Neotoma magister

2006·0.31 MB·

by  Steven B. Castleberry

M S
AMMALIAN PECIES
No. 789, pp. 1–5, 3 figs.
Neotoma magister.
BySteven B. Castleberry, Michael T. Mengak, and W. Mark Ford
Published 24 May 2006 by the American Society of Mammalogists
Neotoma magister Baird, 1857 6.8 (0.3, 117); length of incisive foramen, 11.2 (0.6, 141), 11.1
(0.7, 118); length of maxillary toothrow, 9.6 (0.3, 144), 9.5 (0.4,
Allegheny Woodrat
121); length of nasals, 21.5 (1.1, 131), 21.1 (1.2, 97); length of
palatal bridge, 9.6 (0.7, 142), 9.5 (0.7, 117); length of rostrum,
Neotoma magister Baird, 1857:xliii, 498. Type locality ‘‘Carlisle
22.3(1.1,127),21.9(1.1,95);zygomaticbreadth,52.1(2.1,120),
Bone Cave,Pennsylvania.’’
51.4 (2.0, 107).
Neotoma pennsylvanica Stone, 1893:16. Type locality ‘‘near the
topofStoneMountain(2,000ft.),CumberlandCounty,Penna. DISTRIBUTION. The historic distribution of N. magister
[Pennsylvania], some six miles from Pine Grove, at a point generally followed the Appalachian Mountains and Interior High-
known asLewis’sCave.’’ landregionsoftheeasternUnitedStatesfromwesternConnecticut
Neotoma floridana magister: Schwartz and Odum, 1957:204. to northern Alabama (Goodwin 1932; Poole 1940). The speciesis
Namecombination. extirpated from Connecticut and New York and is restricted to a
small area along the Hudson River Palisades in New Jersey (Fig.
CONTEXT AND CONTENT. Order Rodentia, suborder
3). The southern and western borders of the distribution approxi-
Myomorpha, superfamily Muroidea, family Cricetidae, subfamily
mate the Tennessee River corridor, with the exception of 3 speci-
Neotominae, tribe Neotomini,genusNeotoma,subgenusNeotoma
menscollectedsouthoftherivernearMuscleShoals,ColbertCoun-
(MusserandCarleton2005);partoftheN.floridanaspecies-group
ty,Alabama,thatwereidentifiedasN.magisterbasedonpresence
(Birney 1973a; Burt and Barkalow 1942; Edwards and Bradley
of the maxillovomerine notch (Hayes and Richmond 1993). Al-
2001; Planzetal.1996).N. magisterismonotypic.
though cavedepositsindicateaformermorenortherlydistribution
DIAGNOSIS.ExternalmorphologyofN.magister(Fig.1)is (Richards 1987), the modern distribution in Indiana and Ohio is
similar to that of N. floridana, the only parapatric Neotoma. Al- restrictedtothelimestoneescarpmentsneartheOhioRiver(John-
though N. magister generally is larger in mass and with longer son 2002). To the east, the distribution follows the Blue Ridge
vibrissae,identificationbasedonsinglemeasurementsisunreliable Escarpment in Virginia, south to the northern portion of western
because of morphometric overlap (Ray 2000). Multivariate com- North Carolina. In the BlueRidgeof North Carolina,N. magister
parisonsofcranialcharacters(Fig.2)separateN.magisterandN. occurs at elevations .640 m (Ray 2000). Historical records exist
floridana primarily on size (Hayes and Richmond 1993; Ray from the Piedmont of Maryland and Virginia along the Potomac
2000).North–southclinalvariationdoesnotaccountforthediffer- River corridor near Great Falls, upstreamfrompresent-dayWash-
ences(HayesandRichmond1993).Themostreliablecharacterfor ington,D.C.(Wetmore1923).Thewesternextentofthedistribution
identifyingN.magisteristhepresenceofamaxillovomerinenotch alongtheOhioHillsinnorthwesternWestVirginiaandsouthwest-
(presentinall418N.magisterskullsbutinonly0.2%of457N. ern Pennsylvaniaisuncertain.
floridana skulls—Hayes and Richmond 1993). Additionally,only
FOSSIL RECORD. Fossil remains of N. magister are nu-
1.7% of N. magister skulls exhibited bifurcation of the anterior
merous from several Pleistocene cave deposits in Indiana, Ken-
palatalspine,whereas70.9%ofN.floridanaskullsexhibitedsome
tucky,Maryland,Ohio,Pennsylvania,Tennessee,andVirginia(Gid-
degreeofbifurcation.
leyandGazin1933,1938;Guildayetal.1964,1977,1978;Rich-
GENERALCHARACTERS.Dorsalpelageisgraytobrown- ards1972,1987).TheoldestremainsarefrommiddlePleistocene
ish gray with more brown typically present in adults. Ventralsur- depositsatCumberlandCave,Maryland,andTroutCave,WestVir-
face is white from throat to tip of tail. Tail is long, moderately ginia (Kurte´n and Anderson 1980). The unglaciated southernAp-
haired, and distinctly bicolored. A midventral gland is present in palachians served as a late Pleistocene refugium for N. magister
adultsofbothsexes.Meanexternalmeasurements(inmm;SDand (Birney1973b;Guildayetal.1964)whenhabitatconditionsinthe
n in parentheses) based on adult museum specimens from Ala- northern extent of the current distribution wereunsuitable(Hayes
bama,Kentucky,Maryland,NewJersey,NewYork,NorthCarolina, and Harrison1992).
