of Transportation. Habitat Destruction, Alteration, and Fragmentation Most amphibians require both terrestrial and aquatic habitats to complete their life cycles, which makes them even more vulnerable to habitat loss, alteration, and Page 8 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont fragmentation than most other wildlife species. Loss of either habitat may have negative impacts on amphibian populations, yet both have been greatly modified by anthropogenic disturbances. Some of the major land-use practices affecting amphibians include agriculture, silviculture, industry, and urban development. These practices often lead to the filling and draining of aquatic breeding habitat (especially wetlands and vernal pools), the removal of trees and other vegetation in the terrestrial uplands used by adults for feeding and refuge, and the alteration of the hydrodynamics associated with stream and river ecosystems (Semlitsch 2003a). Habitat destruction and habitat alteration are two distinct processes, but both have detrimental effects on amphibian populations. Additionally, they both contribute to habitat fragmentation, out of which arises a host of additional problems. Habitat destruction is the complete, and often permanent, loss of an ecosystem and its former biological function. This occurs when wetlands are drained or filled; forests are converted to parking lots, housing developments, agricultural fields, etc.; or when an ecosystem becomes so chemically contaminated that it is uninhabitable. On the other hand, habitat alteration occurs when an ecosystem function is changed, although not completely or permanently. An example of habitat alteration is when trees are cut in a forest as part of a forestry operation. Removal of the canopy will have an effect on the microhabitat and microclimate of the forest, perhaps reducing or eliminating amphibians, but the trees will grow back and suitable habitat will once again exist. The effects of habitat destruction and alteration become most pronounced when they occur on a sufficient scale that habitat fragmentation occurs (Dodd and Smith 2003). As many amphibian populations exhibit metapopulation structure, fragmentation of both aquatic and terrestrial habitats may lead to numerous issues. As the number of suitable breeding sites decreases, the total number of individuals available to recolonize extirpated populations or form new populations is diminished. This may lead to a reduction in the number of source populations as well. The loss of aquatic habitat may also lead to isolation of populations and the subsequent loss of genetic diversity, as they may be too far apart to allow successful dispersal. Even more important than aquatic breeding sites may be intact terrestrial habitat, because this provides movement corridors that connect local populations. Because amphibians are prone to desiccation, they require Page 9 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont moist microclimates provided by intact canopy cover, coarse woody debris, and leaf litter to successfully migrate between habitats. Barriers introduced by fragmentation, such as housing developments, roads, and agricultural fields, may restrict movement (Semlitsch 2003b). The immediate effects of fragmentation on amphibian populations include the loss of the most vulnerable individuals, populations, and species. As the landscape becomes increasingly fragmented, the natural linkages between habitat patches are lost and barriers to movement are introduced. At this point, they may become isolated from habitats required to complete their life cycles, such as if aquatic breeding sites are cut off from terrestrial sites. There may be an increased risk of predation as edge effects are introduced and cover is reduced. Fragmentation may remain a filter to movement or may become an impenetrable barrier (Dodd and Smith 2003). Long-term effects of fragmentation include the loss of genetic variability, decreased survivorship, isolation, increased edge effects, and greater susceptibility to stochastic events. Small, isolated populations confined to the tiny pieces of remaining habitat become susceptible to the accumulation of mutations and the loss of adaptive potential. Edge effects become more pronounced and may alter the physical structure and microclimate of the habitat. There is often the loss of migratory routes and dispersal corridors, compounding the problems of isolation. These long-term pressures on the populations, combined with loss of individuals to direct mortality, can become too much for the populations to overcome. They become more vulnerable to extirpation, and possibly extinction, due to random stochastic events, such as disease, drought, etc. (Dodd and Smith 2003). Amphibian Movement Patterns and the Effects of Roads Amphibians are frequently characterized by complex life histories, often requiring different habitats at various life stages in order to complete their life cycles. Contrary to popular belief, most amphibians are not entirely aquatic; in fact, some species are completely terrestrial, and other species may require both terrestrial and aquatic habitats. These different habitats may be widely spatially distributed, leading to the development of metapopulation structure and dynamics. This distribution also drives movement Page 10 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont patterns in amphibians, particularly those with the biphasic life cycle, which may require migrations between foraging, breeding, and overwintering sites. These species are therefore quite vulnerable to the effects of habitat fragmentation, and specifically to mortality on roadways that bisect their home ranges (Dodd and Smith 2003). Life History Entirely aquatic amphibians include the salamander species hellbender (Crytobranchus alleganiensis) and mudpuppy (Necturus maculosus), but relatively few amphibians overall are confined to water throughout their lives. The movements of this group are largely limited to spatial and temporal distributions within the water column and to habitat structures within the water body. Being confined to water, they are the group least likely to be impacted by direct mortality upon roadways, as they will not be crossing the road and getting hit by a vehicle. The effects of roads on entirely aquatic amphibians is probably limited to habitat destruction through road construction, impacts from road salt and other chemicals, and other indirect impacts related to roadways (Dodd and Smith 2003). Other amphibians, such as the eastern red-backed salamander (Plethodon cinereus), have entirely terrestrial life histories. These amphibians are characterized by direct development; eggs are deposited on land and hatch into miniature adults, or they give birth to live young (Dodd and Smith 2003). As is true for any amphibian species, they are prone to desiccation and therefore require a humid environment, such as is provided by the moist microclimates under deciduous forest leaf-litter and coarse woody debris (Pough et al. 1987; deMaynadier and Hunter 1998). They may also inhabit subterranean or arboreal habitats, and may move between subsurface, surface, and arboreal habitats depending on season, life stage, and/or reproductive needs (Dodd and Smith 2003). Although they may be driven to cross roads periodically, this is probably restricted mostly to juvenile dispersal or other density-dependent causes, as roads have been found to be the most important factor hindering amphibian movement. In fact, a study by Gibbs (1998), in which the red-backed salamander was one of the species of interest, demonstrated that forest-road edges are substantially less permeable than forestopen land edges and even forest-residential edges. This indicates that entirely terrestrial Page 11 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont amphibians probably are not inclined to leave the forest to cross a road unless some underlying factor necessitates it for survival. Groups that may be forced to cross roads more frequently therefore, are the amphibians exhibiting a biphasic life cycle, in which primarily terrestrial adults must move to a water body to breed and deposit eggs that eventually hatch into aquatic larvae (tadpoles). These species may be vulnerable to roadway mortality repeatedly throughout their lives if their habitat has been fragmented by a road network. For example, terrestrial adults may have to cross a road to access an aquatic breeding site, and then cross back again to return to terrestrial foraging habitat. Some species may need to cross again to find overwintering habitat (Hels and Buchwald 2001). Upon completion of metamorphosis, dispersing juveniles may be forced to mass migrate across roadways in pursuit of suitable habitat. The juveniles may disperse hundreds to thousands of meters from their natal pools, increasing the odds that they will encounter a road at some point (Dodd and Smith 2003). As this is the group of amphibians most at risk for roadway mortality, the remainder of this paper will focus primarily upon those with the biphasic lifestyle. Aquatic Breeding Habitat Although the migration towards suitable aquatic breeding habitat is necessary for reproduction, the destination water body varies between species. Some species utilize large, permanent water bodies, while others prefer small, ephemeral (vernal) pools. Generally, those species that breed in large, permanent water bodies produce larvae with slower growth and larger body sizes at metamorphosis, which leads to increased fitness as adults (Semlitsch 2003). An example is the American bullfrog (Rana catesbeiana), whose tadpoles take 2 to 3 years to complete metamorphosis. This is in contrast to