Neotoma magister - Baird, 1858
Allegheny Woodrat
Other English Common Names: Allegheny woodrat, Appalachian Woodrat
Taxonomic Status: Accepted
Related ITIS Name(s): Neotoma magister Baird, 1857 (TSN 555661)
Unique Identifier: ELEMENT_GLOBAL.2.101808
Element Code: AMAFF08100
Informal Taxonomy: Animals, Vertebrates - Mammals - Rodents
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Mammalia Rodentia Cricetidae Neotoma
Genus Size: C - Small genus (6-20 species)
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Concept Reference
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Concept Reference: Hayes, J. P., and M. E. Richmond. 1993. Clinal variation and morphology of woodrats (Neotoma) of the eastern United States. J. Mamm. 74:204-216.
Concept Reference Code: A93HAY01NAUS
Name Used in Concept Reference: Neotoma magister
Taxonomic Comments: Neotoma magister formerly was regarded as a subspecies of Neotoma floridana. Evidence from analyses of variation in mtDNA, allozymes, and morphology indicates that Neotoma magister is a highly distinct lineage (Hayes and Harrison 1992, Hayes and Richmond 1993, Edwards and Bradley 2001). Baker et al. (2003) and Musser and Carleton (in Wilson and Reeder 2005) recognized N. magister and N. floridana as distinct species.

Baker et al. (2003) used the name "Appalachian woodrat."
Conservation Status
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NatureServe Status

Global Status: G3G4
Global Status Last Reviewed: 17Apr2006
Global Status Last Changed: 05Sep1996
Rounded Global Status: G3 - Vulnerable
Reasons: Fairly large range in the eastern United States, but extirpated or declining over about 35% of the range; still common in other areas; decline has been rapid and the cause is not yet fully understood.
Nation: United States
National Status: N3N4 (04Feb2005)

U.S. & Canada State/Province Status
United States Alabama (S3), Connecticut (SH), District of Columbia (SH), Indiana (S2), Kentucky (S3S4), Maryland (S1), New Jersey (S1), New York (S1), North Carolina (S2S3), Ohio (S1), Pennsylvania (S3), Tennessee (S3), Virginia (S3), West Virginia (S3)

Other Statuses

IUCN Red List Category: NT - Near threatened

NatureServe Global Conservation Status Factors

Range Extent: 200,000-2,500,000 square km (about 80,000-1,000,000 square miles)
Range Extent Comments: The range extends from western Connecticut (formerly), southeastern New York (virtually extirpated), northern New Jersey, and northern Pennsylvania southwestward through western Maryland, Tennessee, Kentucky, West Virginia, and northern and western Virginia (from the Blue Ridge westward) to northeastern Alabama (observed in several cave systems) and northwestern North Carolina (Hall 1981), with isolated populations north of the Ohio River in southern Ohio (where recent surveys failed to locate this species; W. Peneston, pers. comm., cited by Mengak 2002) and southern Indiana (Whitaker and Hamilton 1998). Although Hall (1981) showed N. magister in the northwestern corner of Georgia, the Tennessee River is generally accepted as the southern range limit.

Number of Occurrences: 81 to >300
Number of Occurrences Comments: This species is represented by a very large number of occurrences or subpopulations (probably more than 200).

Population Size: 10,000 - 1,000,000 individuals
Population Size Comments: Total adult population size is unknown but likely is much larger than 10,000.

Viability/Integrity Comments: The number of occurrences or subpopulations with good viability probably is substantially smaller than the total number of extant occurrences.

Overall Threat Impact: High
Overall Threat Impact Comments: Possibly widespread deforestation and habitat fragmentation contributed to the initial decline of the species, isolating populations and eliminating dispersal and travel corridors (Sciascia, pers. comm., 1994). Deforestation and associated reduction/elimination of food resources currently may be a threat to some local woodrat populations. Causes of the continuing decline are not yet fully understood, but some hypotheses have been offered. Probably the explanation lies in a combination of these and other factors that may differ locally in importance (Linzey 1990).

Parasitism by the raccoon roundworm (Baylisascaris procyonis) has been identified as a significant cause of mortality and a probable major factor in the decline in several states (McGowan 1993; Hicks, pers. comm., 1994; Johnson et al. 1997; LoGiudice 2000). Although rarely fatal to raccoons, infestation may cause cerebrospinal nematodiasis in other species and has caused declines in some populations (Kazacos 1983). New York released and monitored woodrats in formerly occupied areas and all the animals died (50 total, including the released adults and their progeny); 11 of the 13 recovered carcasses were infected by the roundworms (McGowan 1993; Hicks, pers. comm., 1994). Stone observed that woodrat decline in New York coincided with a marked increase in raccoon numbers (Linzey 1990). Hayes (unpublished research proposal) suggested that woodrats may be especially vulnerable for two reasons: 1) among other novel items, woodrats are known to carry back to their nests the feces of other animals, which might include raccoon scats contaminated with B. procyonis eggs, and 2) a "relatively long generation time increases the probability that individuals will become infested prior to reproduction." Both of these reasons recently have been documented (McGowan, pers. comm., 1994). Additionally, McGowan (pers. comm., 1994) found that woodrats may colonize areas where an infected woodrat has recently died, thus perhaps maintaining spread of the parasite.

