Small Mammal Survey of the Nulhegan Basin Division of the Silvio 0. Conte NFWR and the State of Vermont's West Mountain Wildlife Management Area, Essex County Vermont • Final Report March 15, 2001 C. William Kilpatrick Department of Biology University of Vermont Burlington, Vermont 05405-0086

A total of 19 species of small mammals were documented from the Nulhegan Basin Division of the Silvio 0. Conte National Fish and Wildlife Refuge (NFWR) and the West Mountain Wildlife Management Area Seventeen of these species had previously been documented from Essex County, but specimens of the little brown bat (Jr{yotis lucifugu.s) and the northern long-eared bat (M septentrionalis) represent new records for this county. Although no threatened or endangered species were found in this survey, specimens of two rare species were captured including a water shrew (Sorex palustris) and yellow-nosed voles (Microtus chrotorrhinus). Population densities were relatively low as reflected in a mean trap success of 7 %, and the number of captures of two species, the short-tailed shrew (Blarina brevicauda) and the deer mouse (Peromyscus maniculatus), were noticeably low. Low population densities were observed in northern hardwood forests, a lowland spruce-fir forest, a black spruce/dwarf shrub bog, and most clear-cuts, whereas the high population densities were found along talus slopes, in a mixed hardwood forest with some • rock ledges, and in a black spruce swamp. The highest species diversity was found in a montane yellow birch-red spruce forest, a black spruce swamp, a beaver/sedge meadow, and a talus slope within a mixed forest. The lowest species diversities were found in northern hardwood forests, a black spruce/dwarf shrub bog, and most clear-cuts. ·

Introduction

Two portions of land in Essex County, Vermont, recently purchased from the Champion Paper Company through the Vermont Land Trust have been designated as the Nulhegan Basin Division of the Silvio 0. Conte National Fish and Wildlife Refuge (NFWR) and the West Mountain Wildlife Management Area. Much of this area of Northern Highlands (Northeast Kingdom) is a moist, boggy, ·swampy area ofboreal forest (Johnson, 1998) that includes a number of low mountains, less than 3000 feet in elevation. The Nulhegan Basin of the Conte NFWR, administered by the U.S. Fish and Wildlife Service, consists of26,000 acres in Lewis and parts of Ferdinand and Bloomfield townships north of Route 105. This area includes the drainage of the three major branches of the Nulhegan River and a number of bogs. The West Mountain Wildlife Management Area (WMA), administered by the Vermont Agency of Natural.Resources, consists of

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22,100 acres in Ferdinand, Brunswick, and Maidstone townships south of Route 105. This area includes a number of ponds and bogs along Paul and Wheeler Streams. Thirty-six species of small mammals (i.e., weighing less than 500 g) are known from Vermont and include one species listed as endangered (Indiana bat), one listed as threatened (small-footed bat), and eight species considered rare (long-tailed shrew, pygmy shrew, silver-haired bat, red bat, hoary bat, southern bog lemming, rock vole and pine vole). Those species listed above as rare are known from fewer than 5 0 specimens collected from Vermont. Although little survey work for mammals has been reported to date from Essex County, 21 of the 36 species of small mammals known from the state have been documented from this county (Kirk, 1916, Osgood, 1938a; G9din, 1977; Chipman, 1994). The objective of this study was to survey the small mammal diversity in the newly established Nulhegan Basin Division of the Silvio 0. Conte National Fish and Wildlife Refuge (NFWR) and the West Mounta~ WMA. This survey was conducted by surveying wetland and upland sites that were characteristic of natural communities of this region including northern hardwood forests, red spruce-northern hardwood forests, black spruce swamps, dwarf shrub bogs, alluvial shrub bogs, and beaver meadows. In addition, specific habitats (hot spots), including swift-flowing streams, talus slopes and clear-cuts, associated with the occurrence of rare species of small mammals were surveyed. Bat activity and diversity was investigated by use of mist nets and acoustical monitoring.

Materials and Methods

• Between 7 June and 15 October 2000 small mammals were collected with Sherman live traps and Museum Special snap traps from 19 sites (Appendix I). Traps were baited with either rolled oats or peanut butter and were set in the evening and checked in the early morning. Most sites were trapped for three consecutive nights with 60 to 100 traps per night. Traps were set in lines with the traps spaced about 20 feet apart. The upland sites sampled included two northern hardwood forests, three red spruce-northern hardwood forests, a lowland spruce-fir forest, and a montane yellow birch-red spruce forest. Other sites sampled included a black spruce swamp, a black spruce/dwarf shrub bog, two dwarf shrub bogs, an alluvial shrub swamp, a beaver meadow/sedge marsh, four boulder fields (talus slopes) with varying degrees of canopy and slope, and three clear-cut sites in early successional stages. Additional trapping methods were employed in an attempt to obtain additional specimens of shrews. For water shrews, thirty Museum Special snap traps baited with peanut butter were tied to vegetation and set along the edge of swift -flowing streams at five sites. For other shrews, 10 pit traps set 5 feet apart with a connecting drift fence were established at six sites including a northern hardwood forest, a black spruce/dwarf shrub bog, two dwarf shrub bogs, an alluvial shrub swamp and a: clear-cut.

• 2 The presence of bats was surveyed by the placement of mist nets across logging roads, trails and streams and by acoustical surveys using the Anabat II receivers. Bats were captured using 38 mm mesh, 2-ply-50 denier black nylon (Avinet) mist nets of 6 m and 12 m lengths. Mist nets were oriented horizontally • over streams at six sites and along logging roads at four sites for a total 30 net nights. In addition, an acoustical survey was conducted on 18 nights either near the area being netted or at areas where netting was not possible. The acoustical surveys were conducted using :frequency-division (Anabat II) detectors to evaluate general bat activity and to compare species detected with the acoustical survey with species captured in the field. · In order to assess habitat diversity not solely based on species richness (number of species) but also on the abundance of each species (species eveness) the following diversity indices, which account for heterogeneity in species eveness, were calculated. Simpson's index (Krebs, 1989) was calculated by: . D=LJJP where D is Simpson's index arid pi is the proportion of species i in the community. This diversity index predicts the probability of collecting two organisms that are the same species. More commonly used is its complement, 1 - D, which is the probability of collecting two organisms that are different species. The Shannon-Wiener index is derived from information theory and was calculated by the following: · H' = -Ilpi)(IogiJJi) where H' is the Shannon-Wiener index of species diversity and pi is the proportion of the sample comprised of the ith species. This index is a measure of the amount of uncertainty in predicting the species of an individual chosen at • random. Voucher specimens ofrepresentative specimens (Table 1) of all species collected were prepared as museum skins and skulls and are deposited in the Zadock Thompson Natural History Collections of the University of Vermont. Tissues and ectoparasites were collected from all specimens prepared as vouchers. For all specimens of the genus Peromyscus captured both salivary and blood samples were collected. The blood samples have been sent to the CDC for testing for hantavirus and Lyme disease. Salivary samples were used to identify mice of the genus Peromyscus unambiguously to species using the genetic markers at the salivary amylase locus following the methods of Aquadro and Patton (1980) with the modifications of Kilpatrick et al. (1994).

Results

A total of 19 species of small mammals (Table 1) were documented from the Conte Refuge and the West Mountain WMA of Essex County, Vermont. In addition, nine species oflarger mammals were observed during the survey including: woodchucks (Marmota monax), beavers (Castor canadensis),

• 3 TABLE 1.--Small mammals collected in a survey of the Conte Refuge and the West Mountain Wildlife Management Area of Essex County, Vermont.