a vernal pool specialist, such as the eastern spadefoot toad (Scaphiopus holbrookii), in which metamorphosis occurs rapidly in a matter of days to weeks at the expense of reduced body size (Dodd and Smith 2003). This speedy growth process is necessitated by the temporary nature of the aquatic habitat; these larvae must develop quickly before the pool dries out. But the key distinction of vernal pools is the lack of predatory fish and tadpoles (such as the American bullfrog). Larvae in permanent water bodies often possess anti-predator defenses, while those in vernal pools do not. Therefore, a vernal Page 12 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont pool’s hydroperiod must be long enough to allow the larvae to complete metamorphosis, but short enough to preclude predatory fish and tadpoles (Semlitsch 2003). A discussion of vernal pools is necessary when considering where to place road crossings, for these pools should be prioritized in any road-crossing program. They may possess a more diverse array of amphibian species than larger water bodies; the species may also be of more threatened status (Dodd 1992 as cited in Semlitsch 2003). However, vernal pools are often overlooked, even in wetland protection regulations (Preisser et al. 2000). This is an unfortunate oversight, for considering the irreplaceable function that they provide as breeding habitat for so many amphibian species, they should be of primary concern. For example: 14 species of frogs were found breeding in temporary ponds within upland areas in north-central Florida (Moler and Franz as cited in Dodd and Cade 1998), 20 amphibian species were discovered in a temporary pond on the coastal plain of Georgia (Cash 1994 as cited in Dodd and Cade 1998), 15 amphibian species were noted in a temporary pond in north-central Florida (Dodd 1992 as cited in Dodd and Cade 1998), and 16 species of amphibians were documented in temporary ponds in northern Florida flatwoods (O’Neill 1995 as cited in Dodd and Cade 1998). In New England, vernal pool obligates include the wood frog (Rana sylvatica), eastern spadefoot toad, spotted salamander (Ambystoma maculatum), marbled salamander (Ambystoma opacum), and Jefferson salamander (Ambystoma jeffersonianum) (Preisser et al. 2000). Metapopulation Dynamics The wide spatial distribution of suitable terrestrial and aquatic habitats for biphasic amphibians, especially the vernal pool specialists, often leads to the development of metapopulation structure (Dodd and Smith 2003). “A metapopulation is a set of local populations… connected by processes of migration, gene flow, extinction, and colonization. Two primary factors control amphibian metapopulation dynamics: 1) the number or density of individuals dispersing among ponds, and 2) the density and distribution of wetlands in the landscape that determine dispersal distances and the probability of successfully reaching ponds” (Semlitsch 2003). Page 13 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Taking into account the factors noted by Semlitsch, loss of wetlands, and particularly vernal pools, increases the distances between ponds and leads to the isolation of populations. This is particularly true for amphibians, as they cannot migrate long distances due to the physiological limitations of desiccation. This can lead to loss of gene flow or extirpation of local populations if the remaining wetlands are too far apart to allow for recolonization (Semlitsch 2003). General Effects of Roads on Amphibian Populations One can extend this reasoning to include the effects of roadways on metapopulations. Roadways may fragment habitat, making it more difficult to migrate to breeding sites. Also, high mortality incurred upon roadway crossing could serve to decimate an already stressed population. Vos and Chardon (1998) discovered that road density has a negative effect on the occupation probability of a pond by moor frogs (Rana arvalis). In areas with high road densities, occupation probability was reduced to 55 %. In the part of the study adjacent to a highway, occupation probability was lowered to less than 30 %. They point out that although the size and availability of breeding habitat is generally thought to be the limiting factor determining population size in amphibians, there are now indications that the distribution of suitable terrestrial habitat is becoming more critical due to high mortality in the terrestrial phase. This high mortality is being attributed more frequently to road kill as more roads are built and traffic densities continue to increase. Aquatic breeding amphibians are most vulnerable to roadway mortality due to their movement patterns, population structures, and preferred habitats (Hels and Buchwald 2001). Amphibians are small-bodied (which makes them less visible to drivers) (Semlitsch 2003), slow-moving, and non-cognizant of the dangers posed by vehicles (Ashley and Robinson 1996). Generally, two factors interact to determine where, when, and how severe roadway mortality of amphibians occurs: the movement patterns of the species and the intensity of the traffic. Movement Patterns Amphibians most often encounter roads during seasonal migrations between breeding sites, summer habitat, and hibernation sites, or during dispersal to new habitat patches (Ashley and Robinson 1996; Vos and Chardon 1998). Ashley and Robinson Page 14 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont (1996), in a study conducted along a causeway on Lake Erie, observed seasonal highs in spring and autumn, consistent with life history patterns of reproduction and dispersal. Depending on the species, unimodal (leopard frog) and bimodal (bullfrog) patterns of road crossing were detected between April and October, again consistent with the life history pattern of the species. Amphibian mortality was significantly associated with the adjacent roadside habitat and vegetation communities preferred by various species. This indicates that some species were attempting to cross to desirable habitat on the other side of the causeway. The amphibians crossed mostly at night when traffic flow was light, which may have reduced the overall mortality (Ashley and Robinson 1996). Movement patterns may also be influenced by environmental factors, such as temperature and moisture (Sinsch 1988; Bayliss 1995; Bartlet 2000; as cited in Muths 2000). Roadway Mortality The Ashley and Robinson (1996) study along a Lake Erie causeway documented that short-lived species, which produce many young and are among the more common and abundant amphibian species, constituted the largest numbers of road kill: northern leopard frog (Rana pipiens), American bullfrog, green frog (Rana clamitans), and American toad (Bufo americanus americanus). It has been noted that a species often found killed along a road may simply reflect the presence of a large and thriving population (Huijser and Bergers 1997; Mallick et al. 1998; both as cited in Hels and Buchwald 2001). Hels and Buchwald (2001) describe three factors they believe affect a species vulnerability to road mortality: 1) velocity of the species, 2) diurnal movement pattern of the species, and 3) diurnal movement pattern of vehicles. In their study, activity patterns of amphibians were concentrated at night, when traffic flows were lowest, yet they still report annual mortality of 25 % of the reproductively active adult population of spadefoot toad (Pelobates fuscus) and 21 % of the combined reproductively active populations of the common frog (Rana temporaria) and moor frog. The probability of being killed increases with increasing traffic intensity (e.g., 34% - 61% probability of being killed crossing a road with 3,207 cars per day, increasing to 89% - 98% on a busy highway). Survival increases exponentially with velocity of the amphibian, and decreases exponentially with increasing traffic intensity. Up to a traffic Page 15 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont intensity of 625 vehicles per hour (15,000 vehicles per day), the velocity of the amphibian has a large influence on its probability of being killed. Above this traffic intensity, the probability of being killed is very close to 100% for all species regardless of velocity (Hels and Buchwald 2001). Hels and Buchwald (2001) additionally note several other interesting observations. For example, the angle of crossing also affects probability of being killed. Survival is greater with perpendicular crossing, with survival steadily decreasing as the crossing angle deviates from perpendicular. Unlike most animals, amphibians, due to their small body size, are killed mostly through direct hits by a vehicle’s wheel and may remain unharmed if they stay still under a passing vehicle, although some larger vehicles may kill through wind speed alone. Finally, they also note that estimates of amphibian road kill are often underestimated, as small body sizes may mean they go undetected, repeatedly being run over may obliterate the corpse, or they may be eaten by scavengers. Roadway Mortality Impact But how much of an impact does road mortality actually have on amphibian populations? Fahrig et al (1995) found that total number of dead and live frogs per kilometer (km) decreased with increasing traffic intensity. The proportion of dead frogs and toads increased with increasing traffic intensity, and the density of live frogs and toads decreased with increasing traffic intensity. Collectively, these results indicate that traffic mortality has a significant negative effect on local density of amphibians. At a rate of 24 – 40 vehicles per hour, Kuhn (1987, as cited in Clevenger et al. 2001) found that 50 % of a cohort of migrating common toads (Bufo bufo) was killed. A study by