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Amphibian Populations of Acadia National Park: Patterns and Processes in Relation to Anthropogenic and Natural Changes in Wetland Ecosystems

Proposed Start Date: October 1 1997

Proposed End Date: September 30 2001

Proposal To:

US National Park Service Office of Scientific Studies New England System Support Office Boston, MA

Submitted by:

Cooperative Park Studies Unit, University of Maine Biological Resources Division U.S. Geological Survey Orono, ME 04469

Project Officer: Allan O'Connell, Leader, Cooperative Park Studies Unit, University of Maine

Investigators: Malcolm Hunter, Professor of Wildlife Ecology, University of Maine William Glanz, Associate Professor of Zoology, University of Maine Aram Calhoun, Assistant Research Professor of Wetland Ecology, University ofMame

Project Number: NARO-N-OOl.OOO Title: Status and Monitoring of Amphibian Populations in Northeast National Parks

1 Project Number: NARO-N-OOl.OOO Project Title: Status and Monitoring of Amphibian Populations in Northeast National Parks

Proposal Title:

Amphibian Populations of Acadia National Park: Patterns and Processes in Relation to Anthropogenic and Natural Changes in Wetland Ecosystems

ABSTRACT

We propose to study how the dynamics of amphibian populations in Acadia National Parkare affected by Ilatura! changes (e.g. beaver dams and ecological succession) and anthropogenic changes (road building, path construction, and beaver and fire management) in wetland ecosystems. _Within _major wetland classes, we will measUre the patterns of amphibian species distribution and abundance in relation to wetland vegetation, fish populations, soil type, basic water chemistry, and physical parameters (water depth, temperature, hydrogeomorphic setting). We will conduct concurrent studies of amphibian reproductive success, focusing on those variables most influential in analyses of abundance patterns. Wetland types will be sampled over a range of disturbance and successional conditions, allowing us to investigate whether differences in amphibian population dynamics among sites can be explained by wetland attributes driven by varying disturbance histories.

INTRODUCTION

_A primary management function of the National Park System is the conservation ofnafilral biodiversity. Amphibians are important components of many forest and wetland ecosystems in the United States. In eastern forests, for example, they can be the dominant vertebrate group in terms of both abundance and biomass (Merchant 1972~ Burton & Likens 1975). As their life histories usually include both aquatic and terrestrial habitats, amphibians may be significant members of several adjacent ecological communities, contributing both species diversity and ecological interactions to each (Gibbons 1988). Many national parks have been created to preserve such communities; _species inventories, monitoring, and research, therefore, are needed to aid park management in this conservation role.

2 Recent studies, however, have noted major declines in amphibian numbers at numerous sites in a variety of habitats (Barinaga 1990, Wake 1991, Griffiths & Beebee 1992). Potential causes of declines include various human impacts such as habitat loss (Griffiths & Beebee 1992, Blaustein et al. 1994b, DeMaynadier & Hunter 1996), acid precipitation (e.g. Freda 1986, Horne & Dunson 1994) heavy metals (Freda 1991), global warming, and UV-radiation increases (Blaustein et al1994a and 1994b). Pechmann and Wilbur (1994) urged caution in interpreting such declines; they note that few studies have sufficient long-term data on abundance to distinguish significant declines from natural population fluctuations. Pechmann et al (1991) documented dramatic density fluctuations over 12 years in four amphibian species at one site with no evidence for long-term declines. Clearly, long-term monitoring and detailed studies will be needed to firmly test whether human activities are causing major decreases in the abundances of amphibian species.

Many national parks in the United States are suitable locations for studying long-term trends in amphibian populations. Parks preserve large areas of suitable habitats, eliminating large-scale habitat loss as a potential influence on local amphibians. Such areas, nevertheless, are subject to atmospheric chemical inputs from outside their boundaries, and to sometimes dramatic habitat alterations by human visitors, by other wildlife species, and by park management activities. By detailed studies of amphibian population trends at park sites varying in human disturbance and ecosystem chemistry, a clearer picture ofiong-term processes affecting amphibian populations would emerge. In this propo~ we describe a set of studies to test the effects of anthropogenic and natural . ecosystem changes on amphibian populations in Acadia National Park, Maine.