Taxon Number Captured Voucher Specimens Insectivora Talpidae Parascalops breweri 2 2 Soricidae Sorex cinereus 11 9 Sorex fumeus 4 4 Sorex palustris 1 1 Blarina brevicauda 4 4 Chiroptera V esp.ertilionidae Myotis lucifugus 10 2 Myotis septentrionalis 24 2 Rodentia Sciuridae Tamias striatus 4 2 Tamiasciurus hudsonicus 2 1 Glaucomys sabrinus 1 1 Muridae (Sigmodontinae) Peromyscus maniculatus 41 27 Peromyscus leucopus 1 1 (Arvicolinae) Clethrionomys gapperi 95 57 Microtus pennsylvanicus 13 10 Microtus chrotorrhinus 2 2 Zapodidae Zapus hudsonius 26 16 Napaeozapus insignis 63 28 Carnivora Mustelidae Mustela erminea 3 2 Mustela frenata 2 1 TOTAL 309 172

4 snowshoe hares (Lepus americanus), coyotes (Canis latrans), red foxes (Vulpes vulpes), black bears (Ursus americanus), stripped skunks (Mephitis mephitis), white-tailed deer (Odocoileus virginianus) and moose (Alces alces). • Small Mammal Diversity A total of 4305 trap nights were spent surveying 19 sites with an average trap success of7.0%. The effort of sampling at each site ranged from 70 to 392 trap nights (Tables 2-4) and the success varied from 0.8 in a lowland spruce forest to 21.6% among the large boulders of an old stream bed within a mixed forest. The effect of the time of year a habitat was sampled on both trap success and species diversity is shown in Figure 1. Both low trap success (1.9 %) and low species diversity (1 -D = 0.39) was / observed in northern hardwood forest (Table 2). Only five species of small mammals ( chipmunks, red squirrels, deer mice, red-backed voles, and woodland jumping mice) were captured or observed in this habitat (Table 2). The woodland jumping mouse was common in this habitat representing 55 % of the captures. A total of 410 pit trap nights produced no shrews in this habitat. Much higher trap success (7.9 %) and slightly higher species diversity (1 - v D = 0.45) was found in red spruce-northern hardwood forest (Table 2). Although only four species of small mammals ( chipmunks, deer mice, red-backed voles, and woodland jumping mice) were found, both the woodland jumping mice and the red-backed vole were relatively common in this habitat accounting for 26 and 57 % of the captures respectively. An additional site (U-7), trapped later in the season (October), produced higher trap success (12.0 %). Although two additional • species (northern flying squirrel and short-tailed weasel) were captured at this site, the estimates of species diversity·were similar to that found in other red spruce- northern hardwood forest (Table 2). The lowland spruce-fir forest had the lowest trap s_uccess (0.8 %) of any r/ site sampled. The species diversity estimates range from aD-1 of 0.46 to~ Shannon-Wiener index of 1.0 both of which are slightly below the average values for upland sites of 0.4 7 and 1.16 respectively. The montane yellow birch-red spruce forest had the highest species diversity for small mammals of any of the upland sites sampled (Table 2). A total of 9 species of small m_ammals were taken from upland sites (Table 2) but only the northern flying squirrel (Glaucomys sabrinus) was restricted to an upland site. Although the trap success in lowlands was similar to that experienced in the uplands, the species diversity was on average higher (Table 3). This increase in species diversity was the result of additional taxa of shrews and voles being present in thes~ lowland sites. The black spruce swamp produced both high trap V success and high species diversity, whereas the black spruce/dwarf shrub bog v produced low trap success and low species diversity (Table 3). Pit traps set for 100 pit trap nights in the black spruce/dwarf shrub bog failed to captured any additional species of small mammals. Both p.warf shrub bogs and the alluvial

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J... Cl) .0 E 1 4 :J C/'Jrn 1 4 z ,(1} (.) CJ) (.) 1 2 Q,) :J 1 2 Cf) (.) Cl) 0. 1 0 0. co 1 0 Cf) J... }- 8 a, ~ 8 ·•...... > 0 ··...... ·...: '·, ::::: ..... C: -<~.;~ ~s~: ca ca ...... ······...... 6 :J Q.) 6 ···,," ·,.," .:.-:::...... E ~ ··.. , ...... :.-::: ··-.... E ...... 4 4 ·,...... ' .·:;/:..... :J ··-...... ::::: (.) ··'-.::-- ...... ···•· ··-.... ·...... :-:.:· ...... 2 2 :-=~~...... '·<,- ...... , ...... • ...... '\._ ...... -::.~· ;·-.... _...... {-:{ ,>:· ......

Jun.e July August September October

. Fig .. 1. Seasonal effect on mean trap success

and cummulative species number f.:::<::;1-

• .., .- TABLE 2.--0verview of small mammals captured in tJllllll[d sites: MU-1 northern hardwoods (Conte Refuge), MU-2 northern hardwoods (West Mountain WMA), MU-3 red spruce-northern hardwoods (Conte Refuge), MU-4 red spruce-northern hardwoods (West Mountain WMA), MU-5 lowland spruce-fir forest (Conte Refuge), MU-6 montane yellow birch-red spruce forest (West Mountain WMA), U-7 mixed hardwoods (West Mountain

Taxon MU-1 MU-2 MU-3 MU-4 MU-5 MU-6 U-7 Sorex cinereus 1 1 Blarina brevicauda 2 Tamias striatus obs. 1 1 obs. Tamiasciurus hudsonicus 1 obs. obs. obs. obs. obs. Glaucomys sabrinus 1 Peromyscus maniculatus 2 6 1 2 1 Clethrionomys gapperi 1 19 5 7 13 Napaeozapus insignis 4 1 1 10 1 2 Mustela erminea 1 Total Trap Nights 210 260 259 282 240 260 150 Trap Success 1.9% 1.9% 10.4% 5.3% 0.8% 5.4% 12.0% Number Species 1 4 4 2 2 6 5 Simpson's Index D* 1.0 0.28 0.55 0.56 0.5 0.31 0.54 1-D** 0 0.78 0.45 0.44 0.5 0.69 0.46 Shannon-Wiener H'*** 0.0 1.46 1.19 0.92 1.0 2.14 1.39 obs. - midden or animal seen or heard (red squirrel) * = Probability of picking two organisms that are the same species. ** = Probability of picking two organisms that are different species. *** = Average degree of uncertainty in predicting the species of an individual chosen at random.

7 TABLE 3.- Overview of small mammals captured in lowland sites: ML-1 black spruce swamp (Big Swamp, Conte Refuge), ML-2 black spruce bog/dwarf shrub bog (Western Bog, Conte Refuge), ML-3 dwarf shrub bog (Ferdinand Bog, West Mountain • WMA), ML-4 dwarf shrub bog (Dennis Pond, West Mountain WMA), ML-5 alluvial shrub swamp (North Notch Shrub Swamp), ML-6 beaver meadow-sedge meadow- :marsh (Upper Yellow Branch Open Wetland Complex, Conte Refuge) Taxon ML-1 , ML-2 ML-3 ML-4 ML-5 ML-6 Sorex cinereus 4 1 1 1 Sorex fumeus 2 1 Sorex palustris 1 Blarina brevicauda 1 Tamiasciurus hudsonicus 1 Peromyscus maniculatus 3 1 2 Microtus pe,nnsylvanicus 3 2 2 1 Clethrionomys gapperi 14 4 4 Napaeozapus insignis 8 Zapus hudsonius 2 6 8 6 3 Mustela erminea 1 - Total Trap Nights 392 328 240 267 270 262 Trap Success 8.9 % 1.2 % 3.8 % 4.5 % 3.7 % 5.0 % Number of Species 8 1 2 4 4 7 Simpson's Index D* 0.24 1.0 0.56 0.48 0.42 0.18 1-D** 0.76 0.0 0.44 0.52 0.58 0.82 • Shannon-Wiener H'*** 2.44 0.0 0.92 1.42 1.57 2.57 * = Probability of picking two organisms that are the same species. **=Probability of picking two organisms that are different species. *** =Average degree of uncertainty in predicting the species of an individual chosen at random .

• 8 TABLE 4.- Overview of small mammals collected in boulder fields and clear cut sites. Taxon BF-1 BF-2 BF-3 CC-1 CC-2 CC-3 Sorex cinereus 1 Sorexfumeus 1 • Blarina brevicauda 1 Tamias striatus 1 Peromyscus maniculatus 15 7 1 Peromyscus leucopus 1 Microtus pennsylvanicus 5 Microtus chrotorrhinus 2 Clethrionomys gapperi 6 16 1 1 Napaeozapus insignis 18 13 1 Zapus hudsonius 1 Mustela erminea 1 Mustela frenata 1 1 Total Trap Nights 90 190 355 70 '90 90 Trap Success 6.7% 21.6% 11.3 % 2.9 % 4.4 % 1.1 % Number of Species 2 5 7 2 4 1 Simpson's Index D* 0.72 0.35 0.30 0.50 0.27 1.0 1-D** 0.28 0.65 0.70 0.50 0.75 0.0 Shannon-Wiener H'*** 0.66 1.72 1.87 1.0 2.0 0.0 *=Probability of picking two organisms that are the same species . **=Probability of picking two organisms that are different species. ***=Average degree of uncertainty in predicting the species of an • individual chosen at random .