van Gelder (1973) on common toad breeding migration estimated that 60 cars per hour would kill 90 % of the adult toads. In yet another study, Heine (1987, as cited in Clevenger et al. 2001) predicted that at 26 vehicles per hour, the estimated survival rate of the common toad is zero. Ehmann and Cogger (1985, as cited in Fahrig et al. 1995) conservatively estimated that 5,480,000 reptiles and frogs are killed annually in Australia. Ashley and Robinson (1996) documented 27,846 road-killed young-of-the-year leopard frogs, while total amphibian road mortality averaged 11.65 amphibians per km per day between the months of April and October. Clearly, road mortality may have long-term impacts on amphibian populations (Semlitsch 2003). Page 16 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Exactly how much of an impact road mortality will have depends on the species. Some populations may be depressed by road mortality while others may experience little impact (Ashley and Robinson 1996). It mostly depends on whether road mortality is additive or compensatory for a given population. In general, for adults, density- independent mortality factors are more important, while for larvae both densitydependent and density-independent factors matter (Duellman and Trueb 1994 and references therein, as cited in Hels and Buchwald 2001). If a population is mainly regulated by density-independent factors, then road mortality would be an additive effect, and the impact to the population would be greater. If a population is regulated more by density-dependent factors, then road mortality would be compensatory, and the overall impact on the population would be much smaller. However, traffic intensity may eventually increase to the point where it has an additive effect on an otherwise densitydependent population (Hels and Buchwald 2001). For threatened species, road mortality often does have an additive effect that is detrimental to populations and metapopulations (Semlitsch 2003). Management Considerations: Predicting Crossing Locations An important management consideration is how to best predict where amphibian roadway crossings are likely to occur, so that measures may be implemented to reduce roadway mortality, especially for species experiencing additive effects. As mentioned previously, those species most likely to be crossing a road are those with the biphasic lifestyle. Therefore, two habitat requirements must be within migratory proximity of a road: terrestrial (for most species, preferably deciduous forest) and aquatic. The remaining questions then concern the distance an amphibian will be likely to travel to access a given habitat type and any evident patterns to their movement directions and pathways. The answers to these questions vary quite widely based on species, although a good rule of thumb would seem to be a movement distance estimate of several hundred meters. A review of the literature offered the following wide-ranging figures, a summary of which appears in Table One. Semlitsch (2003) states that amphibians usually live within a few hundred meters (m) of their natal aquatic habitat, although some do disperse to new sites. Vernal pool specialists often breed in a single pool throughout their Page 17 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont reproductive lives and live in the surrounding uplands, dispersing an average of 125 m into the uplands after breeding (Semlitsch 1998, as cited in Preisser et al. 2000). Juvenile salamanders may disperse up to 670 m and juvenile frogs up to 1 km from their natal pool (J. Victoria, personal communication; Berven and Grudzien 1990, both as cited in Preisser et al. 2000). The mean distance traveled from breeding ponds for boreal toads (Bufo boreas) is 721.46 m (females) and 218.15 m (males); 92 % of all movements are within 700 m of the breeding site (Muths 2003). Spadefoot toads move a maximum of 1200 m between their hibernation sites and their breeding ponds (Nöllert 1990 in Hels and Buchwald 2001). Moor frogs migrate a maximum distance of 350 m and common frogs a maximum distance of 600 m (Haapanen 1970 in Hels and Buchwald 2001). Striped newts (Notophthalmus perstriatus) will travel up to 709 m, while eastern narrowmouthed toads (Gastrophryne carolinensis) will travel up to 914 m (Dodd 1996, in Dodd and Cade 1998). The average of all the above figures is 628 m. This indicates that amphibians will travel considerable distances for their small sizes to reach suitable breeding and foraging habitats. These numbers also indicate that any road within approximately 100 m to 1,000 m of a wetland could potentially become the site of an amphibian roadway crossing. Movement directions and patterns also vary considerably between species, making the identification of corridors difficult, if even possible. Some species are reported to move in specific directions when entering or leaving a breeding site, and it has been suggested that some species follow migratory corridors (Stenhouse 1985; Verrell 1987; both as cited in Dodd and Cade 1998). For example, the mole salamander (Ambystoma talpoideum) travels to and from breeding sites in a nonrandom manner (Semlitsch 1981, as cited in Muths 2000). Certain toads may also move in linear patterns away from breeding sites (Bartlet 2000, as cited in Muths 2000). Shoop (1968, as cited in Dodd and Cade 1998) detected a travel corridor approximately 30 m wide for the spotted salamander in Massachusetts. Other studies have found less specificity. Stenhouse (1985, as cited in Dodd and Cade 1998) noted that individual spotted salamanders tend to enter and exit ponds at the same location each year, although no specific pathway is followed among a group. Dodd and Cade (1998) found that striped newts and eastern narrow-mouthed toads favor certain directions of movement over Page 18 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont others even though no specific corridor is followed, indicating that they probably move directly to the breeding pond from their terrestrial habitat. It does appear from some of the studies that amphibians may follow some kind of general pathway or direction during breeding migrations, making it likely that an under-road passageway in association with appropriate funneling techniques could prove successful in channeling the amphibians away from death on the roadway. Table X. Summary of distances traveled by amphibians between habitat types. Species Distance Traveled Between Aquatic and Terrestrial Habitats (meters) Amphibians, general 400 Vernal pool specialists 125 Juvenile salamanders 670 Juvenile frogs 1000 Boreal toad, female 721 Boreal toad, male 218 Spadefoot toad 1200 Moor frog 350 Common frog 600 Striped newt 709 Eastern narrow-mouthed toad 914 Average distance traveled, all species 628 Page 19 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Global Amphibian Decline A literature review of global amphibian declines was used to strengthen the need for reducing amphibian mortality on Vermont roadways. Globally, amphibian populations are in dire conditions due to anthropogenic impacts on the environment. Many of the identified factors behind these declines are global problems that have no attainable short-term solution. The combination of these mechanisms is actually enough to lead scientists to worry about the future of amphibians as a whole. Because of this, it is critically important to reduce sources of amphibian mortality wherever it is feasible. The following section of this report will describe some of the sources of global population decline Global Population Trends A large-scale study of global amphibian population trends was compiled which strongly suggested population decline is occurring at a global scale (Houlahan et al 2000). The study consisted of 936 data sets from publications, reports, and unpublished data from 37 countries and included 157 species. This study found that a rapid global amphibian decline from 1960-1966 was followed by a less intensive decline from 19661997. The decline rates were approximately 15% annually and 2% annually respectively with a total of 61 documented extinctions (Houlahan et al 2000). A majority of the historic declines were observed in Europe, but more recent rapid declines have occurred in Costa Rica, Panama, and Australia. Until recently, most scientists believed these declines were simply cyclical downturns in natural population dynamics. However, new evidence indicates that these declines suggest a more catastrophic cycle (Dalton 2000). Many of the recent population declines are of special concern due to the fact that they occurred in protected areas such as reserves and parks with no clear causative agent (Halliday 1998). Several well-documented localized population declines have been observed in relatively undisturbed sites. These declines are strong evidence for some type of global factor that is likely an indirect anthropogenic effect (Halliday 1998). A majority of the species-specific declines have occurred in isolated high-altitude ecosystems (Houlahan et al 2001). There are countless examples of species-specific declines that serve to illustrate the global connection between local declines. The Golden Toad (Bufo periglenes) was Page 20 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont discovered in a Costa Rican Cloud Forest and went from fairly common with an estimated population of 1,000 down to a single toad within a few years. The gastric brooding frog (Rheobatrachus silius), only discovered in 1973, was found in relative abundance until its rapid decline to extinction in 1981 (Phillips 1990). Anthropogenic Causes of Amphibian Population Decline A single clear causative agent for global amphibian decline has yet