BACKGROUND

Amphibians of A9adia National Park

Fourteen species of amphibians have been recorded on Mount Desert Island, six salamanders and eight anurans (Table 1). Based on the known geographic ranges of Maine's amphibians (Hunter et al. 1992) there is only one other Maine species likely to occur on MDI, the blue-spotted salamander Ambystoma Iaterale and its polyploid hybrids; It is unlikely, but possible, that two other species, the spring salamander Gyrinophilus porphyriticus and the mink frog Rana septentrionalis occur there.

Each ofthese species has a distinct set of habitat requirements (Hunter et al. 1992, Stockwell and Hunter 1989), but it is possible to make some broad predictions about the potentialimpacts of environmental change. In particular, it is likely that the loss or modification of wetlands would be of prime importance, given that all of these species; with the exception of red-backed salamanders, are linked to freshwater wetlands (broadly defined) at some point during their life cycle. Because of this relationship, we have included a brief overview of the status of wetlands on MDI based on two sources: National Wetland Inventory (NWI) maps, recently updated by the U.S. Fish and Wildlife and a recent monograph, "The wetlands of Acadia National Park and vicinity" (Calhoun et al. 1994).

3 Table 1: AmphiBian species recorded on Mount Desert Island.

Spotted Salamander Ambystoma maculatum EastemNewt Notophthalmus viridescens Dusky Salamander Desmognarhus fuscus Red-backed Salamander Plerhodon cinereus Four-toed Salamander Hemidacrylium scutatum Two-lined Salamander Eurycea bislineata - American Toad Bufo americonus Spring Peeper Pseudacris clUcifor Gray Treefrog Hyla versicolor Bullfrog Rona catesbeiona Green Frog Rona clamitons Wood Frog Rona sylvatica Leopard Frog Rona pipiens Pickerel Frog Rona palustris

Wetlands of Acadia National Park

Marine aquatic beds, intertidal shellfish flats, salt marshes, freshwater marshes, forested wetlands, and peatlands provide co~ and definition to the otherwise relatively rugged landscape of Mount Desert Island. Acadia National Park has 12,846 hectares (31,730 acres) of wetland area (this number does not include wetlands in the Marine and Riverine Systems that app~ar as linear features on wetland maps or wetlands smaller than the minimum mapping unit of 0.3 hectares). Eighteen percent of all land located within ANP is classified as wetland with another 11 % in fee ownership lands and 5% in lands held under conservation easements.

The majority ofweflands fall within two systems: Marine (36%) and Palustrine (32%). Aquatic beds (69%) and areas lacking vegetation (e.g., unconsolidated bottom, rocky shore)(31%) constitute the Marine System. Palustrine wetlands (freshwater wetlands with persistent vegetation such as sedge meadows, cattail marshes, or wooded swamps) are dominated by forested or scrub-shrub wetland· communities (86%). Ofthe 9000 wetland units mapped by NWI in the region; more than 40% were palustrine. The majority of palustrine wetlands are <0.5 hectares in size. Wetlands impa~ted by -... beavers, fire, and road construction fall largely within the Palustrine System.

Impacts on Amphibians _ -

In recent years probably the greatest impacts on the freshwater wetlands ofMDI have come through changes in hydrology, specifically the blocking or unblocking of drainages because of the activities - of beave is and humans. Humans commonly construct roads that have no culverts or inadequate culverts, thus impounding runoff water upslope and making downslope sites drier. Beaver dams convert streams and the various wetland and upland ecosystems that border the streams into a particular kind of wetl~d (usually a deep marsh) and after the dam is abandoned these marshes succeed to wet meadows, shrub swamps, forested wetlands, and other ecosystems. These changes

4 have many ramifications for the entire ecological community of such sites (Brown et al 1996, Dubuc et a11990, Grover 1993, Naiman et al1986, 1988), including the amphibians.

Habitats for some amphibian species may be created, thus benefitting those species. Where roads block a large volume of surface runoff, vernal pools may form and be used as breeding sites by certain species, most notably wood frogs and spotted salamanders. The flowages created by beavers are likely to be breeding sites for several species (eastern newt, bullftog, green frog, American toad, spring peeper, gray treefrog, leopard frog, and pickerel frog) and year-round sites for the first three species named (Hunter et al 1992, Grover 1993).