• 9 shrub swamp produced similar results with moderate trap success and species diversity (Table 3). Among the wetlands, the highest species diversity was observed in the beaver meadow with a total of seven species captured (Table 3) and signs of moles observed. A total of 11 species were captured from wetland • sites (Table 4) and the water shrew was only taken from a wetland site. The attempts to obtain additional captures of shrews by the placement of snap traps along swift-flowing streams and the establishment of pit traps with drift fences resulted in the capture of a single masked shrew and nine rodents. A total of 910 trap nights with pit traps produced one masked shrew from a dwarf shrub bog and three red-back voles from a black spruce/dwarf shrub bog. Snap traps set along two swift-flowing streams (Paul Stream and the Logger Branch of the Nulhegan) at five different sites for a total of 450 trap nights produced two red­ backed voles and four woodland jumping mice. Traps were set in four boulder fields for a total of 695 trap nights and produced a total of 8 8 captures of 11 different species including the masked shrew, s:r;noke shrew, short-tailed shrew, deer mouse, white-footed mouse, meadow vole, yellow-nosed vole, red-backed vole, woodland jumping mouse, long-tailed weasel and short-tailed weasel. The trap success within boulder fields ranged from 0.0 % to 21.6 % with an average 12.7%. One of the boulder fields was quite open with secondary growth ofraspberries, small saplings and grass from which a masked shrew and meadow voles were captured. The other three boulder fields were more mesic with more than 70 % canopy cover. Although one of these sites (which was very steep near Dennis Pond) produced no specimens after 60 trap nights the other two produced both high trap success and high species diversity (Table 4). Two specimens of the yellow-nosed or rock vole (Microtus • chrotorrhinus) were taken from one of the boulder fields (Table 4) but no specimens of the long-tailed shrew (porex dispar) were obtained. The yellow­ nosed vole was only taken from talus slope habitat. Three clear-cuts were trapped for a total of 250 trap nights (Table 4). The mean trap success of2.8 % was low but one site near a small patch of mixed hardwoods had a yield of 4.4 %. A total of six species were captured from clear­ cuts including the short-tailed shrew, chipmunk, deer mouse, red-backed vole, woodland jumping mouse and meadow jumping mouse.

Small Mammal Distributions and Abundance

Moles .. Only one of the two taxa of moles known from Vermont was captured during this survey. The presence of moles was observed by the presence of surface ridges along and across roads and trails in only a few areas of the Conte Refuge, primarily along 4 Mile Road between its juncture with Lewis Pond Road and the parking area to Mollie Beattie Bog and along Lewis Pond Road where it crosses over the Upper Yellow Branch of the Nulhegan River. One hairy-tailed mole (Parascalops breweri) was collected with a harpoon trap along 4 Mile Road approximately 1 mile west of its intersection with Lewis Pond Road. Although no • surface ridges or other evidence of moles was observed in the West Mountain WMA, a specimen was found by Brett Engstrom and Marc Lapin along South America Pond Road on the west side of Paul Stream above Ferdinand Bog.

Shrews. Shrews made up only 6.5 % of the captures, however, four of the six species of shrews known from Vermont were taken during this survey. Three species of the genus Sorex were captured during this survey (Table 1). The water shrew (S. pa/ustris) was taken along a tannic pool in a beaver meadow. Although this water shrew was the characteristic size and color of this taxon, the fringes of hair on the hind feet were not conspicuous and it had a very strong musky odor. In addition, this water shrew was heavily parasitized as a flea, a tick, and a number of liver flukes were collected from this animal. The masked shrew (S. cinereus) accounted for the greatest number of shr.ews captured and was taken in a diversity of habitats including montane yellow birch-red spruce forest, lowland spruce-fir forest, black spruce swamp, beaver meadow, and alluvial shrub swamp (Tables 2 and 3). Four specimens of the smoke shrew (S. fumeus) were taken from a black spruce swamp, a beaver meadow, and among the mossy covered boulders of a mixed forest (Tables 3 and 4). Only four specimens of the short-tailed shrew (Blarina brevicauda) were taken during this survey. This shrew was captured in a diversity of habitats including a montane yellow birch-red spruce forest, a dwarf shrub bog and in a clear cut (Tables 2-4).

Bats. Two species of bats were captured during this survey (Table 1). The northern long-eared bat (Myotis septentrionalis) was the most common species captured representing 71 % of the total captures whereas the little brown bat (M. /ucifugus) represented 29 % of the captures. The northern long-eared bat was taken from four different sites and the little brown bat from only two sites (Appendix II). Considerable variation in which species were captured was observed between different nights at a single location. Along an overgrown road within the forest on West Mountain one little brown bat and seven northern long­ eared bats were taken on 10 August. On 12 August eight little brown bats and two northern long-eared bats were taken from the same site. The acoustical survey showed a high level of bat activity at a number of sites where netting captured no specimens. These sites include Lewis Pond, Dennis Pond, and the Lower Black Branch of the Nulhegan. No additional species of bats could be identified by the acoustical survey.

Rodents. Ten species of rodents were captured during this survey (Table 1) and two additional species, the woodchuck (Marmota monax) and the beaver (Castor canadensis) were observed. Beaver are common and frequently observed along water courses and in many lowland sites. Although three woodchucks were . observed dead on route 105 during the summer, within the area being surveyed ground hogs were observed at only a single location near Paul Stream about 2 mi. SW route 102. No gray squirrels (Sciurus carilonensis) were observed or captured. Three species of sciurids were captured (Table 1), however, both red

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squirrels (Tamiasciurus hudsonicus) and chipmunks (Tamias striatus) were observed and heard much more frequently than they were captured. Chipmunks were observed or captured at most.upland sites including northern hardwood, red spruce-northern hardwood, and montane yellow birch-red spruce forests. Red • squirrels were observed, heard or captured from most upland sites and from a black spruce swamp (Tables 2 and 3). A single specimen of the northern flying squirrel (Glaucomys sabrinus) was taken in mid October from a mixed forest in the West Mountain WMA (Table 2). The most frequently captured small rodent was the red-backed vole ( Clethrionomys gapperi), representing 31 % of the total captures. This small mouse was taken from a variety of upland and lowland sites (Tables 2-4) where spruce or fir was present. Two other species of voles were captured during this survey but each was restricted to rather specific habitats. The yellow-nosed or rock vole (Microtus chrotorrhinus) was taken from mossy covered rocks at a single site within West Mountain WMA (Table 5). Although the meadow vole (Microtus pennsylvanicus) was taken from five different sites (Tables 3 and 4), these can all be characterized as open with growth oflow vegetation such as grass, sedges, or herbaceous annuals. The woodland jumping mouse (Napaeozapus insignis) was the second most abundant species captured, representing 20 % of the total captures. This ta:xon was generally restricted to upland sites (Table 2) and was more abundant than the red-backed vole at some sites. The woodland jumping mouse was also captured in one of the black spruce swamps surveyed (Table 3), in a boulder field within a mixed forest and in a clear-cut (Table 4). The meadow jumping mouse (Zapus hudsonius) was not found in any upland sites (Table 2) but was found • common in most lowland sites (Table 3) and was taken in one clear-cut (Table 4). Meadow jumping mice were not nearly as abundant as woodland jumping mice accounting for only 8.4 % of the total captures. Although both of these genera of jumping mice hibernate, woodland jumping mice were captured on 14 October. Jumping mice have been found active as late as early November, although most enter hibernation by 20 October (Whitaker, 1972; Whitaker and Wrigley, 1972). Mice of the genus Peromyscus were the third most abundant species captured, representing 14 % of the captures. Although these mice were generally much less abundant than expected, they were very abundant at a few sites (Table 4). Nearly all specimens captured were identified as deer mice (P. maniculatus) by analysis of salivary amylase samples. Several specimens had rather heavy parasite loads as was evident from the large number of eggs in their livers. Deer mice were taken from a variety of upland and lowland sites (Table 2 -4). A single specimen of a white-footed mouse (P. leucopus) was captured from the boulder field near Pauls Stream (Table 4). This is the same site where Peromyscus were especially abundant.