Pennsylvania,Tennessee,Virginia,Washington,D.C.,andWestVir-
FORM AND FUNCTION. Neotoma magister has long vi-
ginia (Hayes and Richmond 1993) for males and females,respec- brissae, usually .51 mm (Ray 2000). Vibrissae number up to 50
tively, are: length ofhead and body (definedastotallengthminus
length of tail vertebrae), 227.8 (14.3, 132), 224.1 (12.0, 110);
length ofhind foot, 42.7 (2.0, 133), 42.2 (2.0, 112); length ofear,
30.6(2.3,72),29.8(2.8,59).Meanexternalmeasurements(inmm;
range and n in parentheses) based on adult museum specimens
from Alabama, Kentucky, Maryland, North Carolina, Tennessee,
Virginia,Washington,D.C.,andWestVirginia(Ray2000)are:total
length, 397.4 (311–451, 91); length of tail, 180 (147–210, 91);
length of hind foot, 41.7 (35–46, 92); length of ear, 28.3 (23–34,
42). Mean bodymasses(in g;SDand n in parentheses)formales
and females, respectively, are 357.0 (56.5, 48) and 337.0 (53.5,
36) fromacrossthedistribution(Hayesand Richmond1993).
Meancranialmeasurements(inmm;SDandninparentheses)
from Alabama, Kentucky, Maryland,NewJersey,NewYork,North
Carolina,Pennsylvania,Tennessee,Virginia,Washington,D.C.,and
WestVirginia(HayesandRichmond1993)formalesandfemales,
respectively, are: breadth at mastoids, 20.0 (0.8, 129), 19.8 (0.7,
110);breadthofmetapterygoidfossa,4.3(0.3,142),4.3(0.3,117);
breadth of rostrum, 8.4 (0.4, 144), 8.4 (0.4, 121); breadth of zy-
gomaticplate,4.7(0.3,145),4.7(0.3,121);condylobasilarlength,
52.1(2.1,120),51.4(2.0,107);greatestlengthofskull,54.0(2.0, FIG. 1. AdultNeotoma magisterinRandolphCounty,West
116), 53.3 (1.8, 83); least interorbital constriction, 6.8 (0.2, 141), Virginia.Photographby StevenB. Castleberry.
2 MAMMALIANSPECIES 789—Neotoma magister
FIG. 3. Distribution of Neotoma magister in the eastern
UnitedStates.
uals. Ovaries are covered by fat and folds of the oviducts. Uterus
isbicornusand convergesintoa broadU-shape.
Water consumption by captive N. magister ranged from 0 to
250mloffreewaterper48h(Newcombe1930;Poole1940).Free-
ranging N. magister generally have year-round access to water in
FIG.2. Dorsal,ventral,andlateralviewsofcraniumandlat- streamsandseepagewaterinornearcavesandrockoutcrops.
eralviewofmandibleofNeotomamagisterfromBuncombeCoun-
ty, North Carolina (female, Georgia Museum of Natural History ONTOGENYANDREPRODUCTION.Thebreedingsea-
25729). Greatestlengthofcraniumis48 mm. sonisvariabledependingonlocation.Specimensfromsoutheastern
Pennsylvaniaproduced 2 or3 littersper yearbetweenmid-March
and October (Poole 1940). In western Virginia, breeding occurs
oneachsideofthemuzzle(Howell1926).Blackvibrissaearestiff, year-round, but most young are born between May and October
whereas white vibrissae are softer. A few vibrissae are blacknear (Mengak2002a).Thegestationperiodis30–36days(Poole1940).
the base becoming white on the distal half. Molt in N. magister Typical litter sizes range from 1 to 4, with means of 2.0 (Poole
progressesuniformlyovertheentirebody(Poole1940).Dentalfor- 1940)and2.3(Mengak2002a).Sexualmaturityisattainedat3–4
mulaisi 1/1,c0/0, p 0/0,m 3/3,total16. months (Poole 1940). Individuals may give birth at 10 months
Themidventralglandbecomesactiveduringthebreedingsea- (Poole1940).
