MAMMALIAN SPECIES no. 431, pp. 1-10, 4 6s Aplodontia rufa. Published 23 April 1993 by The American By Leslie N. Carraway and B. J. Verts ty of Mammalogists Aplodontia Richardson, 18296 Aplodontia Richardson, 18295:334, Type species Aplodontia le poring Richardson (= Anisonys rufa Ratineaqie). CONTEXT AND CONTENT. Order Rodentia, Suborder Sciuromorphe, Superfamily Aplodontoidea, Family Aplodontidae, Ge rvs Aplodontia (Hall, 1981; Miler and Kellogg, 1955; Simpson, 1945). One living specice i recognize! (Fall, 1981), Aplodontia rufa (Rafinesque) Mountain beaver Anisonys? raf Rafiresque, 1817:45, Type locality “Neighbour hood of the Columbus River” Aplodonta leporina Richardson, 18298:435. No locality, but nest alumbva River implied. Based on a specimen (ao. 17) inthe ‘Hoson's Bay Museum Aplodontiarafa: Merriam, 1886:316, First formal we of current ‘same combination Aplodntia major Merriam, 186316 Trp ay “Pace ca "y, Cal" Aplodontia major Merci, 1899:20. Type locality “Queril [sc] Lake, Oxympic Ms, [Grays Harbor Co], Washington, Aplodontia olsmpica Merriam, 189920, Type locality “uciult {sie} Lake, Olympic Mts. (Grays Harbour Co. Washington.” Aplodontia pacifiea Merriam, 1899:19. Type lealty “Newport, mouth Of Yaquina Bay, [Lincoln Co.) Oregon.” Aplodontia phaca Merraat, 1890:20, ype locality “Pt. Reyes, Marin Go, California.” Aplodontia chryseola Kellogg, 1914:295. Type locality “Jackson Lake, Siskiyou County, California, tude 5900 feet. Aplodantia nigra Taylor, 1914297. Type locity “Point Arena, Mendocino County, Calfraia.” Aplodontia humboldtiana Taylor, 1916a:21. Type lcaliy “Cae lotta, Humboldt County, Caloris.” CONTEXT AND CONTENT. Context same as fr gers Seven subspecies are recognized currently (Dalquest and Scheer, 1945; Hall, 1981) Ar californica (Peters, 18642179), Type locality “nese main top.” Type locality restricted to crest of Sierra Nevada Mountains, (Cater, by Hall 1941}. (major Merriam, 1886, s synonym) 1 humboldiiana Tayler, 19L6a:21, soe above, 1 nigra Taylor, 1914:297, see above 1. pacifica Merriam, 189919, see above 1. phaca Merriam, 1899:20, see above 1 ‘rainieri Merriam, 1899:21. Type locality “Paradise Creek, southside MM. Rainier, (Pierce Co.) Washington (a, 5200) ft,” (columbiana Tayler, 19166, a synonym). rufa (Rafinesque, 1817:48), see above, (leporina Richardson, 18298, olympica Mercian, 1899, chryseola Kellogg, 1914, and grisea Taylor, 19166, are synonyms) DIAGNOSIS. The mountsin beaver (Fig. 1) hes the general appearance of a medium-siod muskeat (Ondaira zibethicus) except ‘he tail i wellfured and exceedingly short, Other characteristics of the specios inci a pelagetniformily dark brown except for = ‘rhe spot below each ar: skull (Fig. 2) attned dorsoventally and broudened posteriory: cheekteeth lacking complex folds of enamel snd each appears a single pul of dentine surrounded by enamel, snd maxillary cheek eth wth labial spinelke projection of enarel, ‘whereas the mandibular cheekteeth havea similar gu projection, ‘The auditory bulla is ask-shaped, a portorntal process lacking, the palate i broad and extends beyond the soothrows, and the ‘coronoid process is high and curved posterionly (MeLaugin, 1984), ‘The origin of the maseter muscle fe Bmited tothe ventral surface ofthe aygoma (Kurtén and Anderson, 1980). The baculum (Fig. 3) js thin, convex drsally,eoncave ventrally, and forked distally each distal