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Reintroduction of the Eastern Woodrat (Neotoma floridana) in Southern Illinois
Aaron K. Poole, Brian A. Novosak, Aaron C. Gooley, David M. Ing, Robert D. Bluett, Timothy C. Carter, and George A. Feldhamer

Southeastern Naturalist, Volume 12, Issue 1 (2013): 1–10

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2013 SOUTHEASTERN NATURALIST 12(1):1–10 Reintroduction of the Eastern Woodrat (Neotoma floridana) in Southern Illinois Aaron K. Poole1,2, Brian A. Novosak1,3, Aaron C. Gooley1,4, David M. Ing1,5, Robert D. Bluett6, Timothy C. Carter7, and George A. Feldhamer1,* Abstract - Populations of Neotoma floridana (Eastern Woodrat) are decreasing in parts of their geographic range in the southeastern United States, and the species is stateendangered in Illinois. Once found throughout the Shawnee Hills region of southern Illinois, woodrats were restricted to four known populations in Jackson and Union counties by the late 1980s. We used reintroductions to establish viable populations of Eastern Woodrats at previously occupied sites in Illinois. From April 2003 through March 2009, we released 422 Eastern Woodrats live trapped in Arkansas and Missouri into 5 historically occupied sites in southeastern Illinois. Recapture rate 1 month after release was 12.5%. The continued presence of woodrats at release sites, reproduction, and wide dispersal beyond reintroduction sites all suggest preliminary success of the reintroduction of this r-selected species. Introduction Historically, Neotoma floridana Ord (Eastern Woodrat) was widespread in suitable habitats throughout southern Illinois, where it was considered common (Howell 1910), but is now one of the most endangered mammalian species in the state (Nÿboer et al. 2006). Nawrot and Klimstra (1976) found direct evidence of past habitation (e.g., skeletal remains, scat, stick nests) at 24 sites throughout Jackson, Union, Johnson, Pope, Gallatin, and Hardin counties. Additional sites seemed to offer suitable habitat, but no direct evidence of past habitation was observed, possibly because artifacts were destroyed by weathering and disturbance (Nawrot 1974). Four extant, remnant populations remain in Jackson and Union counties (Fig. 1). The largest, most secure, and best studied of these populations is at Pine Hills (Crim 1961, Ing 2008, Layne 1958, Monty 1997). Smaller populations remain at Fountain Bluff (Wagle and Feldhamer 1997), Horseshoe Bluff, and Little Grand Canyon (Monty et al. 1995, Wagle 1996). Although these populations are separated by only 2 to 14 km, all exhibit significant genetic differentiation and low heterozygosity (Monty et al. 2003). Although Eastern Woodrats can occupy forests, swamps, thickets, grasslands, hedgerows, and abandoned buildings (Rainey 1956), the remnant populations in southern Illinois primarily occupy rocky bluffs in parts of Shawnee National Forest. 1Department of Zoology, Southern Illinois University, Carbondale, IL 62901-6501. 2Mathematics and Science Division, Southeastern Illinois College, Harrisburg, IL 62946. 3US Fish and Wildlife Service, 4701 North Torrey Pines Drive, Las Vegas, NV 89130. 4Cooperative Wildlife Research Laboratory, Southern Illinois University, Carbondale, IL 62901. 5Department of Life Sciences, John A. Logan College, Carterville, IL 62918. 6Illinois Department of Natural Resources, One Natural Resources Way, Springfield, IL 62702. 7Department of Biology, Ball State University, Muncie, IN 47306. *Corresponding author - feldhamer@zoology.siu.edu. 2 Southeastern Naturalist Vol. 12, No. 1 Protocols for reintroducing large, long-lived, K-selected species are fairly well established (Fischer and Lindenmayer 2000, Kleiman 1989). However, this is not the case for smaller, r-selected species such as woodrats (Serfrass 2008). Our objectives were to implement and evaluate a recovery project (Bluett 2003) by: 1) reintroducing (sensu IUCN 1998) and establishing Eastern Woodrat populations at selected sites in southern Illinois where they had occurred historically; and 2) monitoring populations at all sites for persistence, dispersal, and reproductive success. Our efforts to establish self-sustaining populations of Eastern Woodrats at previously occupied sites in Illinois were within the context of a general framework for reintroducing r-selected species (Serfrass 2008). Study Sites The Shawnee Hills region of southern Illinois encompasses 16 counties bordered by the Mississippi and Ohio rivers. Much of this region includes the highly fragmented Shawnee National Forest, which is primarily dominated by a variety Figure 1. Extant and release sites of translocated populations of Eastern Woodrats (Neotoma floridana) in southern Illinois. 