son, especially in males (Poole 1940). The odor of the midventral Youngarepinkandnakedatbirthandweigh15–17g(Men-
glandis‘‘marked’’(Poole1940:263). gak 2002a; Poole 1936, 1940). By day 5, young are covered by
The baculum of N. magister is short with a broad proximal finehairandarefullyfurredat2weeks.Eyelidsareconspicuously
end and lateral wings giving a U-shaped appearance (Burt and black until eyes open. Eyes become sensitive to light at 2 weeks
Barkalow1942).Meanbacularmeasurements(inmm;rangeinpa- and are fully open at 3 weeks (Poole 1936). Tail remains hairless
rentheses) of 4 specimens of N. magister examined by Burt and until ca. 3 weeks. Mean growth rates of juveniles (body mass #
Barkalow (1942) are: length, 6.79 (6.41–7.10); diameter at base: 175 g) and subadults (176–225 g) in western Virginia were 1.26
dorsoventral, 1.74 (1.45–1.98) and lateral, 3.32 (2.93–3.50); di- and 0.95 g/day, respectively (Mengak 2002a). Mother and young
ameter of shaft: dorsoventral, 0.69 (0.60–0.86) and base, 0.82 remain together until individual young weigh $115 g (Mengak
(0.75–0.93). Penis has recurved spines thatexpand in the vagina, 2002a).N.magistercanliveto48monthsincaptivity(Poole1940)
allowingacopulatorylockduringmating(Howell1926).Testesare and to58 monthsinthewild(Mengaketal.2002).
partly abdominal,especiallywhenreproductivelyactive(Patterson Young are reared in nests constructedofbark,grasses,roots,
1933). Mean testes measurements for 3 males from West Virginia and shredded wood fibers(Poole1940). The outsideofthenestis
inFebruarywere18.5mmpolarlengthby11mmdiameter.Scro- constructed of coarse materials, and the inside is lined with finer
tum is shallow, enclosing one-third of testicles and epididymides materials (Newcombe 1930; Poole 1940). Nests have an outside
(Patterson1933).Aprostateispresent.Cowper’sglandsarecaudal diameterof460mmandanestcavitydiameterof120mm(Poole
to ischiocavernous muscle. Spongy tissue and coagulating glands 1940). Neststypicallyarelocated in inaccessiblerockcrevicesor
surround theneck of thebladder in reproductivelyactiveindivid- on ledgesincaves.
789—Neotoma magister MAMMALIANSPECIES 3
ECOLOGY. The presence of rock habitats, such as boulder grayfoxes(Urocyoncinereoargenteus),easternspottedskunks,and
fields, caves, cliff faces, or talus slopes, is the limiting factor for long-tailed weasels (Balcom and Yahner 1996; Poole 1940). Rac-
occurrenceregardlessofvegetationtype(Heisler1941;Newcombe coons(Procyonlotor)hosttheascaridnematodeBaylisascarispro-
1930; Poole 1940). N. magister occurs in a wide variety offorest cyonis,whichcausesfatalneurologicaldiseaseinintermediateand
types,includingnorthernhardwood(composedofsugarmaple[Acer aberranthosts.B.procyonishasbeenimplicatedinpopulationde-
saccharum], yellow birch [Betula alleghaniensis], and American clinesofN.magisterinthenorthernpartsoftherange(LoGiudice
beech[Fagusgrandifolia])andredspruce(Picearubens)–eastern 2001, 2003; Owen et al. 2004). N. magister is at greater risk of
hemlock (Tsuga canadensis)athigherelevations(.800m)inthe infection than other small mammals because of its propensity to
AlleghenyMountainsinMaryland,Pennsylvania,andWestVirgin- collectandcachefoodandnonfooditems,includingfecesthatmay
ia.Thespeciesoccursinmixed-mesophyticandmixed-oak(Quer- contain B. procyonis eggs. N. magister regularly collected simu-
cus)–pine (Pinus) forest types throughout its distribution (Castle- latedraccoonfecesatraccoonlatrinesites(LoGiudice2001).
berry et al. 2002a, 2002c; Fassler 1974; M. T. Mengak, in litt.). A diverse assemblage of ectoparasites, including chiggers,
SandstonerockoutcropsoccupiedbyN.magisterinnorth-central fleas, mites, and ticks, occurs with N. magister in Indiana (Cud-
West Virginia occur on steeper slopes, are wider, have less leaf more 1986). Two flea (Epitedia cavernicolaandOrchopeaspenn-
litter accumulation, and have fewer understory trees than do un- sylvanicus) and 1 tick (Ixodes angustus) species are known from
occupied rock outcrops (Myers 1997). In eastern Kentucky, N. N. magister in West Virginia (Castleberry et al. 2003). The Neo-
magister occupies rock outcrops on steep slopes with high over- toma-specific flea, O. pennsylvanicus, is common throughout the
story tree densities(Bommarito 1999).N. magistertoleratesava- distribution(Benton1971;Castleberryetal.2003;Cudmore1986).
riety of forest stand age and structure conditions but selects for- TheprotozoanparasiteEimerianeotomawasdocumentedinasin-
aging areas with diverse understory vegetation (Castleberry et al. glefemaleincentralPennsylvania(Sands1951).