Conversely, these hydrologic changes could mean habitat loss or degradation for many species. Damming streams will eliminate some upstream habitat for two stream-dwelling species, the dusky and two-lined salamanders, and will modifY their habitat downstream of the dam by changing water flows and temperatures. Inundation by a dam may also decrease the availability of some other key habitats: forested wetlands that harbor spring peepers and gray tree frogs during much of the year; wet meadows that are used by leopard and pickerel frogs; upland forests that are home to red-backed 'salamanders; and vernal pools that provide a fish-free refuge for breeding spotted salamanders, wood frogs, gray tree frogs, spring peepers, American toads, and four-toed salamanders. The loss of surface water runoff due to blockage by a road could dry out downslope wetlands and could even degrade the habitat of red-backed salamanders which require a moist soil litter for foraging activity (Heatwole 1960).

Sorting out the precise mechanisms by which hydrologic changes can lead to habitat loss, addition, and modification may be fairly complex. Beyond the gross and obvious changes, such as a stream becoming a pond, there may be a variety of other biological (e.g. presence of fish), chemical (e.g. pH), and physical (e.g. water temperature) changes that could profoundly affect habitat suitability for a particular species (Home and Dunson 1994, Hecnar and M'Closkey 1996). For example, if a vernal pool near a stream were flooded by a: beaver dam and this allowed fish to invade, the breeding success of amphibians might plummet due to fish predation on eggs and larvae.

Much ofthe preceding discussion implies thatthe effects of beavers on amphibian habitats is an issue . that merits concern. However, given that the construction and destruction of beaver dams are Datural processes, park managers might be inclined to ignore their impacts on wildlife. Nevertheless, there are some arguments for at least an assessment of their impacts on amphibians. First, humans have directly manipulated beaver populations on MDI over the last century, first by eliminating them from the island and then subsequently reintroducing them. Indirect effects on beavers have also been dramatic. The removal of wolves from the island and prohibitions on fur-trapping left beavers without two predators that might tend to regulate populations. The human-set 1947 fire created an extraordinary surfeit of food for beaver, notably aspen, and subsequent succession has begun to reduce that food supply (Dubuc et al 1990). Finally, the extensive construction of roads has given beavers many opportunities to create flowages by blocking a small culvert on sites where a typical beaver dam would be difficult to construct and maintain. After these road-side dams are built, it is

5 a human decision whether or not they will be allowed to stay. In short, humans affect the waxing and waning of beaver populations in many ways and thus need to be concerned with the consequences these fluxes may have for amphibians. Moreover, this is not a problem unique to Acadia National Park; beavers are one of the most widespread and ecologically important mammals in temperate North America (Naiman et al 1988, Smith et al 1991 , Jones et al 1994); managing their populations is an issue in parks and other lands across the continent (e.g. McCall et al1996).

- Other human impacts;

Anthropogenic pollution at Acadia National Park has been of concern to park managers for many years (Anderson 1984). Although Anderson warned of potential negative effects of pollution on amphibians at ANP, subsequent research (see review by Freda 1986, 1991) has underscored the absence of any clear universal patterns. For example, the effects ofacid precipitation-are mediated by complex interactions with heavy metals and the responses vary enormously among different species (Horne and Dunson 1994).

DESCRIPTION OF RECOMMENDED PROJECT

Objectives: -

We have divided our objectives into two sets that are likely to correspond to two graduate student projects focusing on patterns and processes. The principal objective of the patterns work will be to describe the current array of amphibian habitats, reconstruct how they have probably changed during­ the last sixty years, and predict how they may change in the future under various disturbance regimes. This wil,Ilnvolve first determining the current distribution of amphibians in various types of ecosystems (principally palustrine wetlands and streams) and ecosystem conditions (e.g. wetlands with and without significant human or beaver modifications, etc). Secondly, this will require - - doeumentingthe current and past distributions of various wetland types and conditions u_sing aerial photographs and other historical records. Finally, we plan to relate the ciJrrent distributions of amphibian species to historical records, principally Manville (1939) and Davis (1958), to look for --- - patterns of extinctions and colonizations that might be informative. We predict that physical characteristics of the habitat (e.g. wetland type, hydrology, water temperature) will be the- best - predictors of breeding distributions of amphibian species, with certain biological factors (e.g. presence of predatory fish limiting wood frogs or spotted salamanders) also being important for­ certain species. Alternate hypotheses also will be evaluated, including the importance of historical factors (e.g. species presence in the watershed in previous surveys) and oflandscape features (e.g~ - dry environments limiting species intolerant of dessication, distance to other appropriate wetlands limiting leopard frog recolonization, presence of beaver flowages upstream limiting stream salamander species).