Hall (1988) noticed a correlation between the spread of the gypsy moth (Lymantria dispar) and loss of woodrats in Pennsylvania. "There is a possibility that acorns make up a significant part of the food supply of the species in Pennsylvania. Acorns may be important as a winter food source since they can be stored for long periods of time. Studies conducted by the Pennsylvania Game Commission show that acorn production drops to zero for several years following defoliation. Gypsy moth infestation has resulted in considerable mortality of oaks, especially along the poorer soils of rocky ridge tops. These are the areas where many rocky sites are found, which are ideal woodrat habitats." In support of his argument, Hall noted that at three other sites where woodrats survived, other winter food sources were available.

In a similiar theory, McGowan (pers. comm., 1994) speculated that the 1930-1940s permanent loss of the American chesnut (Castanea dentata) may be a factor. This chestnut was extremely hardy and a bountiful producer of mast, and its previous distribution essentially mirrors the historic distribution of the woodrat. Loss of this stable and predictable food source may have led to the continuing woodrat population decline.

Nawrot and Klimstra (1976) argued that the disappearance of a similiar species, N. floridiana illinoensis, was likely due to two unusually severe winters. "Despite a tolerance of Neotoma to many environmental conditions, severe winter weather appears to be a major factor in sudden woodrat population declines. Fitch and Rainy (1956) recorded a drastic decline in Neotoma floridiana abundance in Kansas after two winters of below-average temperatures and prolonged snow and ice cover. Significantly, this decimated population was unable to make substantial gains and continued to decline gradually for several years." Weather also has been proposed as a factor in Neotoma magister decline, a view that is supported by the fact that the decline has been in the northern part of the range. Records in Pennsylvania and West Virginia over the past thirty years indicate "below normal" temperatures in critical months accompanying the observed shrinkage in range (Linzey, pers. comm., 1994).

In some areas, including caves popular with spelunkers, human disturbance has been implicated in the disappearance of woodrat populations (e.g., Kirkland 1986). These woodrats do seem to avoid areas of human habitation or heavy human use, but many sites where they have disappeared are remote and rarely visited by people.

Habitat is generally inaccessible and undesirable for development, but strip mining of coal and limestone is a potential threat in many areas.

Monty and Feldhamer (2002) stated that increased predation by great horned owls has been proposed as a threat, but owl predation seems highly unlikely as a reason for the decline.

Populations often are small and isolated, hence highly susceptible to extirpation (Mengak, pers. comm., 1994).

Short-term Trend: Decline of 10-30%

Long-term Trend: Decline of 30-50%
Long-term Trend Comments: Populations in the northeastern United States have declined, while those in the southern portion of the range appear to have remained relatively stable. However, woodrats appear to be rare in Tennessee (Kennedy and Harvey 2001), and few specimens have been reported from Alabama, where present status is poorly known (T. H. Henry).

The species has disappeared from Connecticut, New York, most of eastern Pennsylvania, and all but one site in New Jersey. This decline has apparently been occurring for many years. Early sources (e.g., Newcombe 1930, Poole 1940) remarked on the woodrat's decreasing range and abundance, and the most recent reports from Connecticut date from the 1930s (Goodwin 1935). Such a decline might be expected in the wake of human settlement, but in recent decades woodrat decline has been dramatic in New York, New Jersey, Pennsylvania, and Maryland. Apparent extirpation has occurred in at least 100 sites in northern Virginia (Buhlmann, pers. comm., 1992).

Since the late 1960s woodrats in Pennsylvania have disappeared from sites where they once occurred, particularly in eastern and northwestern Pennsylvania; surveys of more than 360 sites from which woodrats have been reported yielded 20 metapopulations, but five of these metapopulation areas no longer support woodrats, and seven of them have fewer active colonies than in the past (C. Fergus, no date; www.pgc.state.pa.us, accessed April 2006). The species has disappeared from the southeastern portion of Pennsylvania and has declined in much of the rest of the state; at present, populations exist throughout much of Pennsylvania's southcentral and southwestern counties, with a few remnant populations in eastern counties; mapped distribution indicates that the species no longer occurs in roughly half of the historically occupied range (Pennsylvania Game Commission website, woodrat information last updated in 2006).

In New Jersey, several populations have been extirpated for at least 20 years; the single known remaining population has been relatively stable in recent years, based on trapping results in 1999-2001 (New Jersey Division of Fish & Wildlife).

Woodrats were documented throughout the historical range in southeastern New York as recently as the mid-1960s, when they appeared to occupy all available habitat; those populations were extirpated by 1987; currently, the only woodrats in the state are immigrants that occasionally occupy a small patch of habitat on the NewYork-New Jersey border (this is the northern extreme of the habitat for the last remaining New Jersey woodrat population) (New York State Comprehensive Wildlife Comservation Strategy, 2005 revised draft). Reintroductions attempted in the early 1990s were unsuccessful.

This species may be represented by only one population in Ohio, or possibly it is extirpated; recent surveys failed to locate woodrats there (W. Peneston, pers. comm., cited by Mengak 2002).

In Maryland, woodrat populations have been in decline for over two decades and continue to decline (Dan Feller, Maryland Natural Heritage Program, pers. comm., cited by New York State Comprehensive Wildlife Conservation Strategy, 2005 draft).