to be identified, but numerous anthropogenic influences have been shown to negatively affect amphibian populations. Mechanisms range from direct anthropogenic impacts such as habitat destruction, to more indirect impacts such as pesticide interactions increasing susceptibility to parasites and fungal infections. Global Climate Change Global climate change has been linked to two major vectors of local population declines. Localized warming has produced niche overlapping and increased interactions between species that historically had been isolated temporally and spatially (Walther et al 2002). Climatic warming in Britain has led to a source of population decline from a previously insignificant interaction between populations of newts and frogs. Both species share breeding habitats in shallow lakes and wetlands, but share different temporal breeding niches. Warmer winters have created variable responses within the two populations. Historically, salamanders have entered the breeding areas well after the frog eggs have already hatched, but due to warming they are entering earlier. The frogs have not significantly changed their breeding timing and are now suffering egg mortality from a new source of predation (Walther et al 2002). Climate change has also created strong warming in the air-water temperature averages in the tropical regions of the Pacific Ocean leading to increased occurrence of large-scale climatic oscillation cycles. These cycles drastically affect local climate in many areas and are responsible for the recent widespread devastation observed in numerous ecosystems from coral reefs to cloud forests (Pounds 2001). Climate change has been implicated as an indirect cause for numerous disease outbreaks responsible for amphibian population declines globally (Pounds 2001). Page 21 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Fungal Pathogen Outbreaks Global climate change has also been linked through a very complex system to amphibian mortality through fungal infections. Numerous studies have found that certain amphibian populations are being exposed to increased levels of UV-B radiation, resulting in increased rates of fungal infections (Kiesecher et al 2001 and Pounds 2001). Amphibians that breed in shallow montane lakes have been most impacted by UV-B related mortality. Western Toads (Bufo boreas) are the most studied montane species for this mechanism of population decline. Studies have found that their string of eggs develop normally for a few days and then turn cloudy and white experiencing drastic mortality from a lethal fungus infection (Pounds 2001). The fungal strain Saprolegnia ferax has been documented as the primary cause of these population declines found in the Northwestern United States (Kiesecher et al 2001). Most of these populations have rapidly decreased since the 1980’s and numerous montane populations are now extinct (Kiesecher et al 2001). The mechanism for the Saprolegnia ferax infection is an example of widespread environmental problems working together synergistically. Previous studies had linked fungal mortality to stratospheric ozone depletion, but new research has shown that climate is the primary factor (Pounds 2001). The climatic Figure 1: Relationship between water column depth and hatchling survivorship (Kiesecher et al 2001) oscillations mentioned earlier in this report are directly correlated with snowfall in the Cascades and therefore annual lake levels (Pounds 2001). Fungal infections have been linked to decreased water levels, which increases UV-B exposure in oviposition sites utilized by the Western Toad as shown in Figure 1 (Kiesecher et al 2001). This mechanism has been observed in the Western Toad and other montane species that lay their eggs in shallow water areas. Water column depth is an efficient filter of UV-B radiation, with eggs laid in 50 cm of water receiving half of the UV-B as Page 22 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont eggs in 10cm of water (Pounds 2001). The direct mechanism for fungal infection is not yet know but laboratory manipulation has shown in 33% increase in infection rates for Western Toad eggs exposed to UV-B. In natural lakes, mortality rates of 80% are common in shallow sites (20cm) while only 12% in water deeper than 50cm (Pounds 2001). Increased UV-B radiation from either stratospheric ozone depletion, or waterdraw down may also harm any other shallow water breeders as well(Pounds 2001). Amphibian populations in Central America and the Australian Highlands are experiencing population declines from fungal infections that appear to be from a different mechanism. Tilarian rain frogs (Eleutherodactylus angelicus) in Costa Rica have experienced fungal infections very similar to Western Toads, but the rain frogs lay their eggs in shade, ruling out UV-B exposure as a causative agent (Pounds 2001). Amphibian declines observed in Australia and Central America were caused by a chytrid fungus, which affects cutaneous respiration. Concurrent declines in reptile populations, which are not susceptible to this type of infection, suggest that a different mechanism must be present (Pounds 2001). This type of chytrid fungus had not been previously documented. The fact that it turned up on opposite sides of the world leads scientists to believe that it is an opportunistic fungus which appeared due to compromised immune system ability, increasing the susceptibility of amphibians to infection (Halliday 1998). Numerous other pathogens have recently increased in abundance suggesting that environmental factors are leading to decreased immune system function (Halliday 1998). Herbicides A more direct anthropogenic cause of amphibian mortality has been shown through numerous studies relating the popular herbicide Atrazine to endocrine disruption in frogs. Atrazine is the most commonly used herbicide in the United States, with approximately 27,000 tons applied annually. It is also used in over 80 countries, making Atrazine one of the most important herbicides globally (Dalton 2002). Atrazine contains endocrine disruptors that have been linked to disruptions of sexual development in frogs (Dalton 2002). This source of amphibian mortality was first described in the 1980’s when researchers observed increases in frog mortality and deformities (Netting 2000). Studies have found that levels as low as 0.1ppb cause male tadpoles to develop ovaries in addition to testes. Levels greater that 1 ppb lead to the deformation of larynges, organs Page 23 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont that are used to call for mates. Levels higher than 25 ppb are shown to cause drastic drops in testosterone levels, which have been observed in wild leopard frogs in numerous sites with high levels of Atrazine. Different from most lab studies of toxicity, these numbers are ecologically relevant because American waters routinely have levels above 50ppb and in some extreme cases, concentrations can be measured in parts per million (Dalton 2002). Species Introduction Species introduction is yet another anthropogenic influence that is leading to amphibian population decline. The introduction of predatory fish to water bodies has led to an indirect source of stress and mortality in tadpoles. When in the presence of predatory fish, tadpoles take cover on the lake bottom and remain relatively motionless. This behavior reduces fish predation, but it greatly increases the risk of parasite infestation. Parasitic infestations rarely result in direct mortality, but the attacks damage kidneys and other organs, leaving a legacy of increased susceptibility to disease as adults. Increased stocking programs have introduced predatory fish to countless aquatic systems, greatly increasing this mechanism of amphibian population decline (Netting 2001). Wetland destruction can also lead to increased parasitism by concentrating tadpoles and snails that host the parasites, in the remaining wetland patches (Netting 2001). Another factor increasing parasitism is the potential for some pesticides to reduce vigor and mobility of tadpoles (Netting 2001). Bull-frog (Rana catesbeiana) and exotic fish introduction has become common practice among permanent wetlands in the United States for aquaculture and sport-fishing purposes, and have recently been shown to reduce populations of native anurans (Adams 1999). Bull-frogs are generalist feeders and predators that negatively impact native amphibians through predation and competition. Many of the exotic fish commonly introduced to aquatic systems prey on tadpoles and adult amphibians. Tadpole mortality from fish predation can be incredibly high in some species that evolved in the absence of predatory fish. This source of predation is very prominent in more developed areas, where most wetlands have bullfrog or fish introduction. Some studies have found that it is very rare to find any native amphibians in systems once bullfrogs have become established (Adams 1999). Page 24 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Consequences of Amphibian Loss The loss of amphibian species, through numerous mechanisms, has far-ranging implications for ecological food-webs and ecosystem structure. Amphibian diet consists primarily of insects, followed by other invertebrates and vertebrates. Amphibians convert invertebrate biomass into a food source more available to higher trophic levels (Cofrin Center for Biodiversity, 2003). Due to very small body size, amphibians are also able to consume prey too small to be accessible to birds and mammals. An individual Blanchard’s cricket frog consumes close to 4800 insects a year(Cofrin Center for Biodiversity, 2003). This, coupled with very low metabolic rates, makes the conversion