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Bog Turtle Habitat on the Lake Ontario Coastal Plain of New York State
Peter A. Rosenbaum and Andrew P. Nelson

Northeastern Naturalist, Volume 17, Issue 3 (2010): 415–436

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2010 NORTHEASTERN NATURALIST 17(3):415–436 Bog Turtle Habitat on the Lake Ontario Coastal Plain of New York State Peter A. Rosenbaum1,* and Andrew P. Nelson2 Abstract - We develop a concept of suitable habitat for Glyptemys (Clemmys) muhlenbergii (Bog Turtle) on the Lake Ontario Coastal Plain of central and western New York State and compare it to habitat for this species in other parts of its distribution. At the outset of our studies in 1987, only a single Bog Turtle population was known to survive on the Lake Ontario Plain. We located and visited nine of the 10 historically recorded Bog Turtle sites in northern, central and western New York and completed a survey of 84 selected wetlands in Oswego County, at the eastern end of the Lake Plain. In the course of the study, we confirmed the presence of the Bog Turtle at a site where it was last documented in 1916 and discovered populations at three previously undocumented sites, one of which is now the northernmost recorded site for the species. Bog Turtle habitat on the Lake Ontario Coastal Plain is open-canopy, sedge-dominated medium to rich fens contained within larger wetlands that often include an open pond and extensive Acer rubrum (Red Maple) or Red Maple-Larix laricina (Tamarack) swamp. We discuss events that may have led to the current distribution of Bog Turtle populations in central and northern New York and in western Pennsylvania and implications of this distribution pattern for the conservation of the species. Introduction Glyptemys (Clemmys) muhlenbergii Schoepff (Bog Turtle) is a New York State (NYS)-listed endangered species, a CITES I species, and a federally threatened species under the US Endangered Species Act. The species includes two major groups of populations, a “southern population”, extending from southwestern Virginia south to northern Georgia, and a “northern population”, extending from New York (NY) and western Massachusetts south to northern Maryland, northern Delaware, and southern New Jersey (Fig. 1; USFWS 2001). Bog Turtle sites in western and central NYS, along with a historic site in Warren County, NY, and historic sites in Crawford and Mercer counties, PA, comprise the Prairie Peninsula/Lake Plain Recovery Unit (PP/LPRU; Fig. 2), the northernmost of five recovery units established in the Bog Turtle Recovery Plan and located within the northern population of the species (USFWS 2001). As a group, Bog Turtle sites in this recovery unit are disjunct from the main part of the northern population. The US Fish and Wildlife Service (USFWS) has determined that “no fewer than 10 viable 1Department of Biological Sciences, SUNY Oswego, Oswego, NY 13126. 2Rice Creek Field Station, SUNY Oswego, Oswego, NY 13126. *Corresponding author - 416 Northeastern Naturalist Vol. 17, No. 3 Bog Turtle populations and sufficient habitat to ensure sustainability” must be protected in the PP/LPRU in order to achieve recovery (USFWS 2001). This paper contains the first published description of Bog Turtle habitat in the PP/LPRU. Habitat described for other parts of the Bog Turtle’s distribution include a wide variety of predominantly herbaceous wetland community types (Table 1). Physical features of the vegetation deemed significant by other authors include: 1. an open canopy (Arndt 1977, Buhlmann et al. 1997, Herman and Tryon 1997, Kiviat 1978, Somers 2000, USFWS 2001, Zappalorti 1997), 2. dominance of graminoid species (sedges, grasses, and rushes) (Bury 1979, Chase et al. 1989, Kiviat 1993, Somers 2000, Taylor et al. 1984), 3. presence of hummocks or tussock-forming sedges (Collins 1990, Kiviat 1993, Taylor et al. 1984, Zappalorti 1997), and 4. adjacent wooded wetlands (Taylor et al. 1984, USFWS 2001). Prior to the beginning of our studies in 1987, historic reports of Bog Turtle occurrence within the area now designated as the PP/LPRU included four sites in western Pennsylvania, eight sites in west-central New York, and one site in Figure 1. Distribution of the Bog Turtle, indicating locations of the northern and southern populations. (Modified from USFWS 2001). 2010 P.A. Rosenbaum and A.P. Nelson 417 the Lake George basin in Warren County, NY (Table 2) (Bury 1979, Collins 1990, USFWS 2001). The most recent documentation of a new site involved the capture of a single individual at an Oswego County site in 1985. In the early 1990s, a specimen, collected in 1931 from a previously unrecorded site in Monroe County, NY, was found in the collections of the NYS Museum, bringing the total of historically known sites in the NYS portion of the recovery unit to 10. Records for six of these sites are limited to one or two individual turtles or to empty shells or bones. A site with multiple records in Tompkins County is known to have been destroyed (Collins 1990). At the beginning of our studies, only one site (Seneca #1; Table 2) within the PP/LPRU was known to support an active population of the Bog Turtle. We have monitored the population at this site since 1987, conducting demographic, genetic (Rosenbaum et al. 2007), radio-telemetry, and trapping studies. In 1994, a floristic inventory was initiated in conjunction with our ongoing turtle studies. In 1997, we began a survey of historic sites and other wetlands in Cayuga, Genesee, Monroe, Onondaga, Oswego, Seneca, and Wayne counties. Figure 2. Counties in northern New York and western Pennsylvania containing sites where the Bog Turtle has been recorded. Because the Bog Turtle is threatened by illegal collection for the pet trade, the exact locations of extant and historic sites are not included. Extant means that the county is currently known to harbor the Bog Turtle; historic means there is a reliable historic record of the Bog Turtle from wetlands that still exist; extirpated means there is a reliable historic record of the Bog Turtle from a wetland that no longer exists. 