Small Carnivores. Both species of small mustelids (Mustela erminea and Mustela frenata) that are sometimes captured in Sherman live traps were taken during this survey. All five of the specimens collected were taken during the late summer and

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the fall (12 August to 15 October). Two specimens of the long-tailed weasel (Mustela frenata) were trapped in the West Mountain WMA, a female was collected along an old stream bed with large boulders off of Paul Stream about 2 miles west of route 102 and a male was collected among the large moss covered • boulders off of Notch Pond Road about 1 mile south ofroute 105. Three specimens of the short-tailed weasel or ermine (Mustela erminea) were captured from a variety of sites including an upland mixed forest (Table 2), a black spruce swamp (Table 3) and a boulder field within a mixed forest (Table 4). A female short-tailed weasel was captured in Big Swamp (Conte Refuge) whereas two males were captured in the West Mountain WMA. · Several observations were made of moderate and larger carnivores during the summer. A striped skunk (Mephitis mephitis) was observed one evening along Tin Shack road while returning for bat netting. Coyotes ( Canis latrans) were observed along several roads in the Conte Refuge and the West Mountain WMA and scat was commonly found along most roads early in the summer. Red foxes (Vulpes vulpes) were observed along Route 105~ along Stone Dam Road and along South American Pond Road. Scat of black bear (Ursus americanus)was commonly observed in the summer and bears were observed along Notch Pond and Lewis Pond Roads. Snowshoe hares ~epus americanus) were commonly seen along the roads in the early morning and the late evenings in both the Conte Refuge and the West Mountain WMA. Moose (Alces alces) were commonly seen along the roads in the Conte Refuge and routinely seen at the alluvial dwarf shrub swamp sampled. White-tailed deer (Odocoileus virginianus) were observed on three occasions in • the Conte Refuge and the West Mountain WMA. Discussion

This study documented the occurrence of 19 species of small mammals within the newly established Nulhegan Basin Division of the Silvio 0. Conte National Fish and Wildlife Refuge (NFWR) and the West Mountain WMA. Seventeen of these species were previously documented to occur within Essex County but this represents the first documentation of both the little brown bat (Myotis lucifugus) and the northern long-eared bat (M. septentrionalis) in this county.

Small Mammal Distributions and Abundance

Moles. Moles are aifficult to trap and often are rather rare and local in their distribution (Jackson, 1915). The two specimens of hairy-tailed moles (Parascalops breweri) obtained in this survey represent the first specimens from Lewis and Ferdinand Townships and the second record for Essex: County. Although the sandy soils characteristic of many of the areas surveyed seem suitable for hairy-tailed moles, surface ridges were only observed in a relatively small area between the Northern and Yellow Branches of the Nulhegan River

• 13 within the Conte Refuge. The surface ridges formed by hairy-tailed moles are neither pronounced nor easy to detect except in hard packed soil with little or no surface cover (Eadie, 1939). The collection of a specimen from an area where surface ridges were not observed, demonstrates that it is probable that the hairy­ • tailed mole is much more widely distributed within both the Conte Refuge and the West MoU11tain WMA.

Shrews. Although the diversity of shrews captured was high with a capture of five of the six possible species, the overall numbers were lower than expected with shrews making up only 6.5 % of the captures in live and snap traps. The use of pit traps with drift fences failed to increase substantially either the yield or the diversity of shrews, producing only a single shrew in 910 trap nights. The activity of small shrews seems to be greatly influenced by rainfall as 5 of the 20 captures occurred on a single night (July 29) with exceptionally heavy rain. The water shrew (Sorex palustris) was obtained from a single location, a beaver meadow/sedge swamp. Although this is not an unusual site in which to find a water shrew (Hamilton and Whitaker, 1979), it is typically found along the edges of swift-flowing streams (Conway, 1952). Specific trapping for water shrews along swift-flowing streams, however, failed to yield additional specimens. The relatively poor trap success for water shrews compared with·other sympatric species of shrews has been taken as evidence that water shrews maintain lower population densities (Kirkland and Schmidt, 1982; Nagorsen and Peterson, 1981). The masked shrew (S. cinereus) was the most commonly captured shrew accounting for 55 % of the shrews captured. The masked shrew is the most widely • distributed shrew in North America and has been reported from most habitats other than dry fields or woods (Godin, 1977). However, a recent study by Brannon (2000) which characterized the niche relationships of the masked shrew and the smoky shrew (S. fumeus) in the southern Appalachian mountains found that the smoky shrew had a greater niche breadth than the masked shrew. Brannon (2000) found that a combination of litter moisture, leaf-litter depth, slightly decomposed fallen logs and invertebrate size were the best predictors of masked shrew abundance. In this survey, the masked shrew was taken from five different natural communities (Tables 2 and 3) and from a boulder field in a disturbed habitat along a power line. This shrew was noticeably absence from both the northern hardwood and the red spruce-northern hardwood forest sampled. Smoky shrews (S. fumeus) accounted for 20 % of the shrews captured and were taken from relatively few natural communities (Tables 2-4). Although Brannon (2000) found that smoky shrews have a greater niche breadth than masked shrews in the southern Appalachian mountains, masked shrews were found in more community types than the smoky shrew in thi~ survey (Tables 2-4). Diminutive species such as the masked shrew feed on tiny surface-active and litter-active invertebrates (Whitaker and French, 1984; Ryan, 1986; Kirkland, 1991). Although there is some dietary overlap between smoky shrews and masked shrews (Hamilton, 1930; Whitaker and French, 1984), the smoky shrew

• 14 consumes considerably larger prey, such as earthworms, centipedes and salamanders than does the masked shrew (Hamilton, 1930; Whitaker and Cudmore, 1987). Brannon (2000) suggested that the breadth of prey items consumed by the smoky shrew contributed to its niche breadth and found that a • combination oflitter moisture, presence of extremely decomposed, soft moist logs that were partially buried by soil and forest litter, and number of invertebrates were the best predictor of smoky shrew abundance. Additional work is needed in the northern Appalachian mountains to understand the niche breadth and requirements of shrews of the genus Sorex. Godin (1977) suggested that the short-tailed shrew (Blarina brevicauda) is one of the most abundant small mammals in New England and occurs in nearly all habitats. Although the short-tailed shrew was found in a wide variety of habitats including forests, bogs, and clear-cuts (Tables 2-4), it was not commonly taken during this survey. This is in stark contrast with other areas of the state. A small mammals survey conducted during the summer of 2000 in Addison County (Decher and Kilpatrick, 2001) found this taxon to be the most common shrew. In Ess~x County, the short-tailed shrew represented 1.3 % of the total specimens captured whereas this species represented 17 % of the captures in the Addison County survey. Short-tailed shrew populations have been low in other areas of Essex County since 1997 (unpublished data) and thus the low numbers observed in a wide diversity of habitats may represent a population crash. Winter mortality of up to 90 % of populations of short-tailed shrews has been documented (Barbehenn, 1958; Gottschang, 1965; Jackson, 1961). Population densities are known to vary from year to year (Jackson, 1961; Platt, 1968) and populations of short-tailed shrews occasionally crash, requiring several years to recover (Ozoga and Verme, 1968). Further monitoring of this species should be conducted to verify that the current low population numbers is only a temporal fluctuation.