of insect biomass into amphibian biomass very efficient (Pough et al, 1987). Many amphibian species provide important ecotone interactions by foraging for insects in upland habitats. In some Vermont forests, eastern red-backed salamander (Plethodon cinerus) biomass is greater than birds and small mammals (Cofrin Center for Biodiversity, 2003). The annual biomass production of red-back salamanders in these forests can be greater than that of all birds and mammals (Pough et al, 1987). The importance of amphibians as ecosystem and food-web components stresses the consequences of widespread declines. Amphibians and Culverts Some types of amphibian crossings have been more successful than others. To keep amphibians from crossing the road, there are a couple of options. Silt fences can be used to deter the amphibian from crossing. They are relatively inexpensive, requiring only wooden stakes, and canvas. However, silt fences are labor intensive and necessitate maintenance. A community group or a Boy Scout troop could provide this upholding. Concrete walls also deter amphibians from crossing the road. Concrete walls are more expensive than silt fences, but require much less maintenance. Box-shaped culverts and round culverts are options for amphibian to cross under a road. The choices of pipes are bituminous-coated corrugated steel, High Density Polythene Page 25 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Pipe, corrugated aluminum pipe, and reinforced concrete pipe. The choice for box culverts is concrete (LWDD, 2003). Vermont is beginning to turn its focus towards protecting amphibians from road mortality. Because there is less human population in Vermont, the roads have different amounts of travel than heavily traveled Florida roads. Therefore different measures should be taken to protect them. VTRANS could be guided by the research done at the University of Massachusetts. At University Of Massachusetts, Scott Jackson has led the way for the study of amphibian culvert research. The variables affecting use are said to be size, placement, moisture, hydrology, temperature, and noise (Jackson, 2000). It is important to consider what amphibians prefer, when crossing under a road. “Relatively small amphibian and reptile tunnels may be a cost effective means of mitigating highway impacts where roads and highways are located between wetlands and upland habitats” (Jackson, 2000). Scott Jackson recommends using box culverts over pipes. As for the size, he suggests bigger culverts with a wider diameter. This will be more accessible for animals meaning less mortality. Jackson advises at least two feet by two feet squared at the openings. Open tops that allow light will attract more amphibians. Jackson suggests using concrete. If a pipe is to be used, he also suggests a good standard culvert Image of a box culvert from http://www.acousa.com/wildlife.htm diameter is fifteen inches (Jackson, 2000). Gino Guimarro the senior project manager at Woodlot Alternatives, an environmental consultant firm, states that a square bottom would have more usable bottom area (Guimarro, 2003). He also believes culverts should be up to 36 inches diameter to allow bigger animals such as racoons through (Ruediger, 2003). The substrate of the culvert is important for promoting amphibian crossing. Jackson advocates using sandy soil for the bottom of the culvert as well as providing cover such as tree stumps. Bill Ruediger, the Ecology Program Leader for Highways at the USDA Forest Service’s Washington Office recommends a natural bottom surface such as rock or dirt (Ruediger, 2003). Page 26 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Culvert Showing natural substrate and wing walls to guide amphibians under the road.http://www.fhwa.dot.gov/environment/wildlifecrossings/photo25.htm Wing walls are used to funnel the amphibians into the tunnel. The wing walls ought to extend at forty-five degrees to funnel the amphibians toward the culvert. The walls should extend vertically at least eighteen inches. The vertical walls should extend one hundred to two hundred feet outward and the tops of the walls should be parallel with the ground surface on the side closest to the road. Jackson recommends that the crossing structures should be placed less than one hundred feet apart (Jackson, 2000). Comparison of different culvert types is important for Vermont so that the least expensive measure can be used. “There is evidence that amphibian and reptile tunnels are effective when used with two lane roads…. it is not known how effective they will be for facilitating passage beneath highways of four or more lanes” (Langston et al in Jackson, 2000). Ruediger recommends in order of preference, cement box culverts, cement round culverts, PVC pipe, then corrugated steel as the least preferable (Ruediger, 2003). Culvert Replacement/Upgrade Costs Round culverts are less expensive than box shaped culverts, so in the interest of budget; this paper will focus on them. The four types of materials used for pipe culverts are steel, aluminum, and concrete, and HDPE, or High Density Polythene Pipe. Table 11 shows the cost of production and the installation price for two lane roads. The table also Page 27 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont shows the budget for VTRANS in 2004. $50,722,184 goes to maintenance, which is where the culvert replacement would take place (Hoffman, 2003). Table 1. The price of different culvert materials and installation. Culvert Type 2 lane cost (per foot) + $750 installation HDPE 15 inch diameter 21.04 Concrete Box Culvert 2x2 ft 350-700 Reinforced Concrete Pipe 28 Corrugated Metal Pipe 26.12 Total Budget for VTRANS 2004 Amount of VTRANS available for Culverts $301,404,219 $50,722,184 Dave Lathrop of (??) averages culvert replacement costs between $5,000 $10,000 dollars. He states that amphibian culverts are closer to $5,000 because they are dug at a small depth. However, the figures depend on many variables such as size and length of culvert, traffic control, and depth of culvert (Lathrop, 2003). Gil Newbury of (???) states that the minimum culvert size used under state highways is 18 inches. Also culverts less than 4 feet in diameter are primarily made of plastic. Gil adds that cost depends not only on depth, but also on the wetness of the site. He expects a cost of between $3,000 and $5,000 (Newbury, 2003). Chris Slesar of VTRANS states that access to a site will affect the cost. He uses the example that if a culvert needs to be installed on a steep slope, then more labor will increase the costs. Other variables he lists are traffic volumes and guard rail presence (Slesar, 2003). Chad Allen of VTRANS says that the material cost of HDPE, CMP, and RCP (reinforced concrete pipe) are $21.04/LF, $26.12/LF, and $28.00/LF correspondingly. Therefore if a 35 to 40 foot pipe was fitted under a 2 lane road, then the difference in material only comes out to about $150-$200. For a $5,500 job, this is fairly unimportant. Allen also states that the District's first choice is to install HDPE. The other materials such as CMP, are used in areas where the culvert needs to be installed deep within the ground (Allen, 2003). Page 28 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont The problems encountered are that price of the pipes depends on how much the consumer will buy. Robert Flint of the Florida Department of Transportation explains that the more bulk of pipe bought, the less expensive it will be. He said “ we may have a price for a 24" pipe (Concrete Pipe) that is 8 ft. long. If the quantity needed is 1,000 ft., then the price is $23.00 per foot” (Flint, 2003). Also, a potential problem is the culvert’s function in a storm event. Chad Allen, District 5 Transportation Program Manager, recommends that the culvert equalize water between wetlands or push water away from the road (Allen, 2003). Case Studies: Management Strategies Switzerland Switzerland has a long history of research and action based on animal crossing. Swiss scientists have developed a Geographic Information System (GIS) that shows animal habitat and corridors where many animals travel across roads. In these places, the Swiss have adapted numerous methods for preventing animal collisions with cars. When the concern is only larger game like deer, they have constructed warning signs that can detect animal movement along roadside. When an animal moves across the motion detector the sign flashes on warning oncoming traffic that something is close to the road. Constructing large animal overpasses is another method used by the Swiss. Through many studies the Swiss have determined that the best overpass for animal use would have a minimum width of 50 meters (Bank, 2002). By incorporating vegetation and water, the overpass is like the natural environment (Figure 1). This method has proven successful with a wide variety of users from large game to small amphibians and insects. Overpasses are constructed with video cameras and tracking plots to measure their effectiveness. Tracking plots are built in the crossings and have either ink pads that dirty up the animals feet causing it to leave tracks, or fine sand pits that the animal must cross over. When crossed the fine sand will have imprints of the animal’s feet (Sauvajot 2002). When the target animals are small mammals or amphibians an underpass may be used. There are many different methods for building underpasses but one that the Swiss Page 29 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont have used is a one-way tunnel system. This system funnels amphibians from upland habitats to breeding sites in the spring and fall. The only way for the animal to cross the road is through the one-way