418 Northeastern Naturalist Vol. 17, No. 3 Our goals were to 1) determine whether suitable Bog Turtle habitat remains at historic sites within the area and, if so, whether the continued existence of the Bog Turtle could be established, 2) discover previously undocumented populations of the Bog Turtle, 3) describe the characteristic features of Bog Turtle habitat on the Lake Ontario Coastal Plain (LOCP) portion of the PP/LPRU, 4) compare Bog Turtle habitat on the LOCP with that found in other portions of the range of the species, and 5) explore implications of this information for conservation of the species in the PP/LPRU. Field-site Description Sites selected for study included those from which the Bog Turtle had previously been reported (Genesee #1, Onondaga #1, Monroe #1, Oswego #1, Seneca #1, Tompkins #1, Warren #1, Wayne #1, Wayne #2, and Wayne #3 in Table 2), sites designated as “bogs” in an earlier survey of wetlands in Oswego County (Jones et al. 1983a, b), and other sites containing open peatland communities, most commonly inland poor fens, medium fens, or rich graminoid fens, wetland communities distinguished chiefly on the basis of pH and floristic composition (Edinger et al. 2002). Most of these sites are on the LOCP of NYS, ranging south to Seneca County and extending west to Genesee County and east to Onondaga and Oswego counties. Table 1. Ecological communities associated with the Bog Turtle in various parts of its distribution. Community Source Woodland streams Arndt 1977 Ponds Arndt 1977, Bury 1979 Swamps Bury 1979, Herman and George 1986 Tamarack, Black Spruce swamps Ernst et al. 1994, Herman and George 1986, Netting 1927 Fens Herman and Tryon 1997; Kiviat 1978, 1993; USFWS 2001 Bogs Arndt 1977, Bury 1979, Collins 1990, Ernst et al. 1994, Herman and George 1986, Herman and Tryon 1997 Sphagnum marsh Fisher 1887 Meadow bogs (Bog meadows) Collins 1990, Herman and Tryon 1997 Marshes Arndt 1977, Zappalorti et al. 1993 Marshy meadows Ernst et al. 1994, Herman and George 1986 Sedge meadows USFWS 2001 Wet meadows Arndt 1977, Breisch 1988, Buhlmann et al. 1997, Collins 1990, Herman and George 1986, Kiviat 1978, Taylor et al. 1984, Zappalorti et al. 1993 Pastures Carter et al. 1999; Chase et al. 1989; Herman and George 1986; Herman and Tryon 1997; Morrow et al. 2001a, b; Somers 2000 Abandoned fields Arndt 1977, Chase et al. 1989 2010 P.A. Rosenbaum and A.P. Nelson 419 Table 2. Size of wetlands, ponds (when present), and open fens at active and historic Bog Turtle sites in the New York State portion of the PP/LPRU. “Population status” as in Figure 2. Area values cited to 0.01 ha are quoted from GIS shape files (NYS DEC 1999). Values cited to nearest ha are estimated from aerial photographs using ArcGIS® v. 9. Bog Turtle reports Population Known/reported County, site # Original Most recent Surveyed status Wetland (ha) Pond (ha) Open Fen (ha) specimens Latitude (°N) Elev. (m) Seneca #1 1906 Present x Extant 20 8 6 36 42.9572 142 Wayne #1 1916 Present x Extant 72 Drained 8 2 43.1858 124 Oswego #2 1994 Present x Extant 132.88 less than 1 28 7 43.5811 74 Oswego #3 1995 Present x Extant 115.48 46 10 84 43.3733 105 Oswego #4 2000 Present x Extant 162.96 32 6 21 43.4400 194 Warren #1 1883 — Historic 5131 — 63 8 43.4410 100 Genesee #1 1898 1970s x Historic 932 — 6 18 43.0978 174 Monroe #1 1931 — x Historic 2060 — None left 1 43.0128 189 Onondaga #1 1950 — x Historic 2080 — None left 1 43.1442 117 Wayne #2 1960 — x Historic 30 — None left 1 43.0556 125 Wayne #3 1960s 1983 x Historic 236 Seasonal 14 3 43.1528 127 Oswego #1 1985 — x Historic 29.84 34 8 1 43.3689 173 Tompkins #1 1877 1940 x Extirpated ------ Site destroyed ------ 8 42.4147 400 420 Northeastern Naturalist Vol. 17, No. 3 Methods Site visits We surveyed nine of the 10 Bog Turtle sites within the NYS portion of the PP/LPRU previously documented as historic or extant (Table 2), 84 of the 88 sites listed as “bogs” in the wetlands inventory of Oswego County (Jones et al. 1983a, b), and 12 other sites in Cayuga, Oswego, Lewis, Ontario, and Wayne counties. The plant communities encountered were noted, and a list of plant species was developed for each community. Plant nomenclature used here conforms to the New York Flora Atlas (Weldy and Werier 2008), with synonyms given where necessary for understanding of the Bog Turtle literature. Notes were made on the nature of the substrate, water sources and drainage, and the pH of the water in various parts of the wetland. This procedure approximates a Phase 1 Bog Turtle Habitat Survey as described in the Bog Turtle Recovery Plan (USFWS 2001). We also visited six Bog Turtle sites outside the PP/LPRU in Dutchess and Columbia counties in southeastern NYS. This gave us experience to supplement our literature-based understanding of habitats in other Bog Turtle recovery units. Visual search and trapping Sites selected for further study were revisited and subjected to careful visual search of microhabitats favored by the Bog Turtle and probing of likely subsurface retreats. These searches approximate Phase 2 Habitat Surveys as detailed in the Bog Turtle Recovery Plan (USFWS 2001). Trapping surveys were initiated during May and June at 13 sites where either turtles had previously been recorded, live turtles were discovered in the course of this study, or repeated visits reinforced the impression of suitable habitat. Box traps were constructed of 2.54-cm square plastic coated wire mesh with funnel-type entrances. Four trap designs were used: 1) small—10 x 10 x 40 cm with entrances at both ends flanked by 35 cm long adjustable wings, 2) medium—35 x 35 x 48.5 cm with an entrance at one end, 3) large—35 x 35 x 67.5 cm with an entrance at one end, and 4) large—35 x 35 x 65 cm with entrances at both ends. Traps were not baited. They were set to intercept animals moving