In the early 1980s, extant populations in Indiana were reported from 20 sites (18 bluffs, 2 caves) adjacent to the Ohio River; in 1991-1992, live trapping surveys at 17 sites yielded a total of 101 woodrats at 12 (11 bluffs, 1 cave) of the 17 sites (Johnson 2002). Abundance in Indiana appears to have declined in recent decades. Population density for 6 sites in Indiana averaged 27.5/km of cliff habitat during the early 1980s (Cudmore 1985). During the 1990s density averaged about 11 individuals/km at 12 Indiana sites (Johnson et al. 1997).

Woodrat populations in the core of the range in West Virginia (Stihler and Wallace 1996, Castleberry 2000), Virginia (Mengak 2000), and Kentucky (Thomas 1998) currently appear to be secure.

Intrinsic Vulnerability: Moderately vulnerable

Other NatureServe Conservation Status Information

Inventory Needs: Rangewide surveys are needed to better determine status and trends.

Protection Needs: Areas with woodrats should be protected from excessive human traffic. Mining and logging operations should not be permitted in areas with woodrat populations.

Distribution
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Global Range: (200,000-2,500,000 square km (about 80,000-1,000,000 square miles)) The range extends from western Connecticut (formerly), southeastern New York (virtually extirpated), northern New Jersey, and northern Pennsylvania southwestward through western Maryland, Tennessee, Kentucky, West Virginia, and northern and western Virginia (from the Blue Ridge westward) to northeastern Alabama (observed in several cave systems) and northwestern North Carolina (Hall 1981), with isolated populations north of the Ohio River in southern Ohio (where recent surveys failed to locate this species; W. Peneston, pers. comm., cited by Mengak 2002) and southern Indiana (Whitaker and Hamilton 1998). Although Hall (1981) showed N. magister in the northwestern corner of Georgia, the Tennessee River is generally accepted as the southern range limit.

U.S. States and Canadian Provinces
Color legend for Distribution Map
Endemism: endemic to a single nation

U.S. & Canada State/Province Distribution
United States AL, CT, DC, IN, KY, MD, NC, NJ, NY, OH, PA, TN, VA, WV

Range Map
Note: Range depicted for New World only. The scale of the maps may cause narrow coastal ranges or ranges on small islands not to appear. Not all vagrant or small disjunct occurrences are depicted. For migratory birds, some individuals occur outside of the passage migrant range depicted. For information on how to obtain shapefiles of species ranges see our Species Mapping pages at www.natureserve.org/conservation-tools/data-maps-tools.