termins in equipped with a condyle process, whereas the proximal terminus 1 notched (Burt, 1960). The bifurcation was Interpreted to indicate fasion of separate elements thus « homology between epipubic bones in marnupals and the baculum in other mammals Was inferred (Jellson, 19458). The os eltords (ig. 3) isabout 2 by 6 mim, curved, and forked proximally (Scheffer, 1942) GENERAL CHARACTERS. The inciors are strong, fa ‘ened anteriorly, and without grooves: the enamels ylloish orange Choekteeth ase hyprodont (Rensberger, 1978) and evergrowing, and are arranged in straight, parallel rows inthe maxi, hut slighty foneave rows in the mandible, Pie eharactersaed hy the loss of & Inrrer between the iranserse-median and posterinternal basins and loss of the enterointernal cingular sige that brings the prot one tothe margin of the tooth (MeGrew, 1941). Lower molars ‘exhibit ite reduction of the protocond (MeGrew, 1941). The barrier between the two talonid lakes on pé has diappeared eausing the talon to occupy singe large basin, The palate extends beyond the toothrows, The dental formula i 2/2, € 0/0, p 2/1, m 3/3, total 22. PS ts minute ‘The head is fat and wide; the nose “is litle arched” (Ri ardson, 18290:210), The body is thick, heavy, and covered with arse, dull fur. The under Turis thick and covered with sparse tuard hairs: the color is dark redsh or grayish brown. The veatee Fighter than the drsuns (Maser et al, 1981). The lbs are shor and robust, aud equipped with strong, five-ted feet: the sole naked to the heel. The feet ate plantigrade (Hibbard, 1922). The toe, except forthe partially opposable pollex, have log, strong, curved, harp, and leteraly compreseed claws (Lewis, 1949} the claw on "he pole is short an hunt, and has ben referzed to as anal (il, 1937). The claws are crear or light ta. The tail in scarcely visible 15 it barely extends beyond the fur “Mean (SE) monthly mass of juveniles ranged from 357.4 19.7 g ange, 280-440 g; n = 9) in June to © peak of 584.3 = 16.2 g (range, 557-645 g) in October for males and from 473.1 35.6 g (ange, 330-590 g; x = 8) in June toa peak of 630 = 10.0 grange, 621-640 g;n = 2) tho flowing February for females (Lovejoy and Black, 1974). Males lost mass over winter, whereas females continued to gain, Meen (SE) mass of 15 yearlings (both ‘sexes combined) weighed 53 tines beween April and December was Frc. | courtesy of the Weyerhaeuser Archives, Photograph of an adult Aplodontia raf. Photograph2 MAMMALIAN SPECIES 431 Se BD ese omm SE SS, Fic. 3. Camera luca tracings of dorsal, ventral, lateral, distal tnd proximal views (lop five) ofthe baculum of plodontia rufa of OSUFW 8742 rom 2 mi. 8, Ui. E Eddyville, Lincoln Co., Organ, and dorsal and lateral views (bottom two} of the om eltoridis of at individual from near Redmond, King Co., Washington. Os clitoris redrawn from Scheffer (1942:44, fg. 1). Distal eat left forall, ors, ventral, and lateral views, 709.9 + 9.8 g (range, 580-910 g—Lovejoy and Black, 1974), Mean mass of adults fuctusted seasonally wth the peak for both sexes in July and the annual low point in March for females and April for males (Lovejoy and Black, 1974:368, fg. 1). The mean ‘of 455 weighings for 109 adults ofboth sexes weighed throughout the year was 806 g; maximum mau for mountain beavers in Benton, {Go., Oregon, was 1,070 ¢ for adult females and 1,130 for adult makes (Lovejoy and