2013 A.K. Poole, et al. 3 of species of Quercus spp. (oak) and Carya spp. (hickory), with interspersed in-holdings of croplands. Understory often includes Vitis spp. (wild grape), Lonicera japonica L. (Japanese Honeysuckle), and Toxicodendron radicans L. (Poison Ivy). Steep slopes, often with rocky outcrops, occur throughout much of the area, as do permanent and intermittent streams. Methods Selection of release sites All potential reintroduction sites were within the historical range of the species in Illinois. We used ArcView 3.2 and ArcView 8.2 (Environmental Systems Research Institute) to locate potential reintroduction areas and model connectivity at a macro scale. We developed GIS models to identify steep forested areas (Novosak 2004) using forest cover from the Regional Gap Analysis Project National Elevation Dataset (USGS 2002a) and the National Land Cover Dataset (USGS 2002b). The areas of highest value, including those formerly occupied by woodrats in southern Illinois, were buffered at 0.85 and 1.60 km distances to identify suitable release sites and dispersal corridors because we anticipated that forested corridors that provide connectivity between bluff sites would enhance dispersal and population maintenance (Hilty et al. 2006, Lidicker and Patton 1989). We then used ground surveys to confirm the presence of bluff habitat (rocky outcrops) and ranked quantity and quality of food and cover using the methods described in Novosak (2004). Based on suitability of both macrohabitat (bluff habitat and potential connectivity to other sites) and microhabitat (food and cover), we selected four reintroduction sites in 2004: Garden of the Gods (Saline County), Pounds Escarpment/ Rim Rock (Gallatin County), High Knob (Gallatin County), and Buzzard’s Point (Gallatin County). These sites were 2 to 4 km from each other to facilitate natural dispersal of translocated woodrats among sites through corridors of suitable habitat. Castleberry et al. (2001) documented a maximum distance of 704.4 m traveled in one night in the similar N. magister Baird (Allegheny Woodrat), and individuals have been known to travel 1.5 km upon release (LoGuidice 2006). In 2008, we added another reintroduction site based on the same habitat criteria— Lusk Creek (Pope County), approximately 16.5 km southwest of Garden of the Gods (Fig. 1)—to extend the potential dispersal range. Trapping From April 2003 through March 2009, we translocated Eastern Woodrats to southern Illinois from locations in Butler, Carter, Iron, Shannon, and Wayne counties in Missouri, and Cross, Lee, St. Francis, and Woodruff counties in Arkansas. The same subspecies (N. f. illinoensis) occurs in the three states (Hall 1981). Source sites were at similar latitudes as release sites, but did not always include bluff habitat. We set Tomahawk live traps (16.5 x 16.5 x 48 cm; Tomahawk Live Trap Co., Tomahawk, WI) in areas believed to contain woodrats or where sign was evident. Trapping sessions and the numbers of traps operated varied according to area and weather conditions. We baited traps with a mixture of peanut butter, sunflower seeds, cracked corn, and apple, and provided polyester 4 Southeastern Naturalist Vol. 12, No. 1 fiber for bedding. We marked animals with two individually numbered Monel #3 ear tags (Model 1005-3, National Tag and Band Co., Newport, KY). We recorded body mass (nearest g), gender, and reproductive status. We assigned age classes according to body mass: <150 g = juvenile, 150–200 g = subadult, and >200 g = adult (Rainey 1956). Descended (scrotal) testes indicated reproductively active males. A perforated vagina, swollen teats, or lactation was evidence of reproductive activity in females. Using a 3.5-mm biopsy punch, we collected a small amount of ear tissue for future genetic analyses. We held animals in separate cages for the least amount of time feasible prior to release (usually one to two nights maximum) in a dark, quiet room. Woodrats were fed acorns or “rat chow” (Mizuri rodent feed 6F, size M30), as well as apple slices. We transported animals in an enclosed, temperature-controlled vehicle to the release site. Release We did not acclimate woodrats to the reintroduction sites as in a true soft release (Serfrass 2008), but we attempted to minimize the stress on animals from gathering food and nesting material in an unfamiliar area. We constructed artificial nests by placing nest materials—including half-gallon cardboard juice or milk containers, polyester fiber bedding, and covered with sticks—at release sites with about 1 kg of acorns or “rat chow”. Artificial nests were constructed in microhabitats similar to those woodrats use naturally to construct nests, i.e., deep, dry rock crevices, hollow logs, and the base of hollow trees. We then released woodrats in the artificial nests. We maximized post-release population persistence by two to four “pulsed releases” of 6–32 individuals (depending on capture success from source populations) for a total of 50 to 