The major objective of the second graduate student project will be to understand the processes of

6 po.pulatio.n dynamics and habitat selectio.n that lead to. the Db served distributio.n patterns. Primarily this will invo.lve an attempt to. do.cument the relative breeding success o.f each species in the vario.us types and co.nditio.ns o.f habitat o.utlined abo.ve. In practice we will need to. be selective, fo.r it will no.t be feasible to. do.cument breeding success fo.r every species in every type and co.nditio.n o.f habitat in which it o.ccurs. Ano.ther impo.rtant o.bjective o.f this wo.rk will be trying to. understand the environmental facto.rs(e.g. presence Dr absence o.ffish, water chemistry, water temperatures) that shape breeding success and ultimately distributio.ns. We hypo.thesize, fo.r example, that water temperatures will best predict larval gro.wth and develo.pment rates fo.r mo.st species, but that presence o.fpredato.ryfish will depress larval survival o.fvulnerable species, such as wo.o.d fro.gs and spo.tted salamanders, thereby limiting o.verall reproductive success o.fthese species.

Metho.ds:

To. describe the general distributio.n and abundance o.f streams and palustrine wetlands o.n MDI, and ho.w they have changed o.ver the last fifty years, we will use aerial pho.to.graphy spanning fro.m 1953-1991 (so.me inco.mplete 1930s and 40s pho.to.graphy is alSo. available), NWI maps fo.r Acadia Natio.nal Park (1991), and vegetatio.n co.ver type maps co.mpleted by Gary Wagner in 1979. A GIS wetland and watershed database at the Park will allo.w us to. query wetland types by classificati"o.n and identify distributio.n and abundance patterns. Mo.re impo.rtantly, we will be able to. attribute changes in wetlands due to. the co.nstructio.n o.f ro.ads and beaver dams because these are usually easy to. see o.n aerial pho.to.s.

Assessing the distributio.n o.f amphibians in vario.us wetlands by eco.system type and co.nditio.n will require a" diversity o.f appro.aches. The first task will be selecting a stratified rando.m sample o.f wetlands fo.r detailed field wo.rk. Based o.n o.ur current understanding o.f amphibian habitats o.n MDI we wo.uld stratify o.ur sample as sho.wn in Table 2. These 5-year increments are based o.n o.ur estimates o.f the rates o.f successio.nal change fo.llo.wing impo.undment and the breaching o.f impo.undments.

Table 2. Amphibian habitats we expect to. sample o.n Mo.unt Desert Island (indicated by +) subject to. examining data o.n current and past distributio.ns o.f palustrine wetlands and streams.

No.t Iml2Qynde:d Cyrre:ntI~ Iml2Qynd~d Iml2Qyndm~nt In Past 50 yrs < 5 yr >5 yr Breached past 5 yrS Streams + + + + Fo.rested Wetland + + Shrub Swamp + + + + Wet Meado.w + + + Marsh + + + +

7 For each of these strata we will randomly pick at least ten sites, ideally 30 if there were enough sites and time to sample them all. We would prefer to work solely within Acadia National Park but it may be necessary to seek permission to work on some private land to have a balanced and complete sample. We will make a special effort to have several sites in the watersheds selected -for studies of atmospheric deposition effects (IS. Kahl and K. Tonnesson pers. comm., Kahl and Fernandez 1996), evenifit means overriding the random selection process.

In addition to these sites, we will undertake some ad hoc sampling at additional sites for which there is historical information, in order to detect any extinction or colonization events that might be of special interest. For example, the leopard frog and the American toad were recorded by Davis (1958) at a variety of sites across !vfDI; both species have now disappeared from the eastern half of the island (B. Connery, pers. comm). The reasons for these local extinctions may lie in either successional change or disturbance by humans and/or beaver. Th_e very localized distributions of some species on.MDI, such as the gray treefrog and the four-toed salamander, may also pe related to effects of humans or beavers, through habitat fragmentation or other habitat-alterations.