along game trails, small drainage channels, and other apparent runways. Clumps of vegetation placed on top of the traps provided shade for trapped animals. Traps were checked every 24 hours, and animals were removed for data collection (e.g., body measurements and blood samples) and released. Occasionally, traps were moved from their original position as suggested by trapping success or as required by changes in water level. Trap arrays were based on size and structure of the habitat and utilized 24–165 assorted traps (mean = 66.2) for 19–126 days (mean = 34.8). Radio telemetry At all sites where the Bog Turtle was found, some animals were outfitted with transmitters and monitored to assess their seasonal activities and 2010 P.A. Rosenbaum and A.P. Nelson 421 habitat use. In this way, information was obtained on various stages of the Bog Turtle’s life cycle, such as nesting, feeding, basking, brumation, and aestivation. As Bog Turtles often share facilities such as hibernacula, following one individual by telemetry sometimes led to the discovery of additional individuals. When possible, males and females were monitored in order to compare possible ecological and behavioral differences between the sexes. Various models of transmitters and receivers were utilized during our studies (1992–2007). In all instances, transmitter packages (transmitters and glue) weighed less than ten percent of the turtle’s body weight. Larger turtles (greater than 80 g) were generally outfitted with larger transmitters and smaller units were utilized on smaller turtles (less than 80 g). Transmitters were glued on the posterior surface of the carapace with the antenna, if used, circumscribing the edge of the shell using techniques similar to those described by Eckler et al. (1990). Results Documentation of Bog Turtle sites At this writing, the Bog Turtle is known to exist at five sites in the NYS portion of the PP/LPRU (Table 2). The species has been known at Seneca #1 since 1906. Over the course of our surveys at this site (1987–2008), 36 Bog Turtles have been documented, 10 of which were head-started from native stock. Field (1987–1999), telemetry (1992–97, 1999), and trapping studies (1997) have been conducted at this site. A Bog Turtle shell was discovered at Oswego #3 in 1995, and live turtles were documented in 1997. Subsequent field surveys (1997–2009), trapping (1997, 2002, 2005), and telemetry (1997–99, 2002, 2005) studies have documented a vigorous population of at least 84 individuals. The New York State Department of Environmental Conservation (NYS DEC) considers this to be one of the best sites for the Bog Turtle in NYS (A. Breisch, NYS DEC, Albany, NY, pers. comm.). A new population of Bog Turtles was discovered at Oswego # 4 in 2000 as a result of trapping a wetland we identified as a potential Bog Turtle site on the basis of vegetation, floristics, and other habitat features. Field surveys (2000, 2001, 2002), trapping (2000, 2001), and telemetry (2000, 2001, 2002) studies have documented 21 specimens at this site. In 2004, trapping resulted in the capture of a single Bog Turtle at what we believe to be the historic site in Wayne County (Wayne #1) described by A.H. Wright (1918a, b; 1919). Wright provided no locality data for this “open moor” or “moor-like area” other than the name of a wetland in Wayne County, west of Oswego, at an elevation of 420 feet. In addition to trapping (2004) studies, we conducted field surveys (2004, 2005) and telemetry (2004, 2005) studies at this site. Field surveys (2005, 2006), radio telemetry (2005–2007) studies, and trapping (2006) studies have documented seven Bog Turtles at Oswego #2. 422 Northeastern Naturalist Vol. 17, No. 3 This is a very large site with extensive suitable habitat (i.e., 28 ha of open fen). At 43°35' N (UTM 18N 4826), it is the northernmost recorded site for the Bog Turtle. Several of the “historic” sites (Table 2) in the NY portion of the PP/LPRU have also been surveyed. At Oswego #1, from which a single individual was reported in 1985, repeated field surveys over numerous field seasons by us and others and trapping surveys by us (2000, 2004) have failed to uncover any Bog Turtles. We have designated this site as “historic” in Table 2. Unsuccessful trapping and field surveys were also conducted by us at Onondaga #1 (2006) and Wayne # 3 (1991). Both of these sites have also been surveyed by others. Likewise, Genesee #1 has been the subject of unproductive surveys and related studies by us and others. The only surviving historic site in the NYS portion of the PP/LPRU that has not been the focus of recent field surveys by us or others is Warren #1, which is a very large wetland (i.e., 68 hectares of open fen) that is outside of our study area (i.e., the LOCP of central and western NY) and is logistically difficult to access (A. Breisch, pers. comm.). Description of habitat Plant communities. Current and historic sites investigated in Genesee County, Monroe County, Seneca County, and central Wayne County are marl fens, rich graminoid fens, and rich sloping fens, while eastern Wayne County and Oswego County sites are medium fens as described by Edinger et al. (2002). These open-canopy communities are dominated by a variety of different sedges (Cyperaceae). Scattered hummocks of Sphagnum spp. (peat mosses) around the base of small trees and shrubs such as Acer rubrum (Red Maple), Larix laricina (Tamarack), and Toxicodendron vernix (Poison Sumac) provide nesting and basking sites as well as hibernacula. The open-canopy fens of active and historic sites in the PP/LPRU are adjacent to Red Maple or Red Maple-Tamarack swamps. Telemetry carried out on animals captured and released at Seneca and Oswego county sites reveals that some Bog Turtles in central NY spend up to three months of the active field season in wooded swamps adjacent to the sedge-dominated open fens. Disturbance and succession. The substrate at Bog Turtle sites in central and western NY is so