viaduct, this lowers animal death and still allows the amphibians to reach their breeding ground without major difficulty. One major method the Swiss use to ensure the safety of animals at road crossing is teamwork between transportation and environmental planners. Without teamwork and concern for the animals not much can be accomplished (Sauvajot 2002). Germany Germany has gone in slightly a different direction when dealing with overpasses. They have created their overpasses to be used by more then just animals. They have created their overpasses to incorporate humans as well as animals. In farm areas, they have built overpasses with tractor trails as well as pathways for walking and biking. This method is beneficial to animals and the local communities. These overpasses not only connect habitat but also farmland. In addition, Germany has focused on underpasses that are built specifically to prevent noise pollution. Passing cars create noise that prevents some animals Figure 2: Cement underpass without sound buffers from using the tunnels. By building tunnels deeper under the road, it cuts down on some of the noise pollution. In Figure 2, Page 30 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont the underpass is built just under the road’s surface. This underpass does not sufficiently cut down on noise. Different types of fences can also be used to cut down on the noise near roads (Hagood, 2002). The Netherlands The Netherlands has used similar methods as the other countries. Overpasses are built for all types of animals, from large mammals to small insects and many underpasses have been built for small mammals and amphibians. In their underpasses they have built in wood or earth shelves that are built for mammals to cross in watery culverts. In all, the Netherlands have approximately 600 culverts, with only ten built specifically for amphibians (Gray, 2002). The United States The United States has come a long way since concern over wildlife began around roadways. Many states have begun planning and building crossings. Florida for example, has already created many amphibian crossing areas. They have progressed faster then most of the other states and they have an extensive system of crossing culverts. In one two mile stretch on Route 441 in Florida, they have built a 3 1/5 foot high concrete wall with a 6 inch lip at the top (as seen in Figure 3) that directs reptiles and amphibians along the road to an underpass (Southall, 2000). This wall prevents animals from climbing to the roads and it has been a success. Figure 3: Florida roadway Concern throughout the United States has been growing and something must soon be done. Many planning committees have used the methods discovered and implemented in Europe. There is no one solution, each situation is separate and can be handled Page 31 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont differently. The future for animal crossings here in the United States looks promising. As long as there is public concern, animals will be protected. Site Assessment Metric The Purpose of a Metric From measuring the success and excellence of businesses, to quantifying a range of ecological factors, metrics can be used for a multitude of useful applications. In recent years, they have been used to assess biodiversity, degrees of environmental degradation (Sutter 1992), population dynamics, and habitat suitability (Mitch and Gosselink 2000). Indeed, metrics, also often referred to as indices, can be used to evaluate largely anything that a scientist might desire to study. In the environmental field, the use of metrics has proven to be a successful method for quantifying otherwise subjective environmental factors. Assessing Sites of Amphibian Mortality The development of a metric is a somewhat arbitrary practice. However, in this instance a metric is simply a useful by-product of a landscape/habitat assessment. VTRANS, the state of Vermont’s transportation department, has expressed a desire to implement a program designed to prevent amphibian mortality due to road crossings. Each year, large quantities of amphibians are flattened by passing cars, which presents both environmental and safety problems. VTRANS has experimented with small-scale structures designed to prevent amphibians from crossing state highways, however, they are interested in more permanent solutions to a problem they anticipate will otherwise be long-term. Inherent Limitations of Metrics As is true with any metric, there will be flaws inherent to the final product, most often related to incomplete, insufficient, or inaccurate data. Please refer to the “Recommendations for Further Research” section of this paper for information on missing data related to our metric. In any instance, it is recommended that any necessary, or otherwise helpful, data be obtained as soon as possible in order to facilitate the further development of the metric. Lack of funding may preclude extensive data collection, Page 32 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont although the hiring of college interns or other relatively inexpensive help could assist in making the data collection possible. Page 33 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Alternatives Table Decrease Mortality Habitat Connectivity No Action Temporary Road Closing No Change 100% decrease when implemented No Change Temporary Barriers to Crossing Close to 100% No Change Reduced by almost 100 Percent Slight Decrease No Change Decrease Warning Signs Add Temporary Funneling Walls to Culverts Replace Existing Culverts to be More Suitable Towards Amphibians Humans Moving Amphibians in Buckets Permanent Funnel Wall Construction Specific Culverts for Amphibians Wilderness Overpass Public Support/ Involvement Volunteers Needed None None Costs No Change Maximum None Low Minimum Maximum Low Minimum Minimum Low Increase No Change Barriers will be visible from road, but will decrease roadkill Slight Decrease due to more signs Walls will be visible from road, but will decrease roadkill Minimum Maximum Low Decrease Large decrease only when implemented Increase Decrease roadkill Minimum Minimum Low No Change Minimum Maximum Low Decrease Increase Decrease Roadkill May be visible from road, will decrease roadkill Maximum Minimum High Decrease Increase Decrease roadkill Maximum Minimum Decrease Increase Personal Preference Maximum Minimum Medium Very High Aesthetics Continue to see dead amphibians on road Need 1 sentence qualifiers in footnote for maximum, etc Page 34 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Alternatives Analysis and Discussion No Action This alternative suggests the Agency of Transportation to continue with the same policies that they currently have. There would be no increase in spending or change in policy. This alternative does not address the problem. It does nothing to prevent amphibian mortality, raise public awareness, or raise the level of connectivity. Closing Roads This alternative addresses the problem and would prevent amphibian mortality, but only a couple times a year. It does nothing to address the problem during the rest of the year. It does nothing to increase connectivity. By closing the roads the public may be upset but it will cause them to question why this is happening, raising overall awareness of the topic. Temporary Barriers to Crossing By building temporary barriers the mortality rate will be decreased almost completely. The barriers will prevent amphibians from getting onto the roads. They are cheap and easy to install but will need volunteer help with installation, upkeep and deconstruction. One problem with this alternative is connectivity. Barriers from the road with no crossing mechanism completely decrease connectivity. Warning Signs Warning Signs are fairly inexpensive and they have little upkeep. They are easy to implement and do not need much public support. This alternative does little to prevent amphibian mortality. It does nothing to prevent amphibians from going onto roads. It only makes a suggestion to drivers to be careful and does nothing to enforce driver safety. This alternative does not help with connectivity. Add Temporary Funneling Walls to Culverts This alternative will decrease mortality because it will direct amphibians toward the culvert entrance. It will increase the amphibian populations because less will be killed in the roads. However, the wall could confuse the amphibians that don’t move along the fence. This would make them likely to be predated on by birds. Habitats would be Page 35 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont connected using this alternative, unless wear and tear happens to the silt fence. The aesthetics of a silt fence would be intermediate because they are used commonly for house construction. This alternative requires a great amount of labor. The fence must be constructed in the late spring or early summer and taken down in the fall and could be done by civilian groups such as boy scouts. The prices of the drift fences are minimal in comparison with a more permanent lipped concrete wall. Replace Existing Culverts This alternative would decrease amphibian mortality by making the culverts more accessible for amphibian to cross. This alternative would increase habitat connectivity and would have the potential to increase aesthetics. The public support would be minimal as well as the volunteers needed. In this case, existing culverts will need to be replaced anyway. The cost would depend on what alterations needed to be done and could range greatly. Humans Moving Amphibians in Buckets This alternative would require a lot of volunteer work moving amphibians across the road. It would mean a decrease in mortality in the high migration