Bats. Capturing bats in flight with mist nets continues to be one of the most difficult and time-consuming aspects of biological surveys. Mist nets are often biased in their sampling because bats vary in habitat preference, height of flight, and flight speeds. A total of34 bats were captured in approximately 150 net hours, however, no bats were captured at six of the ten areas where nets were set (Appendix Il). Capture of the northern long-eared bat (Myotis septentrionalis) and the little brown bat (M lucifugus) represent the first documentation of either of these species from Essex County. In this survey, northern long-eared bats accounted for 71 % of the captures and little brown bats 29 %. This is consistent with most summer surveys of woodland bats in the northeast where the two most commonly captured species are the little brown bat and the northern long-eared bat (Krusic, 1995; Toth, 1999; Reynolds, 2000a; 2000b ). As an example, a total of 40 bats including little brown bats, northern long-eared bats and big brown bats (Eptesicus fuscus) were captured in 2s·net nights from five sites in the Green Mountain National Forest surveyed in July 2000 (Reynolds, 2000a) . Northern long-eared bats represented

• 15 48 % and little brown bats 40 % of the captures (Reynolds, 2000a). In the Green Mountain National Forest survey, most bats were captured along trails (53 %) whereas only 3 % were captured along streams .. In our survey, most captures also occurred along trails or narrow over grown logging roads (56 %), however, 41 % • of the captures were made along small streams and the remaining 6% along a road.

Rodents. Rodents represent about 80 % of the specimens captured and include nine species (Table 1). Although three species of sciurid rodents were captured during this survey, the survey techniques were not designed to determine the abundance of these species. The eastern chipmunk (Tamias striatus) was trapped or observed at represe:gtative of most upland communities (Table 2) and on the edge of one clear-cut (Table 4). Although primarily associated with deciduous wooded areas (Snyder, 1982), the chipmunk was taken or observed in two red spruce-northern hardwood forests. · Density ofchipmunk:s is reported to be temporally and geographically variable, ranging from 0.3 to 37.6 per ha (Yerger, 1953). In this survey, chipmunks were observed to be very common at two northern hardwood sites in the Conte Refuge. The presence of the red squirrel (Tamiasciurus hudsonicus) was determined by trapping , observing or hearing these squirrels as well as by observation of their middens. The presence of the red squirrel was observed in nearly all upland sites (Table 2) and one wetland site (Table 3). Layne (1954) found that red squirrels occurred in seven distinct habitat types in New York including mixed hardwoods with scattered conifers. Cool, moist conditions in boreal coniferous forest not only provide abundant food ( seeds and fungi) but also may aid in the preservation of middens and promote fungal growth (Rusch and Reeder, 1978). Densities have been reported at 0.3 to 2.0 squirrels per ha, however, significant declines in densities has been reported following selective and clear cutting (Steele, 1998). Although red squirrels appeared to be abundant at most mesic sites with some coniferous forest, their abundance can not be estimated. A single specimen of the northern flying squirrel (Glaucomys sabrinus) was trapped in a northern hardwood-red spruce forest in mid October. During the fall and winter flying squirrels are frequently captured in trap set on the ground (Connor, 1960, 1966) and considerable time is spent foraging on the ground. In New York, optimal habitat for northern flying squirrels contained beech, sugar maple, red maple, red oak, various birches and some hemlock and white pine, however, they were also taken from pure beech-:-maple forest, and stands of red spruce, balsam fir and hemlock (Connor, 1960). Population densities are reported to very from 1 squirrel per 3 ha to 10 per ha in favorable habitats (Wells-Gosling and Heaney, 1984). With the capture of only a single specimen, little can be said about either abundance or habitat requirements. · Although three species of voles were captured during this survey, their abundance range from common in the case of the red-backed vole (Clethrionomys gapperi) to rare in the case of the yellow-nosed vole (Microtus chrotorrhinus) .

• 16 ,.

Red-backed voles were the most abundant small mammal, based on numbers captured, and were taken in nearly all community types sampled (Tables 2-4) . They were not taken from either dwarf shrub bogs or alluvial shrub bogs and appear to be uncommon in northern hardwood forests. With the exception of the • lowland spruce-fir forest site, they were taken in most sites in which spruce was present. This vole is generally found in mesic habitats in coniferous, deciduous, and mixed forests with abundant litter of stumps, rotting logs, and exposed roots (Merritt, 1981). Although Kirkland (1977a) and Martell and Radvanyi (1977) have reported increased abundance of red-backed voles in clear cuts, Lovejoy (1975) found that minimal changes occurred in numbers of voles after logging of moist sites in a northern hardwood forest in New Hampshire. Although red­ backed voles were trapped in clear-cuts (Table 4), they were much less abundant in t~s habitat than at other sites. In Vermont, populations densities have been reported between O and 37 per ha (Miller and Getz, 1972, 1977). The yellow-nosed or rock vole (Microtus chrotorrhinus) was obtained fro:rii one of four talus slopes sampled in this survey (Table 4). This vole is usually associated with rocks, boulders and talus (Wyman, 1923; Osgood, 1938b; Martin, 1971), however, Kirkland (1977b) found the rock vole inhabiting clear-cuts in both northern coniferous and mixed deciduous forest. This vole was not trapped in any of the clear-cuts surveyed j.n this study (Table 4). Other components of the habitat associated with the rock vole are either the presence of surface or subsurface streams (Wyman, 1923; Martin, 1971; Kirkland and Knipe, 1979) and an abundance of mosses and forbs (Kirkland and Jannett, 1982). A high percentage of specimens of this vole that have been captured were obtained from subsurface sites (Timm et al., 1977; Kirkland and Knipe, 1979) indicating that • rock voles spend a considerable amount of time in subterranean activities. This behavior coupled with its habitat requirements may explain why the rock vole is underrepresented in collections compared to other sympatric species of voles. Fewer than 35 specimens of the rock vole have been collected from Vermont. The two specimens collected in this survey are from Brunswick Township and represent the third locatio:n from which this vole has been taken in Essex County. Previously this vole has been reported from Brighton (Osgood, 1938b) and Ferdinand (Chipman, 1994) Townships. Habitat selection by the meadow vole (Microtus pennsylvanicus) is thought to be influenced byrelative ground cover of grasses and herbs (Getz, 1970), soil moisture (Wrigley, 1974) and surface temperature and humidity (Getz, 1971). Although it is most commonly found in grasslands, preferring moist areas, it has also been found in woodlands (Reich, 1981). In this survey, the meadow vole was not a common species captured, occurring at 26 % of the sites (Tables 3 and 4) surveyed and repres_enting only 4.2 % of the specimens trapped. It was taken from dwarf shrub bogs, an alluvial shrub bog, a beaver meadow and among the boulders of a disturbed habitat along a power line but was only abundant at the latter site. The dispersal tendency of M pennsylvanicus is well documented and I ani sure more substantial populations of meadow voles occur in the Nulhegan Basin Division of the Silvio 0. Conte NFWR and the West Mountain WMA.

• 17 -l

In New England, the woodland deer mouse (Peromyscus maniculatus gracilis) and the white-footed mouse (P. leucopus noveboracensis) are forest­ dwelling species that may be found at the same site (Farren and Capen, 1985). These two species are not easily differentiated (Choate, 1973; Rich et al., 1996) which has lead to considerable confusion of the habitat requirements of these two • species in the northeast. All specimens captured in this survey were tentatively identified as deer mice (P. maniculatus) based on pelage and tail morphology (Choate, 1973). Examination of salivary amylase genetic markers (Aquadro and Patton, 1980; Kilpatrick et al., 1994) verified that most specimens were deer mice, however, a single white-footed mice was identified among the mice taken in a boulder field along Paul Stream in the West Mountain WMA.. Although the deer mouse contributed to 13.3 % of the total captures of small mammals and was taken from 52.6 % of the sites surveyed, it was not as abundant as expecte.d. In a small mammal survey in Addison County during the summer of 2000, miqe of the genus Peromyscus represented 47.9 % of the captures (Decher and Kilpatrick, 2001). Although deer mice were relatively abundant at a number of sites (Tables 2-4), they were only taken in number at two boulder sites (Table 4) and a red spruce-northern hardwood site with considerable deadfall (Table 2). This would suggest that the local abundance of deer mice may relate to the amount cifcover, but other sites that seem to have sufficient cover of down logs and stumps produced few deer mice. Many of the deer mice seem to have heavy parasite loads that was manifested by the presence of yellow egg masses in their livers_ Declines in abundance of deer mice in northeastern North America have been reported by Herman and Scott (1984) who could not rule out disease as the cause. Populations of deer mice are frequently infected with the • nematode Capillaria hepatica (Freeman and Wright, 1960; Solomon and Handley, 1971; Herman, 1981; Meagher,.1999). This direct life cycle nematode remains in the liver until death of the host and is thus transmitted through cannibalism, predation, necrophagy or natural death and decomposition (Freeman and Wright, 1960; Layne, 1968). Scavenging ground beetles and shrews may play an important role in the transmission of this nematode (Herman, 1981). Although infected deer mice show reduced liver protein synthesis, hepatomegaly, and splenomegaly (Meagher, 1998), little is known about the effects of this parasite on the dynamics of host populations. Godin (1977) reported that deer mice prefer habitats associated with coniferous forest consisting mainly of red, white, and black spruce and balsam fir, or northern hardwood forest comprised of mainly beech, sugar maple and yellow birch. Several authors (Klein, 1960; Farren and Capen, 1985) have concluded that deer mice prefer habitats with larger diameter trees and with less ground cover than habitats preferred by white-footed mice. Other workers (Kirkland, 1977a;. Martell and Radvanyi, 1977) have found that deer mice were abundant in clear­ cuts. In this survey, deer mice were taken from most wooded sites (Tables 2-4) but from only one of the three clear-cuts sampled. Deer mice were noticeably absent from the three bogs sampled (Table 3) .