soft and deep that mowing or grazing by Bos taurus L. (Domestic Cow) is not and probably has not been a factor in their formation and maintenance. There is little evidence that Odocoileus virginianus Zimmermann (Whitetail Deer) have had a significant effect on the vegetation beyond keeping narrow trails open across the fens. Castor canadensis Auhl (North American Beaver) may be present, but in all active and historic sites we have seen in this area, its activities result in changes in the water level in preexisting wetlands rather than in the establishment of ponds and subsequent beaver meadows in stream channels. Wildfires have not been a significant factor for most sites in this area in historic times, though there is anecdotal record of some burning at 2010 P.A. Rosenbaum and A.P. Nelson 423 Wayne #1. Intentional burning to favor fruits such as Vaccinium corymbosum (Highbush Blueberry) and Gaylussacia baccata (Wang.) Koch (Black Huckleberry) may have occurred prehistorically. There is a history of Sphagnum collection for florist packing material at Wayne #3 prior to 1914 and again for a period of several years starting in 1939 (Stauffer 1959). We cannot be sure if this activity affected portions of the wetland historically occupied by the Bog Turtle, but it is clear that successional change has taken place since moss collection ceased (Stauffer 1959). Peat mining was conducted at Oswego #3 from 1880 to 1902. Again, we do not know the extent of the disturbance at this site, but contemporary accounts describe the effect as dramatic (Rowlee 1897). A review of aerial photographs back to 1938 suggests that the overall outline of the open fen at this site has remained relatively unchanged for the last 75 years. Wetlands adjoining Wayne #1 and #3, Onondaga #1, and Oswego #3 have been ditched and drained for muckland agriculture. Additional drainage was installed at Wayne #1 to alleviate flooding of an upstream access road. Although this has drained a significant area of fen, the site is still easily recognizable when compared to aerial photographs from 1938. Until the early 1990s, a local community used Seneca #1 as a water supply, drawing as much as 250,000 gallons of water per day. The water level of the pond at Oswego #4 was raised and stabilized by a dam constructed in 1937 (M. Kautz, NYS DEC, Cortland, NY, pers. comm.). Size and distribution of sites. Current, recent, and historic sites in westcentral NY consist of large, open fens (6–28 ha) contained within much larger wetland complexes of over 20 ha, and in some cases, of over 100 or even 1000 ha (Table 2). These wetlands are relatively pristine, having been protected by their size from disturbance and development. They are quite isolated, being at or near the headwaters of separate drainage basins and in no case closer than 15 km to each other. As previously stated, although the initial capture of individual turtles has almost always taken place in the open fen, telemetry and mark and recapture studies at Seneca #1, and Oswego #2, #3, and #4, have demonstrated that these animals do enter, spend time in, and traverse the surrounding wooded swamps. Substrate and hydrology. Bog Turtle sites in central and western NY are all characterized by an accumulation of saturated organic soils ranging from only slightly decomposed fibric (peat) through partially decomposed hemic (mucky peat) to more highly decomposed sapric (muck) soils varying in depth and underlying materials. Technical soil types mapped for these sites include Adrian muck, Carlisle muck, Chippeny muck, Edwards muck, Palms muck, Rifle muck, Muck - deep, and Humaquepts and Fibrists (Higgins and Neeley 1978; Natural Resources Conservation Service 2006a, b, c; Rapparlie 1981). Cores revealed organic sediments to a depth of over 8 m at Oswego #3 and over 13 m at a historic site, Wayne #3 (Shipman 1970). Marl deposits are found at the Genesee and Seneca county sites. 424 Northeastern Naturalist Vol. 17, No. 3 At the Seneca County site, rivulets arise from springs at the edge of the wetland basin and flow through the fen to a centrally located pond, while in the Oswego County and eastern Wayne County sites, flooded game trails provide avenues by which turtles and other small animals make their way through the fens. Surface waters of small pools and game trails in the sedgedominated medium and rich fens occupied by the Bog Turtle on the LOCP have pH values ranging from 5.0 to 8.4, though at some sites the sedge fens may grade into Sphagnum-dominated areas of poor fen with lower pH (≤5). There is some outflow of water from the wetlands at all of the current and historic Bog Turtle sites on the LOCP. At most times of the year, the outflow appears to exceed the inflow from surface streams and runoff, suggesting that there is ground water discharge into these systems. Three of the five fens (i.e., Seneca #1, Oswego # 3, Oswego #4) currently known to be inhabited by the Bog Turtle occur at the margins of large ponds (Table 2). One of the other two sites, Wayne #1, was historically associated with a large pond that has now been drained. The remaining site, Oswego #2, has a small pond. Our measurements of the pH of surface waters of the existing ponds have always been higher than readings taken in the adjacent fens on the same day. These data suggest that input and circulation of calcium-enriched ground water may serve to maintain the relatively high pH of these wetlands. Water supplying these wetlands likely has percolated through glacially deposited calcareous sands or gravels or through calcareous bedrock. Floristics. Eighty-one vascular plant species are common to all of the active Bog Turtle sites in Oswego County (Table 3). The active Wayne County site has 54 of these species, and the Seneca County site has 49 (Table 3). Forty-two species are common to all Lake Plain active sites (Table 3). Species absent from the Seneca County site but found at all Oswego County sites as well as the Wayne County