2002c). Althoughpopulationsinthenorthernandwesternperipheries
Mean home-range sizes of 37 individuals radiotracked from of the distribution have experienced dramatic declines in recent
MaytoAugustineast-centralWestVirginiawere6.5haformales years (Balcom and Yahner 1996), N. magister is still believed to
(n519)and2.2haforfemales(n518—Castleberryetal.2001). becommoninappropriatehabitatsinthecentralandsouthernparts
Mean home-range sizes of 7 individuals on the same study area oftherange.N.magisterisrelativelyabundantinportionsofwest-
fromOctobertoDecemberwere0.49and0.78ha,respectively,for ern North Carolina (Ray 2000) and is commonly associated with
males (n 5 3) and females (n 5 4—Hornsby et al. 2005). Mean caveentrancesinthelimestoneregionsofcentralTennessee(Ken-
home-rangesizewas0.18hafor4femalesradiotrackedfromJan- nedyandHarvey1980).InKentucky,thespeciesismoreabundant
uary to April in western Virginia (Mengak 2002b). N. magister insandstonecliffsthroughouttheCumberlandPlateauthaninthe
restrictsitsmovementawayfromrockhabitatsinlatefallandwin- HighlandRimandlargelyabsentintheBluegrassregions(Barbour
ter when little green vegetation is available and cached foods are and Davis 1974; Bommarito 1999; Fassler 1974). Populationesti-
consumed(Hornsbyetal.2005).
mates for 2 sites in the Blue Ridge of western Virginia studied
In eastern Pennsylvania, N. magister consumes variousveg-
intensively over an 11-year period ranged from 0 to 24.1 individ-
etative and fruiting parts of woody plant species, including black
uals, with long-term averages of 6.3 and 10.6 (M. T. Mengak, in
birch(Betulalenta),Americanchestnut(Castaneadentata),flow-
litt.).
ering dogwood (Cornus florida), apple (Malus pumila), bear oak
(Quercusilicifolia),blackcherry(Prunusserotina),rhododendron
BEHAVIOR.Neotomamagisterisagonistictowardconspe-
(Rhododendron maximum), mountain ash (Sorbus americana),
cifics (Poole 1940). Adult home ranges overlap extensively, but
andeasternhemlock(Poole1940).Frondsofthecommonpolypody
eachindividualdefendsauniquedensite(Castleberryetal.2001;
fern (Polypodium vulgare) were common in stomach contents of
Kinsey1977).Incaptivity,adultfemalessecurethebestdensites
N.magisterincentralPennsylvania(Heisler1941).Themostcom-
for breeding and rearing young, adult males and juveniles reside
monfooditemsconsumed(fromhighesttolowestbypercentagein
incommunalaggregations,andsubordinateindividualsavoidcon-
the diet) in xeric oak–pine forests in the Ridge and ValleyofVir-
frontationbyavoidingdominantfemales(Kinsey1977).Whencon-
ginia and eastern West Virginia were blackberry (Rubus) leaves,
fined at high densities, N. magisterformsdominancehierarchies,
fungi, greenbrier (Smilax) leaves, acorns, and oak leaves (Castle-
displays high levels of aggression, and lacks communal aggrega-
berry et al. 2002a). In mesic, mixed oak–northern hardwood and
tions (Kinsey 1977). Males become more aggressive in fall when
red spruce–hemlock forests in the Allegheny Plateau of WestVir-
theycompeteintensivelyfornestsites.
ginia,themostcommonfoodswere(fromhighesttolowestbyper-
Inhigh-densitycaptivepopulations,bodymassandoccupancy
centage in the diet)fungi,acorns,holly(Ilex)andblueberry(Vac-
of nest boxes are positively correlated with social rank (Kinsey
cinium)fruit,fern(Dryopteris),andlichen.Acornsareanimportant
1976).Timespentsittingoutsidenestboxesandfrequencyofdis-
component of the diet, and consumption typically reflects annual
tress vocalizations are correlated negatively with social rank. In-
and seasonalavailability(Castleberryetal.2002a).
termediate-rank individuals have more sexual encounters than do
Neotomamagisteraccumulateslargepilesofsticksandother
alpha individuals. In similar-sized captive groups of N. fuscipes
debris,referredtoasmiddens,underneathoverhangingrockledg-
macrotusandN.magister,thelatterspeciesexhibitshigherlevels
es, which often are at crevice openings (Heisler 1941;Newcombe
of aggression and does not form communal aggregations (Kinsey
1930;Poole1940).Althoughtheirspecificfunctionsareunknown,
1976).
middensarecommondepositionsitesforfooditemssuchasgreen
Adult N. magister generally are not vocal but may vocalize
vegetation and fungi, as well as nonfood items. N. magister de-
when fighting or injured (Poole1940).Incaptivity,variousvocali-
posits feces and urine at latrine sites located on clean, flat rock
zations,include‘‘squeaking’’and‘‘whimpering’’(MengakandZad-
surfaces(Poole1940).Bothsexesdepositscentfromthemidventral
nik2005).Youngcommonlysquealwhileinthenest(Poole1940).
glandnearlatrineareas(S.B. Castleberry,inlitt.).