• 18 Jumping mice were also abundant based on the numbers captured with woodland jumping mice (Napaeozapus insignis) representing 20.4 %, and meadow jumping mouse (Zapus hudsonius) accounting for 8.4 % of the total captures. These two jumping mice are fairly well segregated ecologically in • different stages in the successional sequence of plant communities, with Zapus primarily in meadows and Napaeozapus primarily in woods (Whitaker and Wrigley, ·1972). fu the intermediate stages, as in shrubs or forest-edge situations, the two species are commonly found together. The woodland jumping mouse inhabits cool, moist environments within the forest or forest-edge. Brower and Cade (1966) found no restriction to particular kinds of woods inhabited and Whitaker (1963) found that, in New York, swamps, mesic woods and wet woods were most often inhabited. fu this survey, the woodland jumping mouse was collected in all forested sites sampled with the exception of the lowland spruce-fir forest (Tables 2-4). The meadow jumping mouse lives in a variety of habitats (Quimby, 1951; Getz, 1961; Whitaker, 1963) including meadows, but is often more abundant in thick vegetation along ponds, streams and marshes. fu this survey, Zapus was taken from nearly all wetland sites sampled with the exception of the dense black spruce of Western Bog (Table 3). Zapus was also collected along the damp trail passing through a clear-cut (Table 4). The meadow jumping mouse was absent from nearly all forested sites (Tables 2 and 3) with exception of the black spruce swamp sampled.

Small Carnivores. Both species of small carnivores that were expected to be sampled by the survey techniques employed were captured. Other than to indicate • that both the long-tailed weasel (Mustela frenata) and the short-tailed weasel (Mustela erminea) are present, the number of specimens captured is too small to make any comments about habitat requirements or abundance.

Of the 19 species of small mammals captured during this survey (Table 1), 14 species were documented to occur in the Conte Refuge and 17 were documented from the West Mountain WMA. Three species including the water shrew (Sorex palustris), the yellow-nosed vole (Microtus chrotorrhinus) and the northern flying squirrel ( Glaucomys sabrinus) were each taken from a single location. The water shrew was taken.from the Conte Refuge whereas the yellow­ nosed vole and the northern flying squirrel were taken from West Mountain WMA. Although the first two of these are rare, additional sampling will document that each of these species is more widely distributed within this region. The northern flying squirrel is not rare but is only captured on the ground during the fall and winter. Two other species, the short-tailed shrew (Blarina brevicauda) and the long-tailed weasel (Mustelafrenata) were taken only in the West Mountain WMA. Whereas the absence of the long-tailed weasel from the Conte Refuge is the result oflimited fall sampling, the absence of the short-tailed shrew reflects a low population density and very localized distribution of this

• 19 .; ,.

shrew in the Northeast Kingdom. The causes of this localized distribution is unknown at this time.

Missing Species

Three additional species ofinsectivora, the star-nosed mole, the pygmy shrew, and the long-tailed shrew are expected to occur within the Nulhegan Basin Division of the Silvio 0. Conte NFWR and the West Mountain WMA. The star­ nosed mole ( Condylura cristata) was not found in either the Conte Refuge or the West Mountain WMA, however, it was observed for the first time in Essex County this summer from along Webster Brook in Morgan Township (C. A. Woods pers. comm.). The star-nosed mole prefers damp situations ranging from slow-flowing streams and swamps to damp meadows and woods (Hamilton and Whitaker, 1979). Although these moles are sometimes captured above ground with Sherman live traps or snap traps ( especially during spring flooding), they are more frequently captured in the water in muskrat sets or in minnow traps (Hamilton, 1931 ). It is highly probable that the star-nosed mole occurs in both the Conte Refuge and the West Mountain WMA as suitable habitat is abundant. Two shrews were tentatively identified as pygmy shrews (S. hoyi), but ~xamination of the cleaned skulls verified that both were masked shrew (S. cinereus). No specimens of pygmy shrews were collected during this survey. This is the smallest mammal in New England weighing between 2.2 to 3.6 g (Godin, 1977). This species is difficult to capture by conventional methods, however, pit traps have been effective elsewhere (Prince, 1941). Additional survey work with pit traps in areas such as Big Swamp and the Upper Yell ow • Branch Wetland Complex which have been shown to have high shrew diversity, or other areas identified with high shrew diversity, should be conducted to document the occurrence of the pygmy shrew. The long-tailed shrew (Sorex dispar) is found in relatively few habitat types, primarily among rocks, and especially along talus slopes (Kirkland, 1981; French and Crowell, 1985). The long-tailed shrew is also found in artificial talus created by road building (Conaway and Pfitzer, 1952) and mining operations (Kirkland, 1981), adjacent to cool mountain streams (Kirkland, 1981; French and Crowell, 1985) and in clear-cuts (Kirkland et al., 1976; Kirkland, 1977a). Several workers (Kirkland, 1977a, 1981; French and Crowell, 1985) have noted the similarity in habitat requirements of the long-tailed shrew and the yellow-nosed· vole (Microtus chrotorrhinus). Although yellow-nosed voles were found within the West Mountain WMA and thus suitable habitat for the long-tailed shrew was found~ no specimens of this shrew were captured. Although four talus slopes or boulder fields (Table 4) were trapped during this survey, there are a number of additional talus slopes that should be surveyed for the presence of both long-tailed shrews and yellow-nosed voles within the Nulhegan Basin Division of the Silvio 0. Conte NFWR and the West Mountain WMA. Other summer surveys of bats in Vermont woodlands have captured big brown bats (Eptesicus fuscus) and red bats (Lasiurus borealis) in small numbers