site are either characteristic of medium and poor fen communities and usually absent from richer fens, or they are wetland generalists widely distributed at other sites in the region. The predominant sedge of open fen areas at the Seneca County site as determined by visual estimate was Eleocharis rostellata (Torr.) Torr. (Beaked Spike-rush). In this part of NYS, E. rostellata is associated with marl fens and rich graminoid fens (Edinger et al. 2002). It is not recorded from any of the medium fens of Oswego County. Oswego #3 is dominated early in the growing season by a mixture of Carex exilis Dewey (Coast Sedge) and Trichophorum alpinum (L.) Pers. (= Scirpus hudsonianus) (Alpine Cottongrass), being replaced by mid-summer by Cladium mariscoides (Muhl.) Torr. (Twig Rush or Smooth Sawgrass) and Rhynchospora alba (L.) Vahl. (White Beakrush). At Oswego #4, the predominant sedge of the open fen is Carex lasiocarpa Ehrh. ssp. americana (Fern.) Hultén (American Woollyfruit Sedge), though Cladium mariscoides and R. alba are also present. Oswego #2 is a large and diverse site where Carex lasiocarpa predominates in some open areas and Cladium mariscoides or the combination of Carex exilis, T. alpinum, 2010 P.A. Rosenbaum and A.P. Nelson 425 Cladium mariscoides, and R. alba in others. Carex exilis, T. alpinum, Cladium mariscoides, and R. alba are also common in a wide variety of medium and poor fens in Oswego County and elsewhere on the LOCP. Toxicodendron vernix is present in all of the fens currently known to support the Bog Turtle on the LOCP (Table 3), and in all known historic sites in the NY portion of the PP/LPRU. This species also occurs in many wooded swamps in Oswego County. Vascular plant species that characterize active and historic Bog Turtle sites of the LOCP are listed in Table 4. While each site has some of these species, no site has all of them, and there seems to be no one plant species or set of species that suggests itself as a useful Bog Turtle habitat indicator throughout this region. Discussion Comparison of habitats Plant communities. Bog Turtle sites we observed in southeastern NY are rich, sloping fens and sedge meadows. Sites in other parts of the species distribution represent a variety of ecological communities (Table 1). On the LOCP, Bog Turtle populations occupy medium, rich, or marl fens, with one site—Seneca #1—containing a small area of rich, sloping fen. These open fens typically occur at the margins of ponds and are surrounded by extensive Red Maple-Tamarack or Red Maple swamps. Outside the PP/LPRU, tussocks formed by Carex stricta Lam. (Tussock Sedge or Upright Sedge) as well as Sphagnum hummocks growing up around the base of shrubs and small trees provide nesting, basking and hibernacula sites (Lee and Norden 1966). Tussock Sedge is not part of the communities where the Bog Turtle has been found within the PP/LPRU. Wet meadows, marshes, pastures, and swamps with many of the plants listed for Bog Turtle sites in other parts of the species’ distribution are abundant and common on the LOCP, but Bog Turtles have never been reported from such sites in this area. Sphagnum-dominated poor fens of a more acid nature are particularly abundant towards the eastern end of the LOCP, far outnumbering the relatively rare medium and rich fens of the sort that support the Bog Turtle; however, the Bog Turtle has never been reported from poor fens in this region. Disturbance and succession. The open habitats occupied by the Bog Turtle are generally thought to be successional, dependent for formation and maintenance on grazing by large herbivores such as Cattle or Whitetail Deer, mowing or other agricultural activities, fire, or Beaver (Breisch 1988, Buhlmann et al. 1997, Collins 1990, Herman and Tryon 1997, Kiviat 1978, Lee and Norden 1996, USFWS 2001). Kiviat (1978) developed a conceptual model to illustrate how Bog Turtle habitat may change in the context of various scenarios of succession and disturbance. The wetlands occupied by the Bog Turtle in central and western NY are the direct result of glacial phenomena. They have developed along the 426 Northeastern Naturalist Vol. 17, No. 3 Table 3. Plant species found in all active Bog Turtle sites in Oswego County, indicating those also present in the active Wayne and Seneca county sites. O = Oswego County, W = Wayne #1, and S = Seneca #1. Species O W S Acer rubrum L. (Red Maple) x x x Alnus incana (L.) Moench. (Speckled Alder) x x Amelanchier canadensis (L.) Medic. (Serviceberry) x x x Andromeda polifolia L. (Bog-rosemary) x x Arethusa bulbosa L. (Swamp Pink) x Aronia melanocarpa (Michx.) Ell. (Black Chokeberry) x x x Bidens connata Muhl. ex Willd (Beggar-ticks) x x Boehmeria cylindrica (L.) Sw. (Bog-hemp) x x x Calamagrostis canadensis (Michx.) Beauv. (Bluejoint) x x x Calla palustris L. (Wild Calla) x Calopogon tuberosus (L.) BSP. (Grass Pink) x x Caltha palustris L. (Marsh Marigold) x x x Carex atlantica Bailey (Eastern Sedge) x x Carex canescens L. (Silvery Sedge) x Carex lacustris Willd. (Hairy Sedge) x x x Carex lasiocarpa Ehrh. (Wooly-fruit Sedge) x x x Carex limosa L. (Mud Sedge) x Carex stipata Muhl. ex Willd. (Awl-fruited Sedge) x x x Chamaedaphne calyculata (L.) Moench. (Leatherleaf) x x Chelone glabra L. (White Turtlehead) x x x Cladium mariscoides (Muhl.) Torr. (Bog-rush) x x x Comarum palustre L. (Marsh Cinquefoil) x Cornus sericea L. (Red Osier Dogwood) x x x Decodon verticillatus (L.) Ell. (Water-willow) x x x Doellingeria umbellata (Miller) Nees (Tall Flat-topped Aster) x x x Drosera intermedia Hayne (Spatulate-leaved Sundew) x Drosera rotundifolia L. (Round-leaf Sundew) x x x Dulichium arundinaceum (L.) Britt. (Three-way Sedge) x Epilobium coloratum Biehl. (Purple-leaf Willow-herb) x x Epilobium palustre L. (Marsh Willow-herb) x Eriophorum tenellum Nutt. (Rough Cottongrass) x x Frangula alnus P. Mill. (Smooth Buckthorn) x x x Galium trifidum L. (Three Petal Bedstraw) x x Ilex verticillata (L.) A. Gray (Winterberry) x x x