Range Map Compilers: NatureServe, 2003


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Jackson (01071), Madison (01089)*, Marshall (01095)
IN Crawford (18025), Harrison (18061), Martin (18101)*, Monroe (18105), Orange (18117)*
MD Allegany (24001), Frederick (24021), Garrett (24023), Montgomery (24031)*, Washington (24043)*
NC Ashe (37009), Avery (37011), Buncombe (37021), Burke (37023), Caldwell (37027), Iredell (37097), McDowell (37111), Mitchell (37121), Watauga (37189), Wilkes (37193), Yancey (37199)
NJ Bergen (34003), Morris (34027)*, Passaic (34031)*, Sussex (34037)*
NY Orange (36071), Rockland (36087), Ulster (36111)
OH Adams (39001)
PA Bedford (42009), Berks (42011)*, Blair (42013), Cambria (42021), Cameron (42023), Carbon (42025), Centre (42027), Clinton (42035), Cumberland (42041), Dauphin (42043), Fayette (42051), Franklin (42055), Fulton (42057)*, Huntingdon (42061), Indiana (42063), Juniata (42067), Lancaster (42071)*, Lebanon (42075), Lehigh (42077)*, Luzerne (42079)*, Lycoming (42081), Mifflin (42087), Northampton (42095), Perry (42099), Schuylkill (42107), Somerset (42111), Tioga (42117), Union (42119)*, Venango (42121)*, Westmoreland (42129)
TN Anderson (47001), Bedford (47003), Campbell (47013), Cannon (47015), Carter (47019)*, Cheatham (47021)*, Coffee (47031), Cumberland (47035), Davidson (47037)*, DeKalb (47041), Fentress (47049), Franklin (47051), Grundy (47061), Hamilton (47065)*, Hancock (47067), Hawkins (47073), Hickman (47081)*, Humphreys (47085)*, Jackson (47087), Johnson (47091), Marion (47115), Maury (47119), Morgan (47129), Overton (47133), Perry (47135)*, Pickett (47137), Putnam (47141), Rhea (47143), Rutherford (47149), Scott (47151)*, Sequatchie (47153), Smith (47159), Unicoi (47171), Van Buren (47175), Warren (47177), Washington (47179), Wayne (47181), White (47185), Wilson (47189)
WV Berkeley (54003), Boone (54005), Fayette (54019), Grant (54023), Greenbrier (54025), Hampshire (54027), Hardy (54031), Kanawha (54039), McDowell (54047), Mercer (54055), Mineral (54057), Mingo (54059), Monongalia (54061), Monroe (54063), Morgan (54065), Nicholas (54067), Pendleton (54071), Pocahontas (54075), Preston (54077), Raleigh (54081), Randolph (54083), Summers (54089), Tucker (54093), Wayne (54099)*, Webster (54101)*, Wyoming (54109)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
02 Rondout (02020007)+, Hudson-Wappinger (02020008)+, Lower Hudson (02030101)+, Hackensack-Passaic (02030103)+, Middle Delaware-Mongaup-Brodhead (02040104)+, Middle Delaware-Musconetcong (02040105)+, Lehigh (02040106)+, Schuylkill (02040203)+*, Upper Susquehanna-Tunkhannock (02050106)+*, Upper West Branch Susquehanna (02050201)+, Sinnemahoning (02050202)+, Middle West Branch Susquehanna (02050203)+, Bald Eagle (02050204)+, Pine (02050205)+, Lower West Branch Susquehanna (02050206)+*, Lower Susquehanna-Penns (02050301)+, Upper Juniata (02050302)+, Raystown (02050303)+, Lower Juniata (02050304)+, Lower Susquehanna-Swatara (02050305)+, Lower Susquehanna (02050306)+*, South Branch Potomac (02070001)+, North Branch Potomac (02070002)+, Cacapon-Town (02070003)+, Conococheague-Opequon (02070004)+, Middle Potomac-Catoctin (02070008)+*, Monocacy (02070009)+, Middle Potomac-Anacostia-Occoquan (02070010)+*
03 Upper Yadkin (03040101)+, South Yadkin (03040102)+, Upper Catawba (03050101)+
05 Middle Allegheny-Tionesta (05010003)+*, Conemaugh (05010007)+, Kiskiminetas (05010008)+, Tygart Valley (05020001)+, Upper Monongahela (05020003)+*, Cheat (05020004)+, Lower Monongahela (05020005)+, Youghiogheny (05020006)+, Upper New (05050001)+, Middle New (05050002)+, Greenbrier (05050003)+, Lower New (05050004)+, Gauley (05050005)+, Upper Kanawha (05050006)+, Elk (05050007)+, Lower Kanawha (05050008)+, Coal (05050009)+, Upper Guyandotte (05070101)+, Tug (05070201)+, Twelvepole (05090102)+*, Ohio Brush-Whiteoak (05090201)+, Lower White (05120202)+, Lower East Fork White (05120208)+, Upper Cumberland (05130101)+, South Fork Cumberland (05130104)+, Obey (05130105)+, Upper Cumberland-Cordell Hull (05130106)+, Collins (05130107)+, Caney (05130108)+, Lower Cumberland-Old Hickory Lake (05130201)+, Lower Cumberland-Sycamore (05130202)+*, Stones (05130203)+, Blue-Sinking (05140104)+
06 South Fork Holston (06010102)+, Watauga (06010103)+, Holston (06010104)+, Upper French Broad (06010105)+, Nolichucky (06010108)+, Upper Clinch (06010205)+, Emory (06010208)+, Middle Tennessee-Chickamauga (06020001)+, Sequatchie (06020004)+, Guntersville Lake (06030001)+, Wheeler Lake (06030002)+, Upper Elk (06030003)+, Lower Tennessee-Beech (06040001)+, Upper Duck (06040002)+, Lower Duck (06040003)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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General Description: A large grayish-brown rat with white underparts, large eyes and ears, and a long, furry, bicolored tail. Godin (1977) described the coloration as follows: "In winter the coloration above is buffy gray to pale cinnamon, heavily overlaid with black. The head and the sides of the body are buffy gray, while the axillae, or armpits, are creamy buff. The throat, belly, and feet are white, the fur being white to the roots except along the sides of the belly, where the basal color is pale leaden gray. The tail is sharply bicolored, blackish brown above, white below. The summer pelage is slightly paler and shorter. Immatures are grayer than adults, particularly on the belly."

Total length 35.5-44.7 cm; tail length 16-21 cm; mass 394-500 g (Godin 1977).

The skull is elongated, about 5.0 cm long and 2.5 cm at its greatest width, with "marked interorbital constriction and relatively small auditory bullae. The palate, lacking a posterior median spine, ends about even with the front of the first molar, which is high, flat-crowned, and prismatic. The coronoid process of the mandible is well developed" (Godin 1977).

The sexes are similar in color, and males average a little larger than females. Both sexes have a ventral skin gland which produces a strong, musky odor, especially in breeding males. External genitalia are generally apparent in the males, and females have four mammae.

There are numerous excellent published illustrations of woodrats and their skulls, including Schwartz and Schwartz (1959), Barbour and Davis (1974), Lowery (1974), Whitaker (1980), Wiley (1980), Hall (1981), Hall (1985), and Merritt (1987). Tracks are illustrated in some of these works, and tracks and scats are shown in Murie (1974).

Diagnostic Characteristics: Neotoma magister differs from the Norway rat (Rattus norvegicus) in having a furry, not scaly, tail; longer, softer fur; larger naked ears; and larger bright eyes. "The molars have enamel arranged in prismatic folds and are thus easily distinguished from the tuberculate molars of Rattus " (Adams 1987).
Reproduction Comments: The breeding season is late winter to late summer, with the young born from March to September (Poole 1940, Merritt 1987). Gestation period is about 35 days (32 to 38 days) (Birney 1973). Females normally mate again soon after giving birth, and two or three litters of usually two to four young are produced annually (Poole 1940, Schwartz and Schwartz 1959). Sexual maturity is reached in less than a year, and some early-born females (but not males) breed in the season of their birth (Wiley 1980).