points of the year, but could not guarantee 100% survival and does nothing for any other times throughout the year. It would have no affect on habitat connectivity, and would not affect aesthetics because nothing is being changed. It would be positive because it would encourage community interactions with each other and the environment. This alternative would cost nothing to the state but could have an effect on the traffic, such as causing people to slow down. Permanent Funnel Wall Construction This alternative would decrease mortality and would be beneficial because it would not require much maintenance. This alternative would connect two habitats almost seamlessly. The aesthetics of a concrete wall would be negative. In order to get the funding for a project like this, public support would be needed. Volunteers would not be needed for this project. The cost of this project would be high. Page 36 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Specific Culverts for Amphibians These culverts would nearly eliminate mortality and would maximize habitat connectivity. Amphibian specific culverts are seamless which could have the potential to increase aesthetics. Public support would be needed to encourage VTRANS to install this type of culvert. No volunteers would be needed. The price of these culverts is high. Wilderness Overpass The potential for a wilderness overpass could nearly eliminate mortality by naturally connecting two habitats. An overpass would be aesthetically pleasing to people who view nature as good. Public support for this project would be needed to encourage VTRANS funding. No volunteers would be needed. The cost would be very expensive taking into consideration that bridges cost more than digging a culvert. Page 37 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Recommendations The main goal of this report is to provide management recommendation for the Vermont Agency of Transportation to address the problem of amphibian mortality on Vermont roadways. This report has thus far presented background information on the problem and has provided a list of possible management alternatives. Our recommendation to the agency is not to select a single alternative, but to work within the framework of actions utilizing adaptive management. Due to funding and public support limitations many of the alternatives are not currently feasible. If the project is successful public support and funding will gradually increase making further actions possible. This section of the report will first outline our short-term management recommendations, and then discuss the importance of public support and the information that is still needed to finalize the metric for statewide usage. Our long-term recommendations are based on the compilation of currently lacking datasets as well as increased funding, which will hopefully be available in the future. Current Actions In order to maximize the benefits to amphibians while working within the constraints of funding limitations, we strongly recommend that VTRANS work to implement several of the low-cost alternatives at high-priority sites. The initial actions should be directed at both reducing mortality while raising public awareness. We recommend a combination of amphibian crossing signs and installation of silt-fence funnel walls into existing culverts. We also recommend the continued usage of silt-fence barrier walls at sites where the amphibians do not need to cross the road, similar to the current program on Rt. 2 at Sandbar. The site assessment metric was used to identify 3 high priority sites based on proximity to wetlands and high traffic volume. These sites would be a good starting point to test the effectiveness and feasibility of implementing the alternatives presented in this report. If these sites are successful in both raising awareness of the issue and in reducing amphibian mortality, then more sites should be selected for action. By proving the effectiveness of these actions it will be possible to Page 38 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont gain further public support and funding to allow the implementation of more expensive and more permanent management alternatives. Monitoring One crucial aspect of implementing amphibian-crossing measures is monitoring their effectiveness. In order for the statewide effort to gain momentum and public support, results must be evident. Installation of amphibian crossing structures must be proven to be both used by amphibians and to reduce roadway mortality. Currently, VTRANS documents amphibian mortality by spray painting carcasses on the roadway so that they are not counted again in the future. This method is fairly effective, but it does not capture all of the mortality due to some amphibians being obliterated upon impact, or taken off of the road by scavengers before they are counted. This method is very inexpensive and will be an integral part of any effectiveness monitoring. One-time counts at previously un-assessed sites can also be used to estimate the degree of mortality rates. Monitoring will also be crucial in determining the usage of crossing structures. Jim Andrews recommended using bucket traps set in the ground at the mouth of culvert on the far side of the road. Amphibians simply fall into these buckets after crossing the road and are trapped by the steep smooth walls. The buckets must be checked several times a night to identify, count, and release all of the organisms. This method is very effective but there are two major concerns. Any predator lucky enough to stumble upon a bucket full of tasty frogs is a serious threat, but the use of a raised grate can easily keep predators out. Another concern is the amount of labor needed to maintain these traps. Volunteers are the best option to work through the night checking the traps. Monitoring to prove the effectiveness of crossing structures will be a integral part of gaining public support to continue the evolution of this project in Vermont. Public Support An effective amphibian road-crossing program could end up being quite costly, and if implemented by a state agency, the funds could primarily come from tax dollars. Therefore, public support will be essential to any amphibian road-crossing plan. However, as amphibians are not at the forefront of most wildlife activism crusades, Page 39 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont public support may be more difficult to gain than it would be for more majestic wildlife species. A comprehensive plan for gathering public support must be developed prior to enacting any amphibian-crossing project. Although the intent of this paper is not to develop a plan for developing public support, several brief suggestions will be noted. A concern is that public support will be difficult to procure, because many people do not value amphibians the way they may value more visible and well-understood wildlife species. One way to educate the public about these critters is to hold a series of informational public meetings across the state when the decision is made to fund a crossing program, at least partially, with tax dollars. Another medium that may be used to reach the public is the media, such as descriptive articles in local newspapers or segments on local news programs. An additional suggestion is to appeal to fishermen, birdwatchers, and other outdoors people by including information about amphibians’ importance at the base of the food chain, and the implications of this for other wildlife, in fishing regulation books and at wildlife refuges and other access areas. One way to reach a large segment of the public in a positive manner is through enlightened children. Guest speakers on amphibians, along with live specimens, incorporated into elementary school curriculums would raise the interest and awareness of the students. The students would then, hopefully, go home and inform their families of what they learned; that is, they would relay to the adults the importance of amphibians and how a road crossing program can help reduce mortality. A child’s enthusiasm for a given subject is often a more effective method for swaying an adult’s opinion than is any public service announcement. One of the most important aspects of public support involved with this project is the need for volunteer efforts and ownership. Unfortunately, most of our short-term management recommendations require a large degree of labor. The labor needs include installing, maintaining, and removing the silt fencing every year, to checking bucket traps used in monitoring, to carrying amphibians across to safety in a bucket brigade. We suggest that community groups such as Boy Scout troops or Science classes be approached to take ownership of the labor needs at the sites. As discussed earlier, public involvement can be used to benefit both the public through education and the amphibians. Page 40 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Further Research Needs INTRO PARAGRAPH Culvert Data – More comprehensive culvert data would be useful to determine replacement needs. Specifically, it should be noted that data pertaining to size and the date of last replacement for each culvert are not available in digital format at this time; it will take a substantial amount of time and effort to compile this data into a usable format. In addition, it will be necessary to collect data regarding the average “wetness” of the culverts contained within the study area. The amount of water that flows through a culvert dictates whether or not an amphibian would be capable of passing through it (Andrews 2003). Since such a database does not currently exist, it will be necessary to develop one before the metric can be