Impatiens capensis Meerb. (Spotted Jewelweed) x x x Iris versicolor L. (Northern Blue-flag) x x x Juncus canadensis Gray ex LaHarpe (Canada Rush) x Kalmia polifolia Wang. (Bog-laurel) x Larix laricina (DuRoi) Koch (Tamarack) x x x Lonicera oblongifolia (Goldie) Hook. (Swamp Fly Honeysuckle) x x x Ludwigia palustris (L.) Ell. (Water Purslane) x x Lycopus uniflorus Michx. (Water-horehound) x x Lysimachia terrestris (L.) BSP. (Swamp-candles) x Lysimachia thyrsiflora L. (Tufted Loosestrife) x x x Maianthemum canadense Desf. (Wild Lily-of-the-valley) x x x Menyanthes trifoliata L. (Bog Buck-bean) x Muhlenbergia glomerata (Willd.) Trin. (Spiked Muhly) x x x Nemopanthus mucronatus (L.) Loesener ex Koehne (Mountain Holly) x Onoclea sensibilis L. (Sensitive Fern) x x Osmunda cinnamomea L. (Cinnamon Fern) x x x 2010 P.A. Rosenbaum and A.P. Nelson 427 Table 3, continued. Species O W S Osmunda regalis L. (Royal Fern) x x x Peltandra virginica (L.) Schott ex Schott & Endl. (Arrow-arum) x x Persicaria arifolia (L.) Haraldson (Arrowleaf Tearthumb) x Pinus strobus L. (White Pine) x x Platanthera clavellata (Michx.) Luer (Green Woodland Orchid) x Pogonia ophioglossoides (L.) Juss. (Rose Pogonia) x x Pontederia cordata L. (Pickerel-weed) x Proserpinaca palustris L. (Mermaid-weed) x Rhamnus alnifolia L'Her. (Alder-leaf Buckthorn) x x x Rhynchospora alba (L.) Vahl (White Beakrush) x x x Rosa palustris Marsh. (Swamp Rose) x x x Rubus pubescens Raf. (Dwarf Raspberry) x x Salix discolor Muhl. (Pussy Willow) x x x Sarracenia purpurea L. (Pitcher-plant) x x x Scutellaria lateriflora L. (Mad-dog Skullcap) x x x Solanum dulcamara L. (Climbing Nightshade) x x Solidago uliginosa Nutt. (Northern Bog Goldenrod) x x x Symphyotrichum puniceum (L.) A. Löve & D. Löve (Purple-stem Aster) x x x Thalictrum pubescens Pursh (Tall Meadow-rue) x x x Thelypteris palustris Schott (Marsh Fern) x x x Toxicodendron vernix (L.) Kuntz (Poison Sumac) x x x Triadenum virginicum (L.) Raf. (Marsh St. John's-wort) x x x Trientalis borealis Raf. (Star-flower) x x Typha latifolia L. (Broad-leaf Cat-tail) x x x Utricularia gibba L. (Cone-spur Bladderwort) x Utricularia intermedia Hayne (Milfoil Bladderwort) x x x Vaccinium corymbosum L. (Highbush Blueberry) x x x Vaccinium macrocarpon Ait. (Large Cranberry) x x Vaccinium oxycoccos L. (Small Cranberry) x Viburnum nudum L. (Wild Raisin) x Viola pallens (Banks ex DC) Brainerd (Wild White Violet) x x x Totals 81 54 49 margins of ponds that were established in the lowlands between drumlins, in kettle holes, or behind lake shore barrier bars as the glacier receded and the level of lakes along the southern margin of the glacier fluctuated over the last 10,000+ years (Sutton et al. 1972). This model presents a picture of slow, long-term successional development of the wetlands as contrasted to the relatively short-term changes suggested by Kiviat (1978). Successional conversion of open fens to swamp forests certainly does occur in central and western NY and may have been of significance at Wayne #3, but it seems that some sites may persist in an open state at least for centuries. Our observations suggest that a more immediate threat exists in the form of invasion of open sites by Typha spp. (cat-tails), possibly coincident with cultural eutrophication, or exotics such as Phragmites australis (Cav.) Trin. ex Steud. ssp. australis (Common Reed), Frangula alnus (= Rhamnus frangula) (Smooth Buckthorn), or Lythrum salicaria L. (Purple Loosestrife). Size and distribution. Bog Turtle sites have been described as “small” (Morrow et al. 2001b, USFWS 2001). Sites studied in Delaware, Maryland, 428 Northeastern Naturalist Vol. 17, No. 3 and North Carolina cover no more than 2 ha (5 acres) (Arndt 1977, Lee and Norden 1996, Morrow et al. 2001a). There may be significant opportunities for exchange between small, semi-isolated populations that function as members of a metapopulation distributed along stream drainages (Buhlmann et al. 1997) or, at one site in the southeastern section of NYS, occupy suitable habitats within a large wetland complex that supports a mosaic of habitat types (A. Breisch, pers. comm.; P. Novak, NYS DEC, Schenectady, NY, pers. comm.). Sites we observed in southeastern NYS were small wetlands clearly bordering directly upon upland habitats. There is the possibility of interchange between local populations inhabiting the same watershed in that area. PP/LPRU sites are larger, with open fen habitat covering 6–28 ha or more and total wetland size varying from 20–2080 ha (Table 2). The sites are isolated in different drainages and separated by distances of 15 km or more, suggesting that there is currently no possibility of gene exchange between separate populations. Substrate and hydrology. The substrate at Bog Turtle sites has been described as soft mud (Carter et al. 1999, Somers 2000, Zappalorti 1997), deep mud (Arndt 1977), soft mud and rock (Chase et al. 1989), soft organic or loamy (Kiviat 1993), soft and highly organic (Ernst et al. 1994), or muck (Collins 1990, Zappalorti 1997). Although there is little information available concerning depth of the substrate, we have seen sites in southeastern Table 4. Plant species characteristic of active and historic Bog Turtle sites on the Lake Ontario Coastal Plain. Species Agalinis paupercula (A. Gray) Britt. var. borealis Pennell (False-foxglove) Carex flava L. (Yellow Sedge) Carex hystericina Muhl. ex Willd. (Porcupine Sedge) Carex lasiocarpa ssp. americana (Fern.) Hultén (American Woollyfruit Sedge) Carex leptalea Wahl. (Bristly-stalk Sedge) Cladium mariscoides (Twig Rush) Dasiphora (Potentilla) fruticosa (L.) Rydb (Shrubby Cinquefoil) Eleocharis rostellata (Torrey) Torrey (Beaked Spike-rush) Lobelia kalmii L. (Kalm's Lobelia) Lonicera oblongifolia (Swamp Fly-honeysuckle) Muhlenbergia glomerata (Spiked Muhly) Myrica gale L. (Sweet Gale) Myrica pensylvanica Loisel. ex Duhamel (Northern Bayberry) Parnassia glauca Raf. (Carolina Grass-of-Parnassus) Rhamnus alnifolia (Alderleaf Buckthorn) Solidago patula Muhl. ex Willd. (Roughleaf Goldenrod) Solidago uliginosa (Bog Goldenrod) Symphyotrichum boreale (Torr. & Gray) A.