Black, 1974), A maximum mass of 1,419 alo wes reperted (Maser etal, 1981). The brain mass of mountain beavers ealeulated from eraniat volume was 7.68 g (Mace etal, 1981), Ranges of standard measurements (in ram) were (Ingles, 1965) total length, 300-170; tl length, 20-40: lengths of hind foot, 55 68: ear length (13-21), Measurements for the seme dimensions provided by Bailey (1936), Banfeld (1974), Coman and Guiguet (1965), Dalquest (1948), Hall (1981), Jameson and Peeters (1988), land Psimer (1954) dd not exceed these values by >2 mim, except ‘ar lengths 25 mm were reported by Dalqurst (1948). However, Maser et al. (1081) Isted teasurements (in the same order) for mountain beavers in coastal Oregon as 238-370, 19-55, 48-63, and 21-20, Ranges of selected canal dimensions (nm) for small samples feom throughout the ange (Finley, 1941; Kellogg, 1916; Taylor, 1918) were: basilar length, 51-1~65.4;leguh of nasals, 20.3~30.9, vidth of nasals, 87-11-15 length of incbive foramen, 4.5-8.7; Zygomatic breadth, 46.0°64.0; mastoid breadth, 41.9-61.2: length ‘of upper toothrow, 16.9-21.0; distance between infraorbital fram {ng, 14.1-16.9; and greatest length of mandible, 1,7-54.9. Mest Fic. 2. Dorsal, ventral, and lateral views of cranium, and lateral and dorsal views ofthe mandible of an adult female fpo dont rufa (OSUEW [Oregon State University Department of Fish tries and Wildife mammal collector] 8743; from 2 mi. 8, 1 i. E Eaivile, Lincoln Co, Oregon). Occiptonatl length is 66,7 mun,MAMMALIAN SPECIES 431 ‘of the dimensions for 4. phaea and A. r.pacifca were near the low end of the range, whereas thowe for Al. rainierd apd. californica were near the upper end. DISTRIBUTION. The divtsribution of A. rf (Fig. 4) sin four disune: groups of population. The largest, encompassing ranges ‘of four nominal geographic races, extend fom near Merit Bish Colazabis, south along the Cascade, Olympic, Coast, and Siskiyou ranges to Rio Del, California; another extends from Mt. Shasta, California, souheastward through the Sierra Nevade Range of eas. cern California and extreme western Neva: and the others exist as tiny coastal populations in California, one at Point Arena, Mendocino (Co, the other near Point Reyes, Marin Co, (Dalquest and Scheffer, ods, Hall, 1981), FOSSIL RECORD. Tho family Aplodontidae i descended from the Proscisrnae, an early to middle Oligocene radiation fl lowed by the Allanyinae an Oligocene-Miocene radiation: and that followed by the Aplodentinae, a later Miocene radiation (Kurten and Anderson, 1980: Rensberget, 1975). The apladonteid rodents are represented by the extinct Mylaguuidee and the mostly extinct, Aplodontida; only the monotypic gens Apledontn x extant within Aplodoatidae. The Mylagaulidae represents an early to middle Mio- ‘ene radiation from the early Miocene aplodootids and exhibits extreme divergence and greater specialization. The most notable characteristic af the evolition of the Aplodontoiden isthe marked progression wo greater hypsodoaty (MeGren, 1941; Rersberger, 1982) ‘A indicated by the foil record since the late Oligocene, the geographic distributions of Aplodontia and its ancestors changed litle. There were « few unsuccesfal expansions of the range into Mongolia in the mide Oligocene (Wang. 1987), Nevada and Mon: tana during the Oligocene (Donahoe, 1956: Ressberger, 1981: Shotwell 1958), oorthcental Oregon inthe late