• 20 (Toth, 1999; Reynolds, 2000a). It is probable that additional bat netting will produce captures of these two species in low numbers. Other bats, such a.s hoary bats (;lasiurus cinereus) and silver-haired bats (Lasionycteris noctivagans), are larger and have a relatively high aspect ratio that restricts their flight to open • areas. These open areas can not easily be sampled by mist netting. Netting over open water has, to date, failed to produce any specimens of these larger bats. The habitat types sampled during this survey of bats were consistent with habitat used by the Indiana bat (M sodalis) and the small-footed bat (M leibii) in other parts of their ranges (Hitchcock, 1955; Kurta et al., 1993; Callahan et al., 1997). In addition, a specimen of small-footed bat has been collected from Island Pond (Godin, 1977) and a documented but not verified specimen of the Indiana bat was reported from northern New Hampshire in 1993 (Krusic, 1995). This survey was not adequate in scope to support the conclusion that M sodalis and M leibii do not use the Nulhegan Basin Division of the Silvio 0. Conte NFWR or the West Mountain WMA during the summer active season. It is also possible that the southern flying squirrel (Glaucomys volans) occurs in the Conte Refuge and the West Mountain WMA. Although this species has not been document~d from the Northeast Kingdom, it has been taken as far north as Fairfax in the Green Mountains and Coos County in New Hampshire (Godin, 1977). In addition, the northern flying squirrel (G. sabrinus) is only know from two specimens from the Northeast Kingdom. Two voles, that are difficult to trap, the pine vole (Microtus pinetorum) and the bog lemming (Synaptomys cooperi) were not obtained in this survey. Certainly the bog lemming and very possibly the pine vole occur within the Conte Refuge and the West Mountain WMA. The distribution of the bog lemming in • Vermont is know from a total of 49 specimens captured over the past 90 years (Kirk, 1916; Osgood, 1938a; Godin, 1977; Chipman, 1994; Kilpatrick, unpublished data; C. A. Woods, pers. com.). This rare lemming, however, is known from five different localities from Essex County including Ferdinand Township near the West Mountain WMA. The southern bog lemming is reported to occur in a variety of habitats other than bogs (Linzey, 1983) where they live in a complex series of burrows usually either in sphagnum mbogs or the deep leaf mold of forested areas. Attempts to locate their subterranean burrows were not . successful in ether bogs or forested areas. The distribution of the pine vole in Vermont is based on fewer than 40 specimens (Kirk, 1916; Osgood, 1938a; Miller, 1964; Godin, 1977) but it is known from northeastern Vermont including Essex County. This fossorially adapted vole is known to occur in a variety of habitats (Hamilton, 1938; Benton, 1955; Miller and Getz, 1969). In the northeast, pine voles are usually found in p.eciduous forest (Hamilton, 1938) however, soil type maybe a more important !uniting factor (lv[iller, 1964). This vole often occurs sympatriclywith the hairy­ tailed mole thus the hairy-tailed mole is a good indicator species for pine voles. The presence of hairy-tailed moles in the area surveyed indicates that pine vole habitat is present. Attempts to capture pine voles from subterranean runways used

• 21 by hairy-tailed moles was unsuccessful, in part due to a difficulty in finding runways or surface ridges.

Small Mammal Diversity

Trap and netting success and thus estimates of species diversity are affected by a number of factors including but not restricted to population densities (time of year), species behavior, weather conditions such as wind and rain, moon phase and placement of traps or nets. Even though most sites were sampled for three nights, the diversity estimates still are based on samples taken at one point in time. Sampling of these same habitats at different times will result in different estimates of species diversity. A total of nine species were captured from upland sites with three species representing more than 90 % of the total captures. Red-backed voles ( Clethrionomys gapperi) represented 53%, woodland jumping mice (Napaeozapus insignis)22 % and deer mice (Peromyscus maniculatus) 14 % of the to_tal captures. Six species including the masked shrew, the short-tailed shrew, the eastern chipmunk, the red squirrel, the northern flying squirrel and the short­ tailed weasel were captured only once or twice in upland sites (Table 2). Although the low number of captures of chipmunks, squirrels and weasels is in part the results of these animals not being the primary target of the trapping protocol, the low number of shrews captured in these upland sites is indicative of low population densities in these upland communities. The only shrews captured in these upland sites were two species with broad habitat requirements (Blarina brevicauda and Sorex cinereus). The seven upland sites samples had a mean species richness of3.4 and a mean species eveness (Shannon-Wiener) of 1.16. Of the two northern hardwood sites surveyed one was well below the mean and the other was slightly above these mean values. The northern hardwood with the lowest species diversity estimates (Table 2) was a site that was sampled early in June. Thus, the estimates from the second hardwood forest site with a species richness of 4 and a Shannon­ Wiener index of 1.46 is probably more indicative of this natural community. Average species richness and species eveness was found in two red spruce­ northern hardwood forest sampled, whereas a third site (U-7) sampled in mid October had higher estimates (Table 2). The montane yellow birch-red spruce forest had the highest estimates of species richness and eveness of the upland sites sampled. A total of eleven species were captured from wetland sites but the four most common species represented account for only 76 % of the total captures. Meadow jumping mice (Zapus hudsonius) represented 30 %, red-backed voles (Clethrionomys gapperi) 27%, meadow voles (Microtus pennsylvanicus) 9.5 % and woodland jumping mice (Napaeozapus insignis) 9.5 % of the total captures. The six wetland sites sampled had a higher mean species richness (4.3) and mean species eveness (Shannon-Wiener index of 1.49) than the upland sites. This additional species richness and eveness was contributed in part by shrews. Among

22 the wetlands, four species of shrews were captured and shrews represented 13 .3 % of captures, whereas among the upland sites only two species of shrews were captured and shrews contributed to only 4.7 % of the captures. The lowest species diversity and species richness among the wetland sites was found in a black • spruce/dwarf shrub bog where only red-backed voles were captured (Table 3). Moderate species diversity and eveness was found in other dwarf shrub bogs and an alluvial swamp (Table 3). High indices of species diversity were found in both a black spruce swamp and a beaver/sedge meadow (Table 3) .. This high species diversity at both of these latter sites result from the capture of both forest and open habitat species and richer shrew faunas. · A total often species were captured from boulder fields with three species representing about 87 % of the total captures. Woodland jumping mice (Napaeozapus insignis) represented 35 %, red-backed voles (Clethrionomys gapperi) 26%, and deer mice (Peromyscus maniculatus) 26 % of the total captures. Boulder fields had a mean species richness (3.5) and species eveness (Shannon-Wiener of 1.06) rather sinular to upland sites. The four boulder fields (talus slopes) sampled, however, were very heterogeneous in both canopy cover and slope. The site with no canopy cover (BF-1) had below average species diversity (Table 4). The other three sites were within enclosed canopies. The site with the greatest slope produced no specimens and thus has no estimated species diversity (data not shown). The remaining two talus slopes within forest (BF-2 and BF-3) had the highest population densities observed and higher than average indices of species diversity (Table 4). Not only is this microhabitat very important for some species such as the yellow-nosed vole and the long-tailed shrew, but it also provides abundant cover and hence supports higher densities of several common species of small mammals including deer mice, red-backed voles, and woodland jumping mice. This microhabitat would also appear to be important for small carnivores, no doubt responding to the high population density of small mammals, as 60 % of the captures of weasels came from this habitat (Table 4). Clear-cuts had the lowest mean estimates of species richness (2.3) and eveness (1.0) of any of the habitats sampled. Although 6 species were captured from clear-cuts, the trap success and hence population density was relatively low (Table 4). The site (CC-2) with the highest indices of species diversity among the three clear-cut sites was near a small patch of hardwood forest, and some of the species collected in this clear-cut, such as the eastern chipmunk and the woodland jumping mouse, were clearly forest species. Only red-backed voles were collected from more than one clear-cut site. Clear-cuts have been found to be important habitat for some relatively rare species in West Virginia, including the long-tailed shrew (Sorex dispar) and the yellow-nosed vole (Microtus chrotorrhinus) (Kirkland et al., 1976; Kirkland, 1977a). In addition, an increased abundance of red-backed voles (Clethrionomys gapperi) has been observed in clear cuts (Kirkland, 1977a; Martell and Radvanyi, 1977). Clear-cuts in the Conte Refuge do not appear to provide habitat for rare species such as the yellow-nosed vole or long-tailed shrew as neither of these species were captured in our survey of this habitat (Table 4). Red-backed voles

• 23 were much more abundant in other habitats (Tables 2 and 3) than in these clear­ cut (Table 4). Thus, we have no evidence that clear-cuts are important habitats for small mammals in the Northeast Kingdom.

Conservation and Management Concerns for Small Mammals

Most of the small mammals captured in this survey are relatively comm!Jn in Vermont. Onl three relatively rare species were captured including the water shrew, ygmy shrew and the yellow-nosed vole. The habitats associated with (1,01 rJh these rarer species in this survey include beaver/sedge meadows and talus slopes ~Q\ {\e,CJ)1-, 1r in northei:11 hardw.ood-~ed s~ruce forests. . . . . '(° f ,-i & -So High species diversity was observed at a number of sites mcludmg Big t" l C:i Sr ('>(c, v y Swamp (a black spruce swamp), the Upper Yell ow Branch open wetland complex 0 iJ ( a beaver/sedge meadow), a talus slope within a red spruce-northern hardwood forest and a montane yellow birch-red spruce forest. High population densities were observed at some of the same sites with higher indices of diversity, as well as two red spruce-northern hardwood forests and the boulder field of an old stream channel in a red spruce-northern hardwood forest. Sites identified as being inhabited by rare species and communities with high species diversity and/or population densities should be considered as important habitats for small mammal populations. Habitats that were not examined in this survey, such as small intermittent stream channels, have been shown to support high species diversity of small mammals in Vermont (Decher and Kilpatrick, 2001). Finally, little knowledge is available regarding the roost sites of the bats that uses this area during the summer. Future work should include some radiotelemetry to locate the roost sites used by northern long-eared bats and little brown bats.