& D. Löve (Boreal Aster) Toxicodendron vernix (Poison Sumac) Utricularia intermedia (Flatleaf Bladderwort) 2010 P.A. Rosenbaum and A.P. Nelson 429 NYS where the soil was only a few inches deep over bedrock. At the PP/LPRU sites, the fen mat lies over a deep accumulation of peat or muck composed largely of the remains of the dominant sedges. Kiviat (1993) cites groundwater discharge as a significant characteristic of Bog Turtle habitat in southeastern NYS. Table 1 lists some community types from other areas that are most likely surface-water wetlands. At Seneca #1, groundwater enters the wetland from marginal springs. At other PP/LPRU sites, it appears that groundwater input supplements the input from tributary streams and surface runoff. Considerations of Bog Turtle habitat stress the importance of small, shallow, flowing streams or “rivulets” within the wetlands (Arndt 1977, Buhlmann et al. 1997, Bury 1979, Carter et al. 1999, Collins 1990, Ernst et al. 1994, Herman and George 1986, USFWS 2001, Zappalorti 1997). Such rivulets occur at the Seneca #1 site. However, in other PP/LPRU sites, the rivulets of flowing water characteristic of sites farther south are represented by flooded game trails traversing the sedge mat of the fen. There is little if any current in these channels, leading some Bog Turtle investigators who have seen them to refer to them as “pseudo-rivulets”. Floristics. Taylor et al. (1984) listed Carex aquatilis Wahl. (Water Sedge) as a dominant plant species of Bog Turtle habitat in Maryland. Lee and Norden (1996) cite Carex stricta as an important component of the vegetation of Bog Turtle sites in Maryland, and Zappalorti (1997) found C. stricta and Sphagnum spp. to be important in New Jersey, North Carolina, and eastern Pennsylvania. Arndt (1979) lists Leersia oryzoides (L.) Sw. (Rice Cutgrass) as a dominant species at some sites in Delaware. Kiviat (1993) emphasizes the importance of Dasiphora (Potentilla) fruticosa auct. non (L.) Rydb.(Shrubby Cinquefoil) and Parnassia glauca Raf. (Carolina Grassof- Parnassus) as indicator species for Bog Turtle sites in southeastern NYS. Sphagnum mosses are mentioned in the floristic lists of a number of references (Buhlmann et al. 1997, Carter et al. 1999, Ernst et al. 1994, Herman and George 1986, Taylor et al. 1984). Leersia oryziodes, although common in many wetland communities on the LOCP, is not present at Bog Turtle sites in NYS. Carex stricta occurs only as an occasional component of the vegetation peripheral to some of the open fen mats on PP/LPRU Bog Turtle sites. Carex aquatilis occurs in the vegetation fringing the pond at Seneca #1, but is absent from other sites. Dasiphora fruiticosa and P. glauca occur at Seneca #1 and other historical sites in the western portion of the LOCP, but do not occur at any Oswego County sites. Sixteen vascular plant species were found at all southeastern NY sites visited (Table 5). Fifteen of these also occur at the Seneca County site. Eight of the 16 are found at the active Wayne County site. Muhlenbergia glomerata (Spiked Muhly), Thelypteris palustris (Marsh Fern), and Typha latifolia (Broad-leaf Cat-tail), all components of a wide variety of other wetland communities, are present at all Bog Turtle sites surveyed. 430 Northeastern Naturalist Vol. 17, No. 3 Of the five active LOCP sites, Seneca #1 is most similar floristically to sites in southeastern NYS. Notable floristic differences exist between the flora of the active Oswego County sites and Seneca #1. Wayne #1, in the eastern part of Wayne County, supports a mix of some species found at Seneca #1 and some found at the Oswego County sites. Historic sites from central Wayne County west to Genesee County resemble Seneca #1 floristically. The active Oswego County sites (Oswego #2, #3, and #4), and the historic Oswego #1 site resemble each other floristically. Biogeography and conservation The extant and historic Bog Turtle sites of the PP/LPRU make up a set of highly localized disjunct populations along the northern boundary of the species distribution (USFWS 2001). Most of the LOCP populations, both current and historic, occupy sites in separate drainages leading to Lake Ontario. The historic sites in western Pennsylvania are in drainages that lead to the Ohio River and thence to the Mississippi. The Warren County site is in the Lake George drainage, which currently empties into Lake Champlain and ultimately into the St. Lawrence River. These sites are all at or relatively near the headwaters of their respective drainages. Lee and Herman (2000) suggest that the Bog Turtle was originally a Midwestern species that achieved its current distribution following eastward migration during the Pleistocene. They consider populations inhabiting the PP/LPRU to consist of three units: two disjunct, currently historic areas in extreme western Pennsylvania, the southern shore of Lake Ontario (also Table 5. Plant species found in all southeastern New York (SE NYS) sites visited, indicating those also present in the active Lake Ontario Coastal Plain sites. S#1 = Seneca #1, W#1 = Wayne #1, O#2 = Oswego #2, O#3 = Oswego #3, and O#4 = Oswego #4. SE Species NYS S#1 W#1 O#2 O#3 O#4 Carex flava (Yellow Sedge) x x x Carex hystericina (Porcupine Sedge) x x Carex stricta Lam. (Tussock-Sedge) x x x x Dasiphora fruticosa (Shrubby Cinquefoil) x x Eriophorum viridi-carinatum (Engelm.) Fern. (Cotton-Grass) x x x x Eupatorium perfoliatum L. (Boneset) x x x x Eutrochium maculatum (L.) E. Lamont (Spotted Joe-Pye-Weed) x x x x Juncus nodosus L. (Knot-Rush) x x Lythrum salicaria L. (Purple Loosestrife) x x Muhlenbergia glomerata (Spiked Muhly) x x x x x x Parnassia glauca (Grass-of-Parnassus) x x Scirpus atrovirens Willd. (Bulrush) x x Solidago patula (Spreading Goldenrod) x x x x Symplocarpus foetidus (L.) Salisb. ex