truly completed. Endangered Species Data - It should be noted that the endangered species database for the state of Vermont has not been updated since 1997 (VT FWS 1997); this data may need to be updated accordingly. This may be done, specifically for amphibians, utilizing town-level documented species data for the state. This data, developed as part of the Vermont Biodiversity Project, is available through the Vermont Center for Geographic Information and was most recently updated in 2000 (Buford 2000). Known Crossing Sites – It is important to include all known sites of high amphibian mortality (due to road crossings) in the state, and any known sites of high amphibian movement/migration. This data will aid in determining which areas are most in need of amphibian crossing structures. However, such data has not been compiled as of date (Hoffman 2003) and will therefore need to be collected and put into digital format before addition to the metric. Page 41 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Species Distribution, Density and Diversity – Data is lacking as to exactly where amphibian populations occur in Vermont. For this project, we could only consider wetlands and other habitat where amphibians may be likely to occur. It would be more helpful to know those locations where, indeed, they do occur. More Information On Locations of Vernal Pools - The state currently provides access to National Wetlands Inventory (NWI) wetland location maps for the state. However, NWI maps are developed at a 1:250,000 scale (Tuggle and Wilen 2003), and all of the data available carries a disclaimer regarding the inherent limitations of data developed at this scale (US FWS 2003). Essentially, in treeless areas, wetlands ¼ of an acre and larger are accurately depicted, in forested areas, the minimum wetland size depicted ranges between 1 and 3 acres (CGDB 2003, WGISC 2003). This means that although wetlands provide an ideal habitat for most amphibians, not all wetlands within the state will be included in the analysis of available habitat. This is especially true for small, temporary wetlands, such as vernal pools. We recommend the collection of data on the location of these important habitats. Global Populations - A lack of clear scientific understanding is the largest hurdle in addressing the global amphibian crisis. A majority of our current understanding is from the US, Europe, and Australia, all of which are areas of relatively low amphibian diversity (Houlahan et al 2000). The fact that largescale declines have been observed in many isolated and relatively undisturbed sites can be extrapolated to mean that numerous declines have and are occurring in poorly documented sites with high diversity (Young et al 2001). Latin America is a strong example of the lack of scientific understanding and the implications of this issue. Latin America contains over half of the world’s Page 42 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont amphibian richness, some declines have been documented, but it is estimated that 5-10% of the species in Latin America are undocumented. Because of this it is impossible to fully understand and quantify population declines (Young et al 2001). Cause of Global Decline - Current research has found numerous sources of local population mortality, but a clear global causative agent is lacking. Due to the sensitive nature of amphibian species it is possible that a host of different stresses are working synergistically to harm populations globally. Further research is needed to document all amphibian populations, particularly those in the lesser studied areas of high diversity. Many of the studies suggest that protecting habitat is only a beginning due to the insidious nature of some of the threats such as UV-B radiation. The fact that many of the identified causative agents can not be easily reversed emphasizing the importance of identifying and remediating all other mechanisms such as Atrazine to reduce the stresses on amphibian populations and reduce the risk of global extinctions Long-term recommendations for VTRANS INTRO PARAGRAPH Larger Culverts - Our research has indicated that most amphibians found within the state are likely to move through larger culverts, as they allow more natural light to pass through. They are also more conducive to the long-term maintenance of natural ground cover throughout, providing further appeal to amphibians. Finally, it should also be noted that larger culverts are more likely to have room to allow for both water flow and amphibian passage (Andrews 2003). Focus on Rare, Threatened, and Endangered Species - This is not only for environmental purposes but political purposes as well; amphibians carry a public stigma and thus funds will be more readily available to develop Page 43 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont crossing alternatives if rare or threatened species are at the forefront of the issue (Andrews 2003). Focus on Areas with High Species Density and/or Diversity - It makes the most sense, both in terms of environmental conditions and economic conditions, to first consider sites that are surrounded by a high density and diversity of amphibians. This idea plays into the policy of getting the “most bang for your buck”. Addition of Raised Structures Within Culverts with Regularly Flowing Water - While most amphibians require a moist environment, a culvert with consistently flowing water would not provide an ideal crossing environment. The state should consider installing culverts with modifications that would allow the amphibians to bypass the water flow (i.e. culverts with raised centers, designed to provide a transport route while directing channel flow along either side of the route). Replacement of Culverts on an As-Needed Basis - Logically, it is most cost-effective to replace culverts only when they need to be replaced due to structural damage to either the culvert itself or the road above it. Therefore, we recommend that the final culverts selected to serve as amphibian crossings be replaced on an “as-needed” schedule (unless ecologic conditions dictate replacement sooner), to reduce costs to the state and ultimately, to taxpayers. Focus on Sites with Documented Road Mortality - Since there have been documented amphibian mortality problems at these sites, it would be helpful to both motorists and the environment to take action. Focus on Sites with High Traffic Intensity - If necessary (if the selection processes described above yield a high number of potential sites), traffic volume data may be used to further prioritize sites. If this is the case, we suggest that the above prioritizations be followed, but within each category, Page 44 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont those sites with the largest amount of traffic volume be considered for action first. Adaptive Management - Finally, as with any project designed to address specific environmental problems, the practice of adaptive management will need to be implemented to garner the most successful results. The true outcome of a project can never be known until it has been implemented in the real world; if problems or complications arise, it will be necessary to go back to the original plans and alter them accordingly. GO MORE INTO ADAPTIVE MANAGEMENT Page 45 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont References Ashley, E. P., and J. T. Robinson. 1996. Road mortality of amphibians, reptiles, and other wildlife on the Long Point Causeway, Lake Erie, Ontario. Canadian Field Naturalist 110:403-412. Bartlet, P. E. 2000. A biophysical analysis of habitat selection in western toads (Bufo boreas) in southeastern Idaho. Unpublished Ph.D. dissertation, Idaho State University, Pocatello. Bayliss, P. 1995. 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VT Center for Geographic Information. <http://www.vcgi.org/dataware/default. cfm?page=./search_tools/search_action.cfm&query=theme&theme=018&layers_startrow=41> Croft, Jonathan. 2003. VTRANS Highway District Boundaries. VT Center for Geographic Information. <http://www.vcgi.org/dataware/default.cfm?page=. /search_tools/search_action.cfm&query=theme> Hoffman, Nelson. 2003. Personal Communication. Meeting. October 21 Mitsch, William J. and Gosselink, James G. 2000. Wetlands (Ed. 3). New York: John Wiley and Sons, Inc. North Carolina Corporate Geographic Database (CGDB). 2003. National Wetlands Inventory. <http://www.cgia.state.nc.us/cgdb/nwif.html#quality> Sharp, Steve. 2003. VT Bridges and Culverts: Transportation Structures. VT Center for Geographic Information. <http://www.vcgi.org/dataware/default.cfm?page=./ s earch_tools/search_action.cfm&query=theme&theme=018-&layers_startrow=41> Sutter, GW II. 1993. A Critique of Ecosystem Health Concepts and Indexes. Environmental Toxicology and Chemistry. 12:1533-1539 Tuggle, Benjamin N. and Willen, Bill O. 2003. National Wetlands Inventory Map Products. <http://wetlands.fws.gov/products.htm> U.S. Fish and Wildlife Service. 2003. National Wetlands Inventory Map Data. VT Center for Geographic Information. <http://www.vcgi.org/metadata/Water Wetlands_NWI.htm> VT Fish and Wildlife Service. 1997. Rare, Threatened, and Endangered Species & Significant Communities. VT Center for Geographic Information. <http://www. vcgi.org/dataware/default.cfm?page=./search_tools/search_action.cfm&query=theme> Wyoming Geographic Information Science Center. 1997. National Wetlands Inventory. <http://www.wygisc.uwyo.edu/clearinghouse/metadata/nwi.faq.html> Contact’s List Page 51 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont Relevant appendices Page 52 Reducing Amphibian Mortality on Vermont Roadways Prepared by students of NR 206 - University of Vermont