Malesmakea ‘‘low-pitchedraspysound’’whenfollowingafemale
Neotoma magistercommonlyoccursin habitatsoccupiedby
thenorthernshort-tailedshrew(Blarinabrevicauda),southernred- before mating (Kinsey 1976; Mengak and Zadnik 2005).N. mag-
backedvole(Myodesgapperi),rockvole(Microtuschrotorrhinus), ister produces nonvocal sounds used to orient in the dark when
white-footed deermouse (Peromyscus leucopus), North American ambientsoundsareminimal(DunningandPayne1979).
deermouse(P.maniculatus),cinereusshrew(Sorexcinereus),rock Themidventralglandisusedtoscent-markobjects,leavinga
shrew(S.dispar),smokyshrew(S.fumeus),andredsquirrel(Tam- yellowish brown stain on the adjacent fur (Kinsey 1976). Males
iasciurus hudsonicus—Castleberry et al. 2003; Heisler 1941; scent-mark by pressing the body against objects, using the front
Sands 1951). Rafinesque’s big-eared bat (Corynorhinus rafines- feettodragthebodyalongtodepositscent.Whensexuallyactive,
quii), Townsend’s big-eared bat (C. townsendii), common raven the midventral gland becomes discolored by secretion and soil
(Corvuscorax),long-tailedweasel(Mustelafrenata),easternsmall- picked up by rubbing the gland over rocks and substrate (Poole
footedmyotis(Myotisleibii),easternspottedskunk(Spilogalepu- 1940).Useofscentfororientationincavesandcrevicesislimited
torius),andAmericanblackbear(Ursusamericanus)oftenusethe because cave floors are not a continuous substrate (Dunning and
samerockhabitatsasN.magisterfordenning,roosting,ornesting. Payne1979).
PotentialpredatorsofN.magisterincludegreathornedowls(Bubo Neotomamagisterisstronglynocturnal(Zervanos1969).Red
virginianus), timber rattlesnakes (Crotalus adamanteus), other light is perceived as white light, reducing activity (Zervanos and
snakes, bobcats (Lynx rufus), striped skunks (Mephitis mephitis), Davis 1968). Daily activity begins ca. one-half hour after sunset
4 MAMMALIANSPECIES 789—Neotoma magister
andcontinuesforseveralhoursbeforetaperingoff,butpeaksagain BARBOUR,R.W.,ANDW.H.DAVIS. 1974. MammalsofKentucky.
neardawn (Poole1940; Zervanos1969). UniversityPressofKentucky,Lexington.
BENTON,A.H. 1971. Anannotatedlistofthefleas(Siphonaptera)
CONSERVATIONSTATUS.Throughoutitsrange,N.mag-
of West Virginia. Proceedings of the West Virginia Academy
ister has declined coincident with the decline of the American
ofScience40:35–39.
chestnutand,morerecently,withadeclineinoakabundance(Cas-
BIRNEY,E.C. 1973a. Anassessmentofrelationshipsandeffects
tleberryetal.2002a;MillerandKochenderfer1998;Schuler2004;
of interbreeding among woodrats of the Neotoma floridana
Wright and Kirkland 2000). In Pennsylvania, sites formerlyoccu-
species-group.JournalofMammalogy57:103–132.
pied by N. magister have forests with fewer oak and more conif-
BIRNEY, E. C. 1973b. Systematics of three species of woodrats
erous speciesthan currentlyoccupied locations(BalcomandYah-
(genus Neotoma) in central North America. Miscellaneous
ner 1996). Although N. magister is present in areas with human
Publications,MuseumofNaturalHistory,UniversityofKansas
activity,forestconversionandfragmentationisgreateratextirpated
58:1–173.
sitesthanatcurrentlyoccupiedsites.Clear-cuttingand2-agedre-
BOMMARITO, M. P. 1999. Distribution of the Allegheny woodrat
generation harvesting (removing all trees except 6–10 residual
in the Daniel Boone National Forest: habitat characteristics
trees/ha)in the centralAppalachian hardwood regionofWestVir-
and incidence of raccoon roundworm. M.S. thesis, Eastern
giniahaveminimaldirectimpactonN.magisteriftheforestover-
KentuckyUniversity,Richmond,62 pp.
storyisretainedimmediatelysurroundingoutcropsandintactforest
BURT, W. H., AND F. S. BARKALOW. 1942. A comparative study
ismaintainedon1adjacentside(Castleberryetal.2001).Limited
ofthebaculaofwoodrats(subfamilyNeotominae).Journalof
clear-cutting and 2-aged harvesting near outcrops can benefit N.
Mammalogy23:287–297.
magisterbyprovidingabundantsoftmastandsucculentvegetation
CASTLEBERRY,N.L.,S.B.CASTLEBERRY,W.M.FORD,P.B.WOOD,
thatisconsumedinspringandsummer(Castleberryetal.2002a).
AND M. T. MENGAK. 2002a. Allegheny woodrat (Neotoma
ExtirpationofN.magisterintheNortheastwhereraccoondensities
magister) food habits in the central Appalachians.American
arehighmaybemediatedbytheparasiteB.procyonis(LoGiudice
MidlandNaturalist147:80–92.
2001, 2003), but no evidence supports this relationship in other
CASTLEBERRY,S.B.,N.L.CASTLEBERRY,P.B.WOOD,W.M.FORD,
areas(Owenetal.2004).