Nutt. (Skunk-Cabbage) x x x Thelypteris palustris (Marsh Fern) x x x x x x Typha latifolia (Broad-Leaf Cat-Tail) x x x x x x Totals 16 15 8 4 9 4 2010 P.A. Rosenbaum and A.P. Nelson 431 referred to as west-central New York or the Finger Lakes area), and a historic population in the upper central Hudson River drainage in Warren County. They attribute this distribution to west-to-east dispersal utilizing drainage systems connected by lakes and wetlands formed along the melting ice fronts of the glaciers (i.e., “glacial outlets”) in the Laurentian Basin. Fossil evidence attests to the existence of Pleistocene refugia in Maryland and South Carolina (Bentley and Knight 1998, Holman 1977). A recent study has identified the remains of Bog Turtles from a site utilized by Native Americans approximately 3900 years ago in what is now Livingston County, NY (Madrigal 2008). All known historic and prehistoric sites within the currently recognized PP/LPRU region were covered by the Late or Classical Wisconsin glaciation, which reached its peak about 18,000–20,000 ybp (Fairchild 1909, Isachsen et al. 1991, Prest 1976, Sevon and Braun 1997). Smith (1957), in a consideration of post-glacial biogeography emphasizing amphibians, reptiles, and small mammals, considers the Bog Turtle to be an essentially northern species that was pushed south by the advancing glacier and subsequently reestablished populations in the northern glaciated region, leaving relict populations in the South. Genetic evidence shows relative uniformity among populations in the northern portion of the distribution of the Bog Turtle, which now extends south as far as Delaware and Maryland (Rosenbaum et al. 2007). The ubiquity of a single mitochondrial DNA haplotype (“A”) among populations of this region may be the result of a selective sweep that affected all populations of the northern region that survived the climatic extremes of the late Pleistocene and/or a significant bottleneck that occurred in conjunction with post-glacial colonization of the region from further south. Additional haplotypes (“B” and “C”), found in LOCP populations, are both derived from haplotype “A” (Rosenbaum et al. 2007). These findings suggest that the PP/LPRU populations are derived from and represent a northern expansion and diversification of the genetic strain present at the end of the Pleistocene in the southern portions of the currently recognized northern population of the species. Presuming this to be the case, the literature relating to postglacial events in what is now the PP/LPRU suggests that the Bog Turtle could have entered this area in ways consistent with the distribution of authenticated current, historic, and prehistoric sites by any of at least five different routes: 1) from the south via the Susquehanna drainage, 2) from the south via the Hudson drainage, 3) from the east and south via the Hudson/Mohawk drainage, 4) from the west via drainage channels along the ice front, and/or 5) from the south via the Hudson/ Champlain drainage and subsequent expansion to the west via drainage along the ice front (Fairchild 1909, 1925; Fullerton 1980; Prest 1976). Given the difficulties experienced in finding Bog Turtles at sites where they currently exist, it is highly probable that the species went unreported at sites that have been obliterated by drainage for agriculture and 432 Northeastern Naturalist Vol. 17, No. 3 development since European colonization of the area. The currently known and historically recorded populations on the LOCP, then, may be viewed as relicts spared during the course of local and regional climatic and topographic changes that occurred in post-glacial times, followed by the many and severe changes brought about by human activity in recent centuries. While it may be possible to characterize the biological and physical features of currently occupied and historically reported sites, we suggest that conclusions concerning the detailed biogeographic history of the Bog Turtle in this region will remain highly speculative. In some areas, the Bog Turtle exists in metapopulations, a distribution pattern which allows for occasional genetic exchange and establishment of new populations by means of dispersal along stream and river drainages (Buhlmann et al. 1997). In such instances, the protection of dispersal corridors that connect the units of the metapopulation is of primary conservation importance (Morrow et al. 2001a). However, the currently known populations on the LOCP occupy isolated wetlands (the closest separated by at least 15 km) mostly at or near the headwaters of their separate respective drainages. There is no possibility of genetic interchange between them at the present time nor is the establishment of new populations at new sites by natural means a likely possibility. The current distribution of extant and historically authenticated populations represents the relict survivors of post Pleistocene climatic and cultural events. Conservation of the Bog Turtle on the LOCP is very much a matter of protecting existing populations at their current sites unless the artificial establishment of new populations becomes feasible. Acknowledgments Financial support for field work on Bog Turtles has been provided by the NYS DEC, the USFWS, the NYS Biodiversity Research Institute, the Oswego County Environmental Management Council (OCEMC), and the Bergen Swamp Preservation Society. The OCEMC graciously provided access to the original field records of the Oswego County Wetlands Mapping Inventory project. The NYS DEC provided technical assistance in this study. Over the years, numerous students and colleagues have provided advice and assistance in various aspects of our work. We are especially grateful for the vigorous support of Alvin R. Breisch of the NYS DEC Endangered Species Unit and Robyn A. Niver of the USFWS. Erik Kiviat and three anonymous reviewers provided helpful suggestions on the manuscript. 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