Woodrats are thought to live longer than most small rodents. One female is known to have lived more than three years in the wild (Fitch and Rainey 1956). However, mortality is normally high.

Ecology Comments: Woodrats are basically solitary and unsociable, frequently fighting one another. Each lives alone in its house, except when breeding and raising young. Poole (1949) found "a great deal of individuality" in temperament, behavior, food preferences, etc., among his captive Allegheny woodrats. When upset, woodrats may chatter their teeth or stomp their hind feet (Wiley 1980).

Near the nests are found piles of sticks and trash called middens. Part of the materials found in woodrat houses and middens appears to be cached food in the form of nuts, seeds, berries, cuttings of vegetation, and mushrooms. But there may also be miscellaneous bits of trash, including rags, bits of metal, bones, pieces of glass, paper, etc. Newcombe (1930) and Poole (1940) compiled lists of such materials. The function of this compulsive collecting is unknown.

Scats are deposited in special latrine areas, apparently used by more than one individual, or over a long period of time. Poole (1940) reported seeing such heaps measuring 45 x 25 x 5 cm. A captive kept by Patterson (1933) voided 3 to 84 fecal pellets per day. Similarly, urinating spots are used, leaving dark stains 15-20 cm in diameter on rocks (Schwartz and Schwartz 1959).

Though solitary and territorial, woodrats most often occur in clusters due to patchiness of the rock outcrop, talus, and cave habitat, and conform to the concept of a metapopulation (Hassinger et al. 1996). Home ranges are small, 0.26-0.6 ha (Wright and Hall 1996); usually less than about 90 m across (Burt and Grossenheider 1976). Poole (1940) reported movements of 183 m and 92 m by two woodrats in Pennsylvania. Foraging movements, while often focused within rock habitat, may extend beyond the protection of rocks up to 160 meters from the den site (Wright and Hall 1996). Den shifts tend be less than 100 meters with a median of 40 m (Wright 1998), and woodrats, particularly females, often live their entire lives in the same outcrop (Feller, pers. obs., 1998). There are reports of large unidirectional movements of displaced woodrats, e.g., 1 km and 4 km (McGowan 1993), as well as naturally dispersing individuals, 0.3-1 km (McGowan 1993), 1 km (Feller, pers. obs., 1995), and up to 6 km (Wright, pers. comm., 1998). While woodrats can travel long distances between patches, as distances increase, the chance of successful emigration between patches is likely to decrease, particularly in the absence of protective rock crevices. Barriers to dispersal are not clearly known, as woodrats have been documented to traverse seemingly inhospitable terrain, including roads, small streams, and small fields, though movements are largely within rock habitat (Feller, pers. obs.; Mengak, pers. comm., 1998; Wright, pers. comm., 1998). However, woodrats display unwary behavior when crossing roads (Feller, pers. obs.), and roadkills have been documented (Feller, pers. obs., 1993; McGowan 1993).

Cudmore (1983) estimated population densities of 8.3 to 71.9 woodrats per 1000 m of cliff along the Ohio River in Indiana. According to Burt and Grossenheider (1976), populations of 5-8 adults per ha are probably high.

Predators include owls, skunks, weasels, foxes, raccoons, bobcats and large snakes. Interestingly, examination of the stomach contents of 70 rattlesnakes (CROTALUS HORRIDUS) and 15 copperheads (AGKISTRODON CONTROTRIX) in Pennsylvania and New Jersey revealed no woodrat remains, even though many of the snakes came from areas inhabited by woodrats (Poole 1940). Humans have been killing woodrats for thousands of years--first for food, and much later out of prejudice, because of a resemblance to European rats.

White-footed mice (PEROMYSCUS LEUCOPUS) compete with woodrats for food (Rainey 1956), and in Indiana, opossums (DIDELPHIS VIRGINIANA), raccoons (PROCYON LOTOR) and turkey vultures (CATHARTES AURA) may compete for den sites (Cudmore 1983). Woodrats support many ecto- and endoparasites, including fleas, ticks, mites, chiggers, botflies (Diptera: Cuterebridae), nematodes, and tapeworms (Murphy 1952, Cudmore 1986). Bubonic plague (Schwartz and Schwartz 1959) and rabies (Dowda et al. 1981) have been reported in wild woodrats.

Non-Migrant: Y
Locally Migrant: N
Long Distance Migrant: N
Mobility and Migration Comments: Patterns of genetic (DNA) variation indicate that gene flow can be low among subpopulations of Neotoma magister and that effective dispersal is limited among subpopulations separated by as little as 3 km (Castleberry et al. 2002).

Thomas (2001, cited by Monty and Feldhamer 2002) reported that in Kentucky an adult male moved 3,615 m from the location of his previous capture 49 days earlier. The movement occurred between November and January. The individual remained at the new site for at least 1 year. The longest movement recorded for a female was 405 m. The longest movement between 2 consecutive days of capture was 638 m by an adult male.