ANDM.T.MENGAK. 2003. Fleas(Siphonaptera)ofAllegheny
GENETICS. Neotoma magister has 2n 5 52 chromosomes woodratsinWestVirginiawithcommentsonectoparasitehost
(RayandWebster2004).Twoautosomepairsarelargesubtelocen- specificity.AmericanMidlandNaturalist149:209–212.
tric,2pairsaresmallsubtelocentric,and21pairsarelargetosmall CASTLEBERRY,S.B.,W.M.FORD,P.B.WOOD,N.L.CASTLEBERRY,
acrocentrics.TheXchromosomeislargesubtelocentricandtheY ANDM.T.MENGAK. 2001. MovementsofAlleghenywoodrats
chromosomeismediumsubtelocentric. in relation to timber harvesting. Journal of Wildlife Manage-
PatternsofvariationatN.magistermicrosatellitelociindicate ment65:148–156.
significant genetic differentiation among geographically distinct CASTLEBERRY, S. B., T. L. KING, P. B. WOOD, AND W. M. FORD.
populations throughout the distribution with isolation-by-distance 2000. MicrosatelliteDNAmarkersforthestudyofAllegheny
asthelikelyisolatingmechanism(Castleberryetal.2000,2002b). woodrat (Neotoma magister) populations and cross-species
Although genetically distinct, relatively frequent gene flow occurs amplification in the genus Neotoma. Molecular Ecology 9:
among N. magister subpopulations at geographically proximate 824–826.
rockhabitats.TheremnantpopulationinNewJerseyexhibitsgreat- CASTLEBERRY, S. B., T. L. KING, P. B. WOOD, AND W. M. FORD.
lyreducedgeneticdiversitycomparedtootherpopulationsthrough- 2002b. Microsatellite DNA analysis of population structure
outthedistribution(Castleberryetal.2002b). in Allegheny woodrats (Neotoma magister). JournalofMam-
ApossiblezoneofhybridizationbetweenN.magisterandN. malogy83:1058–1070.
floridanahaematoreiaexistsinBurkeCounty,NorthCarolina(Ray CASTLEBERRY,S.B.,P.B.WOOD,W.M.FORD,N.L.CASTLEBERRY,
2000). Although analysis of mitochondrial DNA (mtDNA) D-loop AND M. T. MENGAK. 2002c. Summer microhabitatselection
sequencesidentified3of5specimensfromtheareaasN.magister byforagingAlleghenywoodrats(Neotomamagister)inaman-
and 2 as N. f. haematoreia, all were identified as N. magister agedforest.AmericanMidlandNaturalist147:93–101.
basedoncranialmeasurementsandpresenceofamaxillovomerine CUDMORE, W. W. 1986. Nest associates and ectoparasites of the
notch. eastern wood rat, Neotoma floridana, in Indiana. Canadian
VariationinmtDNAcytochrome-bsequencessuggeststhatN. JournalofZoology64:353–357.
magisterisincludedinadistinctcladewithN.albigula,N.flor- DUNNING,D.C.,ANDL.N.PAYNE. 1979. Orientationincave-dwell-
idana,andN.goldmani(EdwardsandBradley2002).N.magister ingwoodrats.BehavioralEcologyandSociobiology6:1–9.
isconsideredasisterspeciestoN.floridana(EdwardsandBradley EDWARDS, C. W., AND R. D. BRADLEY. 2001. Molecular phylo-
2001). genetics of the Neotoma floridana species group. Journal of
Mammalogy82:791–798.
REMARKS. Neotoma is the combined form of the Greek EDWARDS, C. W., AND R. D. BRADLEY. 2002. Molecularsystem-
wordsneos,neworyoung,andtomos,cutting(Stangletal.1993). aticsofthegenusNeotoma.MolecularPhylogeneticsandEvo-
The specific name refers to the Latin word for master or teacher. lution25:489–500.
SchwartzandOdum(1957)suggestedthatN.magisterbeincluded FASSLER, D. J. 1974. Mammals of Pulaski County, Kentucky.
asasubspeciesofN.floridanabasedoncranialandexternalmor- TransactionsoftheKentuckyAcademyofScience35:37–43.
phology. Analyses of external and cranial morphology (Hayes and GIDLEY, J. W., AND C. L. GAZIN. 1933. New Mammalia in the
Richmond 1993), mtDNA restriction sites (Hayes and Harrison Pleistocene fauna from Cumberland Cave. Journal of Mam-
1992;Planzetal.1996),andmtDNAcytochrome-bsequences(Ed- malogy14:343–357.
wardsandBradley2001,2002)supporttherecognitionofN.mag- GIDLEY,J.W.,ANDC.L.GAZIN. 1938. ThePleistocenevertebrate
ister as a distinct species. Baker et al. (2003) and Jones et al. fauna from Cumberland Cave, Maryland. U.S. National Mu-
(1997)usethecommonnameAppalachianwoodrat. seumBulletin171:1–99.
WethankM.E. McGheeforaccesstomuseumspecimens. GOODWIN, G. G. 1932. New records and some observations on
Connecticutmammals.JournalofMammalogy13:36–40.