Detennining the distributions of amphibian species will require a variety of methods appropriate for the taxa present (Heyer et al. 1994). We will not describe these methods in great detail here; suffice it to say that we will use standardized methods as much as possible with special reference to the USGS Biological Resources Division's protocols for the North American Amphibian Monitoring Program (Available on the internet at http://www.im.nbs.gov).

To sample the relative abundance of amphibian species across habitats, we will employ multiple techniques, as different species show gross differences in detectability (e.g. active, vocal frogs vs. cryptic red-backed salamanders) and each species varies seasonally in abundance and detectability (e.g. breeding adult frogs compared to larval stages). Furthermore, year-to-year and site-to-site variation in phenology also argue for multiple sampling methods. For most-species breeding in­ wetlands we inten~ to obtain indices of abundance for breeding adults, egg masses, early tadpoles, ana larvae at metamorphosis. Ifthese indices consistently reflect abundance, ranks of abundance for different species should be correlated within a habitat. For example, jf in deep marsh.es all- four indices show species A > species B > species C, we can have confidence in that order of-relative·· abundance. We will test these relationships with appropriate non-parametric correlation statistics.

The most widely applicable field approach to determine species occurrence and abundance will be transect surveys. Within a particular habitat these would be of a standard length and width and, as far as possible, time. For example, in sampling streamside salamanders we will probably use 2 x 10 m transects with about two minutes per transect, but for marsh frogs we would use 2 x 50 m· transects with 12 minutes per transect. For some species conducting a transect search will involve more than just walking and visually searching; for example, turning over logs for red-backed salamanders and dip-netting for eastern newts. A separate set of surveys may be conducted for different life stages: e.g. eggs, larvae, and adults of ranid frogs. Differences in detectability, of course, will severely limit our ability to compare among species and perhaps among habitats.

8 However, many of our key compansons will be between the same species in similar habitats (e.g. green frogs in recently created marshes versus older marshes) and should be reasonably free of bias.

Simple presence/absence information is also of use and much easier to obtain. In particular, for breeding male anurans we can readily determine presence-absence with calling surveys conducted during evening visits at the appropriate times of year. For all species we will record all incidental encounters and this information, combined with records of the total amount of time spent at each site, will be useful in assessing distribution and abundance.

Understanding the interface between habitat selection and population dynalnics will require studying . the reproductive success of certain species in different habitats. For some species this will be reasonably straightforward; for others with eggs or larvae that are difficult to census, it will not be possible to obtain a robust estimate of reproductive success. We will be assessing relative abundances and reproductive success of species among sites using a redundant measures approach. The most direct method of measuring reproductive investment will be counting and monitoring egg masses and estimating their size for the species with conspicuous aquatic egg masses: i.e., the ranid frogs, spotted salamander, and American toad. This simple method may reveal significant differences in reproductive investment in different habitats. Similarly, dip netting and larval traps (funnel traps constructed from 2 liter plastic bottles) will allow us to systematically estimate the densities of all species that have aquatic larvae in lentic environments (all but the red-backed, two-lined, and dusky salamanders). Repeated measurements of larval body lengths and weights will allow us to estimate growth rates and time to metamorphosis, another measure of the suitability of a site for amphibian reproduction. Combining these measures with a vocal index and a transect index (described above.) will decrease error associated with sampling techniques and validate our estimates of both relative abundance and reproductive success. If the majority of indices for a given species shows the same relationships among habitats, as revealed by non-parametric rank statistics, we can consider the rankings of habitat quality to be robust.

For some ofthese parameters (hatching success, growth rates, and time to metamorphosis) we will obtain additional estimates under more controlled conditions by placing egg masses in enclosures. To further elucidate the role of predation on amphibian eggs and larvae, these enclosures will also serve as predator exclosures. Presence of potential predators (e.g. fish, predaceous diving beetles, caddisflies) will be surveyed using the funnel traps and dipnets described above, existing data (primarily for fish in larger bodies of water), and visual surveys for presence/absence.