Recommendations for Future Work

Provide adequate time and funds for future survey work. Although we began our field work in late May, the funding was not received until mid August. Much of the equipment purchased to conduct this survey work was only available for the last few trips of the field season. In addition, funds were eliminated for items such as Anabat II detectors and GPS units when it was learned that such items of equipment could by borrowed. Both the limitation on time period for which some of this· equipment was available and unexpected problems encountered in piecing together functional units from components from different donors and a 6 week delay to replace a lost cable did not allow the best use of the · Anabat II detectors for acoustical surveys of bats. Addition survey work for bats should be high priority. This should include both additional mist netting and acoustical monitoring. We gained a lot of knowledge about where and how to set mist nets to capture bats in northern forests during this survey. Many of our earliest mist net sets produce no captures

24 (Appendix II). At least two Anabat II detectors should be available for future work. One should be used as a remote monitoring station to detect bat activity at a number of additional sites and those sites that show high activity patterns should be sampled with mist nets on subsequent nights. The other unit should be used • either as an additional remote monitoring station or in conjunction with bat netting. Although acoustical monitoring does not measure species abundance or other population biology characteristics, it does provide a lot of information about habitat use with a relatively small amount of effort and cost. A complete analysis of the acoustical monitoring data should be conducted to determine the presence of non-captured species, however, species of the genus Myotis can not be consistently differentiated by acoustical signature. Thus, additional mist netting is need to determine if the state-threatened small-footed bat (M leibii) and.I or the federally-endangered Indiana bat (M sodalis) are present on the Nulhegan Basin Division of the Silvio 0. Conte NFWR or the West Mountain WMA. Additional mist netting should be conducted at a variety of sites including the open water of ponds and larger rivers, trails and small logging roads, streams, and other areas with high bat activity detected by acoustical monitoring, to obtain additional information on species diversity and abundance of bats. As mentioned above, tagging captured bats with radiotransmitters, would allow the identification of summer roost sites. This would provide information on the habitat needs of both common bat species and perhaps some of the rarer bat species . Although 19 sites representing 10 different communities were surveyed for small mammals during the summer and fall of 2000, many areas remain • unsampled. Future work should allow some of the sites that were sampled in 2000 to be resampled at adifferent time of the year, such as sampling the hardwood forest in the fall rather than in June, and thus provide better estimates of the species richness 0fthese communities. The major emphasis, however, should be on sampling new sites. Additional representatives of communities with either high or low indices of species diversity, such as black spruce swamps and lowland spruce-fir forest, should be sampled to obtain a broader view of the small mammal diversities of these communities. A major focus, however, should be to seek out and survey the microhabitats associated with the species that were either found in low numbers or not found in this first survey. This would include additional sampling of talus slopes, small inteffilittent stream channels, swamp edges, and beaver meadows. · Surprisingly, pit traps with drift fences did not prove to be very successful in this survey (1 capture in 910 pit trap nights). This was in part the results of setting pit traps in areas that had demonstrated low trap success with non-pit traps, in order to obtain additional species from these sites. However, in a survey in Addison County (Decher and Kilpatrick, 2001) during the summer of 2000, 84 pit trap nights resulted in a 15.5% trap success and captured 5 species of small mammals. Larger buckets (195 x 160 mm) were used in the Addison County work, whereas smaller buckets (110 x 140 mm) were used in this survey. It is • difficult to know how much of the very low trap success of pit traps in this survey 25 was the result of the smaller sized buckets as compared to the selection of sites with low mammal densities. However, additional pit trapping should be conducted using larger buckets in habitats with both low and high population densities of small mammals. • Nineteen species of small mammals (Table 1) were document in the Nulhegan Basin Division of the Silvio 0. Conte National Fish and Wildlife Refuge (NFWR) and the West Mountain Wildlife Management Area during the first summer survey. The cumulative number of species captured (Fig. 1) has not yet reached a plateau phase, suggesting that additional sampling will produce additional species. In particular, I would expect to find the star-nosed mole (Condylura cristata), the pygmy shrew (Sorex hoyi), the long-tailed shrew (.Sorex dispar), the big brown bat (Eptesicus fuscus), the red bat (Lasiurus borealis)~ the southern bog lemming (.Synaptomys cooperi) and the pine vole (Microtus pinetorum). It is also possible with additional sampling, that the presence of some of the following mammals will be documents: the Indiana bat (Myotis sodalis), the small-footed bat (M leibii), the eastern pipistrel (Pipistrellus subjlavus), the silver-haired bat ~asionycteris noctivagans), the hoary bat ~asiurus cinereus), and the southern flying squirrel (Glaucomys volans).

Acknowledgments

Funding for this small mammal survey was provided through a contract from the Nature Conservancy through its Vermont Field Office. We would like to thank Edward Toth of the Green Mountain National Forest and Susi vonOettingen • of the U.S. Fish and Wildlife service for the loan of Anabat II detectors. Two graduate students, Ryan A. Norris and Mike Malia, did the bulk of the field work under my supervision. I thank Ryan and Mike for their hard work and dedication to collecting the data for this survey. I also thank Dr. Charles A. Woods for the loan of a GPS unit and providing us a place to store equipment during this project.

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• 33 A endix I. Location of sites tra ed in small mammal surve . Communi ·GPS Dates Surve ed Northern hardwoods N44 52.124 W71 45.315 June 8-10 Northern hardwoods N44 38.424 W71 40.064 June 25-27 Red spruce-northern N44 47.418 W71 44.972 July 9-11 hardwoods UM-4 Red spruce-northern N44 40.282 W71 39.973 June 14-16 hardwoods UM-5 Lowland spruce-fir N44 47.122 W71 43.331 June 11-13 UM-6 Montane yellow birch- N44 42.826 W71 42.261 .July 30-August 1 red spruce U-7 Northern hardwoods- N44 44.669 W71 41.043 October 15 red spruce LM-1 Black spruce swamp N44 51.081 W71 44.923 July 27-30; October 1 LM-2 Black spruce-dwarf N44 49.995 W7146.868 July 6-8; October shrub bog 13 Dwarf shrub bog N44 41.316 W71 43.478 August9-11 Dwarf shrub bog N44 43.476 W7139.249 June 28-30 Alluvial shrub swamp N44 45.758 W7142.795 August2-4 Beaver meadow/sedge N44 49.528 W71 54.530 July 16-18 meadow/marsh F-1 Open boulder field N44 44.682 W71 41.910 July 18 BF-2 Wooded boulder field N44 41.117 W7138.187 August 12-14 with little slope BF-3 Wooded boulder field N44 44.990 W71 42.371 August 16; with large moss covered August24; boulders and ledges September 30 BF-4 Steep boulder field near Dennis Pond CC-1 Clear-cut N44 47.138 W71 43.827 June 10-13 CC-2 Clear-cut near island N44 47.441 W71 44.472 July20 patch of hardwoods CC-3 Clear-cut N44 47.585 W71 44.104 Au ust 15

• 34 A endix II. Location and results of bat surve Site GPS Net Dates Nights M. septentrionalis lucifu us ULB N44 52.125 W71 45.248 4 8/2-8/3 0 4 LLB N44 51.482 W7144.568 2 8/2-8/3 0 2 UBB N44 48.224 W71 42.481 1 7/31 0 0 LBB N44 46.768 W71 40.692 3 8/1 0 0 Lewis Pd N44 52.865 W71 46.695 1 7/30 0 0 UYB N44 49.528 W71 45.530 1 7/27 0 0 LYB N44 48.716 W7144.388 6 8/13-8/14 1 8 Goodwin N44 40.827 W7140.792 4 6/13-6/14 0 0 WestMtn N44 43.431 W71 42.900 6 8/10.;8/12 9 10 Dennis Pd N44 43.512 W71 39.250 2 7/18 ,0 0

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