LITERATURECITED
GUILDAY, J. E., H. W. HAMILTON, E. ANDERSON, AND P. W. PAR-
BAIRD,S.F. 1857. Mammals.Reportsofexplorationsandsurveys, MALEE. 1978. TheBakerBluffCavedeposit,Tennesseeand
to ascertain the most practicable and economical route for a thelatePleistocenefaunalgradient.BulletinofCarnegieMu-
railroad from the Mississippi River to the Pacific Ocean. seumofNaturalHistory11:1–67.
House of Representatives, A. O. P. Nicholson, Washington, GUILDAY, J. E., P. S. MARTIN, AND A. D. MCCRADY. 1964. New
D.C. Paris No. 4: a Pleistocene cave deposit in Bedford County,
BAKER,R.J.,ETAL. 2003. RevisedchecklistofNorthAmerican Pennyslvania.NationalSpeleologicalSocietyBulletin26:121–
mammalsnorthofMexico,2003.OccasionalPapers,TheMu- 194.
seum,TexasTechUniversity229:1–24. GUILDAY, J. E., P. W. PARMALEE, AND H. W. HAMILTON. 1977.
BALCOM,B.J.,ANDR.H.YAHNER. 1996. Microhabitatandland- TheClark’sCavebonedepositandthelatePleistocenepaleo-
scapecharacteristicsassociatedwiththethreatenedAllegheny ecologyofthecentralAppalachianMountainsofVirginia.Bul-
woodrat.ConservationBiology10:515–523. letinofCarnegieMuseumofNaturalHistory2:1–87.
789—Neotoma magister MAMMALIANSPECIES 5
HAYES, J. P., AND R. G. HARRISON. 1992. Variationinthemito- woodrat in northcentral West Virginia. M.S. thesis, WestVir-
chondrialDNAandthebiogeographichistoryofthewoodrats giniaUniversity,Morgantown,85 pp.
(Neotoma) of the eastern United States. Systematic Biology NEWCOMBE, C. L. 1930. An ecological study of the Allegheny
41:331–344. cliffrat(NeotomapennsylvanicaStone).JournalofMammal-
HAYES, J. P., AND M. E. RICHMOND. 1993. Clinal variation and ogy11:204–211.
morphology of woodrats (Neotoma) of the eastern United OWEN, S. F., J. W. EDWARDS,W.M.FORD,J.M.CRUM,ANDP.B.
States.JournalofMammalogy74:204–216. WOOD. 2004. Raccoon roundworm in raccoons in central
HEISLER,W.T. 1941. Alleghenywoodratpopulations.M.S.thesis, WestVirginia.NortheasternNaturalist11:137–142.
PennsylvaniaStateCollege,StateCollege,77 pp. PATTERSON, R. C. 1933. Notes on Neotoma pennsylvanicawith
special reference to the genital organization. Proceedings of
HORNSBY, B. S., A. M. RUIZ, S. B. CASTLEBERRY, N. L. CASTLE-
theWestVirginiaAcademyofScience15:38–42.
oBfERARllYe,gWhe.nMy.wFoOoRdDra,tAsNiDnPh.aBrv.eWstOedODa.nd20in0t5a.ctFsatlalnmdsovienmWenetsst PLANZ,J.V.,E.G.ZIMMERMAN,T.A.SPRADLING,ANDD.R.AKINS.
1996. MolecularphylogenyoftheNeotomafloridanaspecies
Virginia.NorthernJournalofAppliedForestry22:281–284.
group.JournalofMammalogy77:519–535.
HOWELL,A.B. 1926. Anatomyofthewoodrat.Monographsofthe POOLE,E.L. 1936. NotesontheyoungoftheAlleghenywoodrat.
American Society of Mammalogists 1. Williams & Wilkins,
JournalofMammalogy17:22–26.
Baltimore,Maryland. POOLE,E.L. 1940. AlifehistorysketchoftheAlleghenywoodrat.
JOHNSON, S. S. 2002. Reassessment of the Allegheny woodrat JournalofMammalogy21:249–270.
(Neotoma magister) in Indiana. Proceedings of the Indiana RAY,D.K. 2000. Phylogeneticsandevolutionofwoodrats(genus
AcademyofScience111:56–66. Neotoma) in the southern Appalachian mountains. M.S. the-
JONES,C.,etal. 1997. RevisedchecklistofNorthAmericanmam- sis,UniversityofNorthCarolinaatWilmington,68 pp.
malsnorthofMexico,1997.OccasionalPapers,TheMuseum, RAY, D. K., AND W. D. WEBSTER. 2004. Karyotypic analysis of
TexasTechUniversity173:1–20. Neotoma(Rodentia:Muridae)fromtheeasternUnitedStates.
KENNEDY,M.L.,ANDM.J.HARVEY. 1980. Mammals.Pp.1–50 JournaloftheNorthCarolinaAcademyofScience120:50–54.
inTennesseerarevertebrates(D.C.EagerandR.M.Hatcher, RICHARDS, R. L. 1972. The woodrat in Indiana: recent fossils.
eds.). Tennessee Wildlife Resources Agency and Tennessee ProceedingsoftheIndianaAcademyofScience81:370–375.