Palustrine Habitat(s): Riparian
Terrestrial Habitat(s): Bare rock/talus/scree, Cliff, Forest - Conifer, Forest - Hardwood, Forest - Mixed, Shrubland/chaparral, Woodland - Conifer, Woodland - Hardwood, Woodland - Mixed
Habitat Comments: Typical habitat is rocky cliffs and talus slopes. These woordrats make midden mounds and stick piles among rocks, but secluded nest sites generally are not within stick houses (see Hayes and Harrison 1992).

Throughout its range, this species associated with extensive rocky areas such as outcrops, cliffs, talus slopes with boulders and crevices, and caves. It occasionally uses abandoned buildings but generally avoids humans. It generally occurs at higher elevations (to about 1000 m) and is rarely found in lowlands or open areas.

In southern New York, New Jersey, and adjacent Pennsylvania, woodrat habitat "consists of extensive boulder fields at the base of ridges with rock outcrops. These talus slopes consist of large boulders (10-20 ft. [3-6 m] across) piled in several layers. Neotoma lives among the cave like spaces formed by the piled boulders" (Sciascia 1990).

Referring to the mountainous area of Pennsylvania, Merritt (1987) wrote, "limestone caves, rocky cliffs and accumulations of residual sandstone boulders marked by deep crevices with underground galleries represent favored habitat." Hall (1985) pointed out that good habitat is found "specifically at water gaps where cliff faces and boulder piles are usually abundant." Unpublished data from Pennsylvania Natural Diversity Inventories (eastern and western offices) make frequent references to woodrat occurrences in sandstone, limestone, or shale outcrops and cliffs, usually with crevices or abundant boulders; also solution caves and abandoned limestone quarries and mines. Associated forest is varied, including cove hardwoods, hemlock-birch, oak-pine, and various combinations of oaks, maples, hickories, beech, and yellow poplar (tulip-tree). Grape, mountain laurel, rhododendron and ferns are frequently mentioned.

In West Virginia, woodrats are common in caves, rock shelters, outcrops with deep crevices, and riverbanks with an abundance of sandstone rocks and boulders.

"In Maryland, this species is found predominately in cliffs, caves and rocky areas in the three western-most counties, and along cliffs of the Potomac River to the vicinity of Washington, D.C." (Feldhammer et al. 1984). In western Maryland "Pottsville Sandstone outcrops [provide] massive, blocky boulders and extensive cliffs with numerous crevices and miniature cave-like situations" (Thompson 1984).

In Indiana, extant populations are restricted primarily to south-facing limestone bluffs on the Ohio River, where there are den sites in the rock and dense red-cedar (Cudmore 1983, Scott Johnson, pers. comm.).

In Kentucky, "cliffs with deep crevices, caves, or large boulders piled in such a way as to form numerous retreats and shelters are favored" (Barbour and Davis 1974). In Tennessee, "rocky cliffs, caves and fissures or tumbled boulders on the sides of mountains are the preferred habitat" (Hamilton 1943), and in North Carolina it is "rocky places and abandoned buildings at elevations above 3000 feet (900 m)" (Adams 1987). Castleberry et al. (2001) suggest that forest clearcutting has minimal impact on woodrat movements, home range, and habitat use, as long as sufficient intact forest is retained adjacent to colonies. They mention, however, that harvesting methods that selectively remove important mast-producing tree species may represent the greatest threat from forest management.

A large house of sticks, leaves, and miscellaneous debris is built, usually within a cave, crevice, or other well-protected place. This may be a mound like a muskrat house (typical construction in the range of other subspecies), but is more often open, giving the impression of a large bird's nest (Poole 1940). Outside diameter is about 35-60 cm (Patterson 1933) and the inner cavity is about 12 cm across (Poole 1940). The nest is lined with shredded bark of grape, red cedar, hemlock, or basswood, grass, fur, rootlets, and sometimes feathers (Poole 1940, Merritt 1987).

Adult Food Habits: Frugivore, Granivore, Herbivore
Immature Food Habits: Frugivore, Granivore, Herbivore
Food Comments: Woodrats are primarily vegetarian, and food preferences vary widely among individuals. Leaves, twigs, fruits, and seeds of many plants are eaten. Newcombe (1930) thought fungi were a significant part of the diet. Acorns are probably an important food, and in Tennessee Goodpaster and Hoffmeister (1952) found quantities of honey locust (GLEDITSIA TRIACANTHUS) seed pods in caches. New Jersey woodrats appeared to be using seed pods of royal Paulownia (PAULONIA TOMENTOSA) for winter food (Beans 1992). Other foods recorded include twigs and seeds of hemlock (TSUGA CANADENSIS) and birch (BETULA LENTA); leaves of rhododendron; chestnuts (CASTANEA DENTATA); fruits of dogwood (CORNUS FLORIDA), apple (PYRUS MALUS) black cherry (PRUNUS SEROTINA), mountain ash (SORBUS AMERICANA) (Poole 1940), pokeberry (PHYTOLACCA sp.), poison ivy (TOXICODENDRON RADICANS) (Linzey and Linzey 1968), grape, and partridgeberry (MITCHELLA REPENS); leaves and shoots of polypody fern (POLYPODIUM VULGARE), basswood (TILIA AMERICANA), black gum (NYSSA SYLVATICA), teaberry (GAULTHERIA PROCUMBENS), blackberry (RUBUS sp.), white pine (PINUS STROBUS), trailing arbutus (EPIGAEA REPENS), mountain maple (ACER SPICATUM) serviceberry (AMELANCHIER sp.) (Heisler 1941), tree-of-heaven (AILANTHUS ALTISSIMA), Virginia creeper (PARTHENOCISSUS QUINQUEFOLIA), and red cedar (Cudmore 1983). Woodrat cuttings noted in rocky riverbank habitat in West Virginia included asters, balsam squaw-weed (SENECIO PAUPERCULUS), riverbank goldenrod (SOLIDAGO RACEMOSA), tasselrue (TRAUTVETTARIA), violet (VIOLA sp.), and ferns (Norris 1991). Animal food (insects, etc.) is rarely consumed. Woodrats eat about five percent of their weight in dry matter daily (Schwartz and Schwartz 1959).