The study sites will be characterized by wetland class according to the Cowardin et al (1979) classification system, vegetation physiognomy, and species composition and distribution. Hydrologic characteristics, including hydrologic regime (timing, depth and duration of flooding), major inputs and losses of water, depth characteristics (particularly in wetland basins), and degree of connectivity to other wetlands, will be described.

Given the expense of exhaustive water chemistry measurements and the generally low correlations

9, between amphibian distribution and water chemistry found by Hecnar and M'Closkey (1996), we plan to measure only a limited suite of such parameters in our work: pH, DO, DOC, nitrate-nitrogen, total phosphorus, conductivity, and turbidity. We will also relate our findings on ~he atmospheric deposition watersheds to other chemical parameters available from that study (Kahl and Fernandez 1996). We will document our methods and study sites carefully, so that future researchers will be able to repeat our surveys and assess whether or not long-term populations changes have occurred_

STUDY PRODUCTS

- We plan to have two graduate students enrolled in the winter of 1998 and reaay for three field seasons, 1998, 1999, 2000, with each finishing by summer 2001. The prirnary products of this study will be the theses of these two graduate students. Both theses are expected to be completed by September 2001. They will include chapters written as journal articles, each focusing on a major· . ()bjective ofthis study, which will be submitted for publication in appropriate ecological and wetland journal-so Each thesis will also include numerous appendices with information on methods and observational data which may not be appropriate in a journal paper, but yet may be useful for future research and monitoring. . These will include detailed descriptions and evaluations of methods employed,-multiyear phenological data on breeding, growth and larval development by amphibian species, and tabular summaries of distributional data and changes in wetland status by site. Identification aids may also be included.

PERSONNEL

Allan O'Connell is a research wildlife biologist and Leader of the Cooperative Park Studies Unit at the University of Maine. He has directed National Park Service natural resource management programs in New York and Maine, and has conducted research on the ecology of Lyme disease, white-tailed deer, coyotes, and spruce grouse in eastern national parks. His work has focused on population changes in relation to habitat fragmentation and landscape variability.

Malcolm Hunteris Professor ofWl1dlife Ecology and the Libra Professor of Conservation Biology at UMaine and is involved in a wide array of research on vertebrates, invertebrates, and plants. With respect to amphibians he has been the principal director of the Maine Amphibian and Reptile Atlas Project, senior editor of The Amphibians and Reptiles ofMaine, and faculty advisor to five graduate students studying the ecology of Maine amphibians. Also of relevance to this project are his s1!Idies· ·on the effects of acid precipitation on wildlife and the animal communities of vernal pools and peatlands.

William Glanz is Associate Professor of Zoology and Cooperating Associate Professor of Wildlife Ecology at the University of Maine. His research includes a variety of projects on the ecology and

10 behavior of vertebrates, including population ecology and habitat use of rodents and bats, and the foraging behavior of birds and mammals. He has directed two projects in Acadia National Park, a biological inventory of wetlands on the Schoodic Peninsula and a study of community composition and seasonal habitat use by bats in the park.

Aram Calhoun has an extensive background in wetland science, formerly as a biologist for the US Natural Resource Conservation Service, and currently as a wetland ecologist at the University of Maine. Major publications include a USFWS monograph on the ecology of red maple swamps in the glaciated northeast and senior authorship of a U.S. Fish and Wildlife Service 1994 publication, "The wetlands of Acadia National Park and vicinity" and contributing author to the "Acadia National Park Water Resources Management Plan." Ajoint project with the New Hampshire Charitable Fund and the Maine Audubon Society on vernal pools has produced a manual entitled" A Maine Citizen's Guide to Locating and Describing Vernal Pools." She is currently working with the Maine Audubon Society and the Maine Department of Inland Fisheries and Wildlife as the Project Coordinator for the Maine Amphibian Monitoring Program and is Project Coordinator for a vernal pool study.