121 individuals at each release site. Releases commenced in early spring after leaf-out when weather became warmer. Subsequent monitoring Approximately 1 month following releases, we used live trapping to monitor translocated animals for survival, reproduction, and movements among and between reintroduction sites. We used the same trapping methods but included large Sherman live traps (13 x 13 x 38 cm; H.B. Sherman Traps, Tallahassee, FL) to facilitate the capture of small juveniles because they can escape from the Tomahawk traps. From September 2011 through March 2012, we visually surveyed surrounding non-agricultural, bluff habitat for woodrat sign to document minimum extent of dispersal. We considered recent scat, fresh food caches, and stick nests as presence of woodrats in an area. From January 2012 through February 2012, we used Tomahawk live traps to document woodrat survival at Lusk Creek. Results From April 2003 through March 2009, we translocated 422 woodrats to Garden of the Gods, Rim Rock/Pounds Escarpment, High Knob, Buzzard’s Point, and Lusk Creek (Tables 1 and 2). The minimum number known alive 2013 A.K. Poole, et al. 5 about 1 month after release at all sites except Lusk Creek was approximately 12.5% based on 47 recaptured woodrats. Because of limited personnel and funding, Lusk Creek was not monitored after initial release. Reproduction occurred based on the capture of 61 “new” (no ear tags or biopsy punch) individuals (Table 3) as well as another 17 unmarked individuals at Lusk Creek taken in 89 trap nights from January through February 2012. Several animals emigrated from reintroduction sites. Based on visual surveys for woodrat sign in bluff areas north and east of release sites during 2011 and 2012, widespread Table 1. Number and gender of translocated Eastern Woodrats (Neotoma floridana) at sites in southern Illinois from April 2003 through March 2009. Site Total number released Number of males Number of females Garden of Gods 121A 39 49 Rim Rock 108 43 65 High Knob 81 26 55 Buzzard’s Point 50B 16 33 Lusk Creek 62 29 33 A33 Unknown sex, data lost. B1 unknown sex, animal escaped. Table 2. Number and date of release of translocated Eastern Woodrats (Neotoma floridana) at sites in southern Illinois from April 2003 through March 2009. Garden of the Gods Rim Rock High Knob Buzzard's Point Lusk Creek April 2003 18 May–June 2003 21 September 2003 7 February 2004 33 23 May 2004 23 February 2005 6 March/April 2005 7 May 2005 6 June 2003 9 July 2003 16 September 2003 11 March 2004 24 April 2004 16 February 2005 1 May 2005 6 April 2005 6 February 2006 19 October 2005 20 February 2006 24 April 2006 14 March 2007 23 February 2007 25 March 2007 25 March 2008 31 February/March 2009 31 6 Southeastern Naturalist Vol. 12, No. 1 dispersal is suggested, extending approximately 8 km northwest of Garden of the Gods, 2.7 km east of Rim Rock, and throughout the area between Garden of the Gods and Lusk Creek (Fig. 2). Discussion There are several possible reasons for the decline and extirpation of Eastern Woodrat populations in southern Illinois (Feldhamer and Poole 2008)—reasons that also likely apply to the decline of closely related Allegheny Woodrats (see LoGiudice 2008). Nawrot and Klimstra (1976) speculated that unusually harsh Table 3. Number of recaptured and “new” (unmarked) Eastern Woodrats (Neotoma floridana) at reintroduction sites in southeastern Illinois from June 2003 through February 2012. Number of Site Number recaptured “new” woodrats Number of trap nights Garden of Gods 12 20 596 Rim Rock 6 9 444 High Knob 22 18 574 Buzzard’s Point 7 14 100 Lusk Creek 0 17 89 Figure 2. Eastern Woodrat (Neotoma floridana) sign (recent scat, fresh food caches, and stick nests) found during visual surveys from September 2011 through March 2012 in southeastern Illinois. 2013 A.K. Poole, et al. 7 winters during 1912 and 1918 could have caused extirpation of colonies in the eastern part of the Shawnee Hills region. Extreme climatic conditions in association with other mortality factors—predation, the nematode parasite Baylisascaris procyonis carried by Procyon lotor L. (Raccoon), irregular mast production, increased density of competitors, continued habitat loss and fragmentation—could have further reduced population numbers and viability. However, this project was based on the assumption that the primary factor in decline was landscape fragmentation and loss of habitat. Our reintroductions were intended to compensate for extirpation of isolated, often small populations because Eastern Woodrats could not naturally recolonize previously occupied sites throughout southern Illinois (Nawrot and Klimstra 1976). Nonetheless, there is more contiguous habitat available now than when the species declined throughout the early decades of the 1900s, during which time the area that now comprises Shawnee National Forest