Water consumption varies greatly among individuals (Poole 1940). Though some captive woodrats drink often, others drink little or nothing. Apparently some woodrats obtain their water from dew or succulent vegetation or from metabolism of foods (Murphy 1952).

Adult Phenology: Nocturnal
Immature Phenology: Nocturnal
Phenology Comments: Primarily nocturnal; most active during first few hours of darkness; less active on moonlit nights (Wiley 1971); more active on cloudy, rainy nights than on clear ones (Rainey 1956, Tate 1970). Active throughout the year but tends to remain "indoors" during bad weather (Poole 1940).
Length: 44 centimeters
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Additional surveys and monitoring are needed throughout the range to better understand status and trends. Appropriate management cannot be implemented until we better understand the causes of woodrat decline.
Restoration Potential: It would be futile to attempt restoration of Allegheny woodrats to areas from which they have disappeared without understanding the causes of that extirpation, and without determining that those causes are no longer present. In most cases today there are too few data to proceed with restocking.

Ohio Department of Natural Resources, Division of Wildlife, Columbus, Ohio, undertook a project in the early 1980s to reintroduce the eastern woodrat. Objectives were to develop reintroduction techniques and to restore a population in a portion of its former range (Schlie and Bookhut 1985). The translocation of West Virginia woodrats to New York was not a restoration effort (see MGMT.RSRCH.PROG).

Preserve Selection & Design Considerations: Aside from the need for favorable habitat and food resources and protection from excessive human traffic, no specific guidelines are available at the present time.
Management Requirements: Crevices and openings in rock or among boulders must be present for den sites, and in some areas this will be the limiting factor. Free water seems not to be a requirement of wild woodrats but may be utilized by them.
Food supplies can be enhanced by planting or managing for native species that provide fruits, seeds, and nuts. Where oaks have been hit hard by gypsy moth, it may be important to assure alternative winter food sources. Gypsy moth defoliation can be prevented and in some contexts with minimal nontarget impacts. Specifically, managers should consider BTK if there is not a potential to eradicate rare Lepidoptera, and "Gypchek" if rare Lepidoptera are present (Schweitzer, 2004, NatureServe online document).

Human intrusion in nesting areas must be discouraged. According to Kirkland (1986), "Although it would be difficult to document a direct cause and effect relationship between the decline of the woodrat and its intolerance of human contact, any management efforts to preserve the eastern woodrat should at least consider this possibility and incorporate into recovery plans safeguards to minimize contact between humans and woodrats."

Pennsylvania Game Commission has formulated provisional protection guidelines, pending more specific guidance that should come out of current research (Hassinger and Dunn 1989). These guidelines are:

1. All caves and limestone mines on public land having either NEOTOMA and/or bats in residence (seasonally or all year) should be designated as "no admittance: restricted areas." Caves with a history of public use or easy accessibility should be gated or fenced to reinforce the "restricted area" concept.

2. Contiguous woodrat habitat, with NEOTOMA occupying any portion thereof, should be protected from any surface disturbance or other form of fragmentation.

3. No surface disturbance should occur within 200 meters [660 ft] (the primary foraging zone) of active colonies.

4. A diversity of mature, mast-producing trees (and all evergreens) should be reserved overtopping and within 50 meters [165 ft] of contiguous woodrat habitat with NEOTOMA occupying any portion thereof.

5. No tree cutting should occur within 200 meters [660 ft] of the "center" of active colonies. Logging roads should be excluded from this zone.

6. If a streambottom occurs within 400 meters [1320 ft] of an active colony, a minimum disturbance corridor (no surface mining, no clearcutting...) of 100 meters [330 ft] in width or wider should connect the colony site to the stream corridor.

7. Blasting - attendant mining - should never be so close as to shift rocks within the colony site.

Restoration efforts should include well-designed experiments that will yield information on possible causes of the decline.

Monitoring Requirements: Occupied habitat should be mapped and a low-impact monitoring program maintained. Regular monitoring is needed in the northern part of the range.

In their annual monitoring of selected woodrat sites, Pennsylvania researchers use the following protocol (Dunn et al. 1989):

A. Habitat is mainly rock piles and cliffs with small openings.

1. Trapping is done between mid-September and mid-October.

2. Weather should be favorable, no storms and temperature normal.

3. Forty traps should be set throughout site, which should be at least one-half acre [2020 sq m] in size.

4. Bait is sliced apple.

5. Live traps are set for two consecutive nights.

6. Traps are checked in the morning (soon after sunrise), then closed for the remainder of day before being reset before sunset.