DepartmentofConservation,Nashville. RICHARDS,R.L. 1987. TheQuaternarydistributionoftheeastern
KINSEY, K. P. 1976. Social behaviour in confined populationsof woodrat, Neotoma floridana, in Indiana. Proceedings of the
theAlleghenywoodrat,Neotomafloridanamagister.Animal IndianaAcademyofScience96:513–521.
Behaviour24:181–187. SANDS, D. E. 1951. A study of theAllegheny woodratincentral
KINSEY, K. P. 1977. Agonistic behavior and social organization Pennsylvania. M.S. thesis, Pennsylvania State College, State
College,73 pp.
inareproductivepopulationofAlleghenywoodrats,Neotoma
floridana magister.JournalofMammalogy58:417–419. SCHULER,T.M. 2004. Fiftyyearsofpartialharvestinginamixed
mesophytic forest: composition and productivity. Canadian
KURTE´N, B., AND E. ANDERSON. 1980. Pleistocene mammals of JournalofForestResearch34:985–997.
NorthAmerica.ColumbiaUniversityPress,NewYork.
SCHWARTZ,A.,ANDE.P.ODUM. 1957. Thewoodratsoftheeast-
LOGIUDICE, K. 2001. Latrine foraging strategies of two small ern UnitedStates.JournalofMammalogy38:197–206.
mammals: implications for the transmission of Baylisascaris
STANGL, F. B., JR., P. G. CHRISTIANSEN, AND E. J. GALBRAITH.
procyonis.AmericanMidlandNaturalist146:369–378. 1993. Abbreviated guide to pronunciation and etymology of
LOGIUDICE, K. 2003. Trophically transmitted parasites and the scientific names for North American land mammals north of
conservationofsmallpopulations:raccoonroundwormandthe Mexico.OccasionalPapers,TheMuseum,TexasTechUniver-
imperiled Allegheny woodrat. Conservation Biology 17:258– sity154:1–26.
266. STONE, W. 1893. Description of a new species ofNeotomafrom
MENGAK,M.T. 2002a. Reproduction,juvenilegrowthandrecap- Pennsylvania.ProceedingsoftheAcademyofNaturalScienc-
ture rates of Allegheny woodrats (Neotoma magister)in Vir- esofPhiladelphia45:16–18.
ginia.AmericanMidlandNaturalist148:155–162. WETMORE,A. 1923. ThewoodratinMaryland.JournalofMam-
MENGAK, M. T. 2002b. Home range and movement of the Alle- malogy4:187–188.
ghenywoodrat(Neotomamagister)inVirginia.Banisteria19: WRIGHT, J., AND G. L. KIRKLAND. 2000. A possible role forthe
3–8. chestnutblightinthedeclineoftheAlleghenywoodrat.Jour-
MENGAK, M. T., AND A. K. ZADNIK. 2005. Behavior of captive naloftheAmericanChestnutFoundation8:30–35.
Allegheny woodrats (Neotoma magister) in Virginia. Banis- ZERVANOS,S.M. 1969. Effectsofphotoperiodonthedailyactiv-
teria26:11–14. ityoftheAlleghenywoodrat(Neotomafloridana).Mammalia
MENGAK, M. T., S. B. CASTLEBERRY, W. M. FORD, J. RODRIGUE, 33:509–515.
ANDN. L. CASTLEBERRY. 2002. Longevityrecordforawild ZERVANOS,S.M.,ANDD.E.DAVIS. 1968. Perceptionofredlight
by woodrats(Neotoma floridana).JournalofMammalogy49:
Alleghenywoodrat(Neotomamagister)inWestVirginia.Vir-
759.
giniaJournalofScience53:167–170.
MILLER, G. W., AND J. N. KOCHENDERFER. 1998. Maintaining
speciesdiversityinthecentralAppalachians.JournalofFor- Associate editors of this account were RONALD GETTINGER, KRIS-
estry96:28–33. TOFERHELGEN,andPAMELAOWEN.EditorwasVIRGINIAHAYSSEN.
MUSSER,G.G.,ANDM.D.CARLETON. 2005. SuperfamilyMuroid- STEVENB.CASTLEBERRY,WARNELLSCHOOLOFFORESTRESOURC-
ea. Pp. 894–1531 in Mammal species of the world: a taxo- ES, UNIVERSITY OF GEORGIA, ATHENS, GEORGIA 30602. MICHAEL
nomic and geographical reference (D. E. Wilson and D. M. T.MENGAK,WARNELLSCHOOLOFFORESTRESOURCES,UNIVERSITY
Reeder,eds.).Thirdedition.JohnsHopkinsUniversityPress, OF GEORGIA, ATHENS, GEORGIA 30602. W. MARK FORD, UNITED
Baltimore,Maryland. STATES DEPARTMENT OF AGRICULTURE FOREST SERVICE, NORTH-
MYERS, R. T. 1997. Microhabitat and ecology of the Allegheny EASTERNRESEARCHSTATION,PARSONS,WESTVIRGINIA26287.