was cleared for farming and grazing (Fralish and McArdle 2009). Whether other potential population mortality factors remain operative is unknown, although occurrence of the raccoon roundworm B. procyonis appears to be negligible in southern Illinois (Birch et al. 1994, Nielson et al. 2007) and particularly on the release sites (Bade et al. 2012). Reintroduced populations have persisted to date and natural reproduction occurred— documented by the 78 unmarked individuals trapped following releases. Only 12.5% of individuals were known to be alive about 1 month after reintroductions, but this certainly is a conservative estimate. Mortality rate was most likely less than 87.5% given the current extent of distribution and reproduction (Fig. 2). We facilitated dispersal by selecting release sites with suitable habitat corridors to increase the probability of establishing a successful metapopulation (Armstrong and Seddon 2008, Wood 2008). Management implications Several species of woodrats are of conservation concern (Feldhamer and Poole 2008). Although previous woodrat reintroductions in Florida (Barbour and Humphrey 1982) and Pennsylvania (Corbett and Shinkle 1997) met with mixed success, our reintroduction was successful based on the extent of natural dispersal and reproduction. Several factors contribute to the success of reintroductions (Bright and Morris 1994, Fischer and Lindenmayer 2000, Kleiman 1989, Wolf et al. 1996), including release in the core of the historical range (although see Lomolino and Channell 1995, Sagarin and Gaines 2002), extent of quality habitat, release of a large number of animals, an omnivorous diet, wild-caught rather than captive-reared source animals, and soft rather than hard release. Wolf et al. (1996) suggested that reproductive potential of the reintroduced species and the duration of the release may be less critical for success. Several of these concepts and practical considerations applied to our project. Although southern Illinois is not in the core of the range for Eastern Woodrats, we released animals at historically occupied sites based on current macro- and microhabitat conditions. We also attempted to release the optimal number of individuals at each site based on available food, shelter, and space. 8 Southeastern Naturalist Vol. 12, No. 1 We maximized post-release population persistence by 2 to 4 “pulsed releases” for a total of 50 to 121 individuals at each release site. All source individuals were wild-caught, and trapping and reintroduction commenced in early spring after leaf-out when weather became warmer. Additionally, sex ratios of released animals at all sites favored females (Table 1). We provided nesting material and food caches to enhance acclimatization and mimic a soft release to the extent possible given that the bluff habitat precluded confining individuals. High rates of dispersal and increased mortality are associated with hard release in some mammalian species (Fritts et al. 1984, Morris et al. 1993, O’Bryan and McCullough 1985). However, because woodrats are highly territorial and dispersal rates are naturally high (Wood 2008), we anticipated many animals would leave the reintroduction sites regardless of hard or soft release. Continued persistence of the reintroduced populations in our study is not assured, and we agree with Dodd and Seigel (1991:343) that “researchers should temper claims of success with a recognition of the need for long-term evaluation.” Nonetheless, given our metapopulation approach with reintroductions of a large number of woodrats over a multi-year time scale, and the documented extent of dispersal from release sites, we believe that the probability of failure due to demographic stochasticity is low, and progress toward improving the conservation status of the Eastern Woodrat in Illinois is encouraging. Acknowledgments This project was funded in part by Federal Aid in Wildlife Restoration Project W- 135-R, a cooperative effort among the US Fish and Wildlife Service, Illinois Department of Natural Resources, and the Department of Zoology at Southern Illinois University Carbondale. D. Woolard, B. Steffen, and J. Hammersley were instrumental in capture, release, and monitoring efforts. We also thank C. Bartman, D. Corgiat, J. Gaffney, M. Hilton, J. Hubert, A. Hulin, J. Kath, J. Kube, R. Lindsay, D. Ludwig, M. Murphy, P. Shelton, and S. Widowski for their assistance. Literature Cited Armstrong, D.P., and P.J. Seddon. 2008. Directions in reintroduction biology. Trends in Ecology and Evolution 23:20–25. Bade, S.R., F.A. Jiménez, A.K. Poole, and G.A. Feldhamer. 2012. Prevalence of Baylisascaris procyonis and implications for reintroduced woodrat populations in southern Illinois. Transactions Illinois State Academy of Science 105:33–39. Barbour, D.B., and S.R. Humphrey. 1982. 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