7. Each captured individual is marked with an ear tag.

8. All animals caught are sexed, weighed, and checked for reproductive condition.

B. Habitat is a cave.

The procedure for monitoring woodrats at a cave site is same as for a rock pile, with 20 traps to be set within the cave and 20 traps outside the cave, surrounding the entrance.

Rather than live-trapping, New Jersey is using a non-intrusive method to monitor its one known population. "By initially collecting the droppings at all locations and periodically resurveying each site and collecting and recording new droppings, we have been able to monitor the level of woodrat activity throughout the summer. In addition to allowing us to monitor the population, the location, size and type of droppings may reveal important information on habitat preferences, productivity, dispersal, food habits and parasites" (Sciascia 1990).

Johnson (1991) described how such methods may also be used in Indiana: "Woodrat populations may also be surveyed by monitoring activity at nests, latrine sites, or communal toilets... In Indiana, active woodrat nests, houses, and latrine sites will be numbered, marked, and plotted on topographic maps. Detailed sketches of the caves and bluff systems will also be used to mark the precise location of nests, houses, and latrine sites for later identification."

In Indiana, status surveys employ live-trapping. "Sites will be visited first to map suitable habitat and search for woodrat nests/houses, piles of cut vegetation, fresh droppings, or other sign... At each site, 40-45 collapsible Tomahawk live-traps (No. 201) will be set and checked for 2 consecutive nights (80-90 total trap-nights). Fewer traps may be used at smaller sites or those with marginal habitat. Traps will be placed near active woodrat sign or throughout suitable habitat and baited with apple slices, peanut butter, and rolled oats. Captured woodrats will be marked with 2 Monel #1 ear tags, examined, and released at the capture site. Sex, age class, reproductive status (females only), standard body measurements, pelage color, and general condition will be noted for each woodrat" (Johnson 1991).

Management Research Needs: The outstanding research need is to discover the causes of the decline in the northeastern part of the range. We also need to learn much more about basic ecology, including preferred foods, home range, and specific habitat requirements.
Population/Occurrence Delineation
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Group Name: Small Murid Rodents

Use Class: Not applicable
Minimum Criteria for an Occurrence: Evidence of historical presence, or current and likely recurring presence, at a given location. Such evidence minimally includes collection or reliable observation and documentation of one or more individuals in appropriate habitat where the species is presumed to be established and breeding.
Mapping Guidance: Separate sites separated by less than 1000 meters should be mapped as separate polygons.
Separation Barriers: Barriers include: wide highways with heavy traffic (subjective determination) and highways with continuous solid barriers that prevent rodent passage; major water bodies, arbitrarily set at those greater than 50 meters across in ice-free areas and those greater than 200 meters wide if frozen regularly.
Separation Distance for Unsuitable Habitat: 2 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Home ranges may be quite small, but at least some species exhibit good dispersal ability that may take them several kilometers from their natal area (Maier 2002). Peromyscus that have been displaced up to 3 km may return home within a few days (see Maier 2002). Displaced Neotoma fuscipes dispersed up to at least 1.6 km from their release point in five nights (Smith 1965). A male Dicrostonyx richardsoni moved more than 3 kilometers per day several times (Engstrom, in Wilson and Ruff 1999). Some species can traverse significant distances of unsuitable habitat. For example, Peromyscus leucopus may move between wooded areas separated by a deforested agricultural gap of up to at least 2 km (Krohne and Hoch 1999). In New Brunswick, a tagged subadult male Peromyscus maniculatus was captured at locations 1.77 km apart after a period of 2 weeks in September, suggesting that dispersal may extend at least this far (Bowman et al. 1999). In Kansas, individual Peromyscus maniculatus were captured at trap sites up to 1.32 km apart (Rehmeier et al. 2004). Dispersal can play a key role in the population dynamics of murid rodents.

Patterns of genetic (DNA) variation indicate that gene flow can be low among subpopulations of Neotoma magister and that effective dispersal is limited among subpopulations separated by as little as 3 km (Castleberry et al. 2002).

Separation distance for suitable habitat is a compromise between the typical small home range sizes of these mammals and their sometimes considerable dispersal ability and the likely low probability that two occupied locations separated by less than several kilometers of suitable habitat would represent independent populations.

Roads, especially divided highways, are major barriers to dispersal in small mammals (Oxley et al. 1974, Wilkins 1982, Garland and Bradley 1984).

Date: 08Mar2005
Author: Hammerson, G., and S. Cannings
Notes: Group contains most members of the family Muridae: mice, voles, lemmings, woodrats, etc.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 17Apr2006
NatureServe Conservation Status Factors Author: Hammerson, G., and J. C. Whittaker
Management Information Edition Author: Sam J. Norris
Management Information Acknowledgments: Information shared by the several Natural Heritage Programs, and the tireless efforts of Barbara Sargent of WVNHP, have made possible the completion of this ESA. Craig Stihler of WVDNR opened his files to me and made valuable suggestions.
Element Ecology & Life History Edition Date: 14Apr2006
Element Ecology & Life History Author(s): Hammerson, G.

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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