LITERATURE CITED

Anderson, I.L. 1984. Potential impact of acid precipitation and poor air quality on the resources of Acadia National Park. Boston, MA. US Dept. Interior, National Park Service, North Atlantic Region, OSS 85-1. 100 pp. Barinaga, M. 199p. Where have all the froggies gone? Science 247: 1033-1034. Blaustein, AR, P.D. Hoffinan, D.G. Hokit, J.M. Kiesecker, S.C. Walls, and J.B. Hays. 1994a. UV repair and resistance to solar UV- B in amphibian eggs: a link to population declines? Proc. Nat. Acad. Sci. USA 91: 1791-1795. Blaustein, A.R, D.B. Wake, and W.P. Sousa. 1994b. Amphibian declines: judging stability, persistence, and susceptibility of populations to local and global extinctions. Conserv. BioI. 8: 60-71. -Brown, D.l, W.A Hubert, and S.H. Anderson. 1996. Beaver ponds create wetland habitat for birds­ in mountains of southeastern Wyoming. Wetlands 16: 127-133. Burton, T.M., and G.E. Likens. 1975. Salamander populations and biomass in the Hubbard Brook Experimental Forest, New Hampshire. Copeia 1975: 541-546. Calhoun, AlK., I.E. Cormier, RB. Owen Jr., A.F. O'Connell Jr., C.T. Roman, and RW. Tiner Jr. 1994. The wetlands of Acadia National Park and vicinity. Univ. Maine Agric. Expt. Sta., Misc. PubI. 721. Cowardin, L.M., V. Carter, F. C. Golet, and E. T. LaRoe. 1979. Classification of Wetlands and Deepwater Habitats of the United States. Office of Biological Services, Fish and Wildlife Service, US Department of the Interior, Washington DC. Tech. Publ. FWS/OBS-79/31.

11 Davis, S.L. 1955-. Notes on the amphibia in Acadia National Park, Maine. M. Sc. Thesis, Cornell Univ., Ithaca, NY. DeMaynadier, P.G., and M.L. Hunter Jr. 1996. The relationship between forest management and amphibian ecology: A review of the North American literature. Environmental Review 3: 230-261. Dubuc, L.1., W.B. Krohn, and RB. Owen Jr. 1990. Predicting occurrence of river otters by habitat on Mount Desert Island, Maine. 1. Wildl. Manage. 54: 594-599. _. Freda, J. 1991 .. The effects of aluminum and other metals on amphibians. Environ. Pollution 71: 305-328. Freda, 1. 1986. The influence of acidic pond water on amphibians: a review. Water, Soil and Air Pollution 30: 439-490. Gibbons, 1. W. 1988. The management of amphibians, reptiles, and small mammals in North America: The need for an environmental attitude adjustment. In Management of Amphibians,­ -Reptiles, and Small Mammals in North America. USDA Forest Service, General TecluUcal Report RM-166. _ Griffiths, R, and T. Beebee. 1992. Decline and faIl of the amphibians. New Scientist (27 June 1992): . 25":29. Grover, A. 1993. Influence of beaver on bird and mammal species richness within wetlands of different sizes in south-central New York. M.S. Thesis, State Univ. of New York, Syracuse. Heatwole, H.-1960. Burrowing ability and behavioral responses to dessication of the salamander, Plethodon cinereus. Ecology 41: 661-668. Hecnar, S.1., and R T. M'Closkey. 1996. Amphibian species richness and distribution in relation to pond water chemistry in south-western Ontario, Canada. Freshwater Biology 3-6: 7-15. Heyer,W.R, M.A Donnelly, R W. McDiarmid, L.C. Hayek, and M.S. Foster. 1994. Measuring and . Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Wasffington, DC. 364 pp. Home, M.T., and W.A. Dunson. 1994. Exclusion of the Jefferson Salamander, Ambystoma jeffersonianum, from some potential breeding ponds in Pennsylvania: effect of pH, temperature: and metals on embryonic development. Arch. Environ. Contam. Toxicol. 27: 323-330. Hunter, M.L., Jr., IAlbright, and J. Arbuckle. 1992. The Amphibians and Reptiles of Maine. Maine Agricultural Expt. Sta. Bull. 838: 188 pp. I

12 Naiman, RJ., C.A. Johnston, and J.e. Kelley. 1988. Alteration of North American streams by beaver. Bioscience 38: 753-762. Naiman, RJ., J.M. Melillo, and J. E. Hobbie. 1986. Ecosystem alteration of boreal forest streams by beaver (Castor canadensis). Ecology 67: 1254-1269 .. Pechmann, J.H.K., and H.M. Wtlbur. 1994. Putting declining amphibian populations in perspective: natural fluctuations and human impacts. Herpetologica 50: 65-84. Pechmann, J.H.K., D.E. Scott~ RD. Semlitsch, J.P. Caldwell, L.J. Vitt, and J.W. Gibbons. 1991. Declining amphibian populations: the problem of separating human impacts from natural fluctuations. Science 253: 892-895. Smith, M.E., C.T. Driscoll, B.J. Wyskowski, C.M. Brooks, and C.S. Cosentini. 1991. Modification of stream ecosystem structure and function by beaver (Castor canadensis) in the Adirondack Mountains, New York. Can. J. Zoo1. 69: 55-61. Stockwell, S.S., and M.L. Hunter Jr. 1989. Relative abundance ofherpetofauna among eight types of Maine peatland vegetation. J. Herpeto1. 23: 409-414. Wake, D.B. 1991. Declining amphibian populations. Science 253: 860.

BUDGET

Budget Justification:

Budgetary infonnation for this draft proposal is summarized in a composite three-year budget on the next page, and by individual year on the following pages.

Salaries for each principal investigator are based on 1 month each year by Calhoun, 0.5 month each year by Hunter and Glanz. These figures are slightly higher than in previous drafts, as they incorporate a 3% raise effective July 1997 and another effective July 1998, .as per collective bargaining agreements. Graduate assistantships are based on 12-month appointments for 2 Ph.D students. Each will have an undergraduate field assistant for the summer months. Fringe benefits are calculated as 30.5% of PI salaries.

Cost-sharing amounts by the University of Maine are given in the annual budgets (pp 15-16). Amounts are based on overhead cost-sharing rates as per cooperative agreement between NPS and UM, and 0.6 month salary cost-sharing by Hunter and Glanz during the first year, and 0.5 months each in the second and third years.

Housing and travel amounts are estimates based on a S-month work season for the graduate students on MDI, and round-trips between brono and MOl by the PI's. Equipment costs in the 1st yeat are for a GIS-capable PC workstation.

13 SUM:MAR.Y THREE-YEAR BUDGET (National Park Ser:vice - Sponsor Share Only)

Year I Year 2 Year 3 - Total

Salaries & Wages PI's 10183 10490 10490 31163

Two Grad Students 28000 28000 28000 84000

Two Field Assistants 6000 6000 6000 18000

Fringe Benefits 30.5% PI's 3106 3199 3199 9504

Housing - 2800 2800 2800 8400

Travel 3450 3450 3450 10350

Supplies 3000 3000 3000 9000 Computer Equipment 3000 3000

Total Direct Costs 59539 56939 56939 173417

Indirect-costs NPS - 13396 12811 12811 39018 (22.5% of direct costs)

Total-NPS 72935 69750 69750 212435

14 THIRD YEAR BUDGET CFY 2000)

NPS Sponsor Share UM Cost-Share Total

Salaries & Wages Calhoun 4715 (l mo.) Hunter 3242 (0.5 mo.) 3242 Glanz 2533 (0.5 mo.) 2533 - PI's total 10490 5775 16265 GradAssts 28000 28000 Field Assts 6000 6000

Fringe Benetits 3199 1761 4960

-Housing 2800 2800 Travel 3450 )450 Supplies 3000 3000 Total Direct 56939 7536 64475

Overhead-NPS 12811 12811 Overhead-UM 128Il 12811 OH-PI's costshare X 45% 3391 3391

Totals 69750 23738 93488

SUMMARy THREE-YEAR BUDGET CFY 1998-2000)

NPS Sponsor Share UM Cost-Share Total

Salaries & Wages PI's total 3Il63 18277 49440 Grad Assts - 84000 84000 Field Assts 18000 18000

Fringe Benefits 9504 5574 15078

-Housing 8400 8400 Travel 103~0 10350 Supplies 9000 9000 Computer Equipment 3000 3000 Total Direct 173417 23851 197268

Overhead-NPS 39018 39018 Overhead-UM . 39018 39018 OH-PI's costshare X 45% 10733 10733

Totals - 2-12435 73602 286037

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