Oligocene-early Miocene (Rensberger, 1983), and southeastern Oregon in the mi. Miocene (Shotwell, 1958). However, the mylagauide expanded thei seographie datribtions into the Great Plaine during the late Miocene nd carly Pliocene (MeGrew, 1941; Shotwel, 1958). dplodontia ‘was reported from thre late Pleistocene faunas in northern Cal fornia, Potter Creek, Samwel Cave, and Haver Cave, all within its present distribution (Kurtén and Anderson, 1980) FORM AND FUNCTION. The kidneys of mountain beavers ae nolbated, the simple pli lacks proce nthe tral al ‘comin he aren Erion, popllscoverd wth tential ‘Spe staid eptnium, nd have no Lys ater areola ‘ere, real pple, opel ven rect tne tone ofthe med lary sobtance (House ea, 1963; Neng ea, 1960; Nu fever an Pieter, 1965; Pier, 1966; Pier ct a, 1960), Fritts. (1960)ropored hath kineys have tort mas redilary substance rate of I'l; however, Singene and Pfeer (1965) reported worl mass :mevullary mbeance rai of 1 529. The tlomera near the srace ofthe crn have mean SE) dlameter of 2 an range 9-115 pn) sl are conned 1116-27 glomerl/ma The lomerl inthe uxtarelary region fave a mean darter of 140 pm range, 112-203 ym) ana Concerted a 9°17 glomerular The eaal vephvon ae srst abundant in the oster corer hve short fogs of Henle hat not ener themed, lc tn spent, so cosine 62 0 of the Kidney nephro (Perea. 1960), The medley Teper hae oly afew tn segments non of wich ee eri with baron tro. Unasal caret of te ideys are that the loop St Hone hes hairpin troy oly ithe tick sri, portion oo lope conect the arterse reine and vena rc fd the clloting dt ecades ori erp i the cores (sor ‘tal, 1960) The anatomy of te kenny euportlaberatry Sings that nontain avers conan produce byperionicure “Iplodenta conse (2) 327 24 ml/kg (>1,000 ob serrations of water ech 24 yuo cup a the Ste ge at Bers 25.7 ml/kg (2.000 observations Ningesr end Pier, 1965). The amout of water conmuned was ively rested to he tout of sacelent fd inte det. Arye produces ite at {tre ofonefourth to one ote body mane daly (Nangeter ta 1960) ean (SE) arn fw ws 9.03 © 0.68 ring the day and 6.7 20.9 ml/h at night an eine concentration wes 220°" 14 rem and 3148 © 34 Oem rempectal ‘The “reduced urine flow and increase in osmolarity at night 7 rst be do nresed bua abortion ol water” (Dicker ad 123 2 123 120 Fic. 4. Geographic dsuibution of Aplodontia rufa. Subspe- cles ate: 1, 7 ealifrnicns 2, 4, humboldtiana; 3, Ar. nigras Sor. pacifca: 5, Aur phaca: 6, Aur rainierss 2, Ar raf (redrawn after Dalquest and Scheffer, 1945; Hall, 1981), Eggleton, 1964:187). Mean ereatnine excretion was 1.78 2 0.07 rg’ during the day and 1-74 + 0.16 mg/b at night (Dicker and Eggleton, 1964), Hematocrts range from 28-42 and the plasms ehlride concentration is 95-103 muliequivalens/1(Nungesser and Peer, 1965). The range ofthe rine plasms ratio i 1:2.0-2.46 (Nangesser et aL, 1960; Nangesser and Preifer, 1965; Plier et al, 1960}, Maximum urine concentration i $00-500 mOum/l the4 toaximum plnma concentration is 400 mnOsm/ (Greenbaum and Dicker, 1963). Reporte, the kidney mitochondria are anaes sy vampresinGroenban and Dike, 1963) however vesopresin reduce urine Bow, plasma sxolarty (6 293 mOsm /I eresine ‘keen andthe rie pana rato 1:14 (Dicker so Eggo 1964), Variations i nubular water reabnrption ene utes and tedium concereatins te yary tery inthe trine Senki Nien ‘snd Pio, 1910), The “aban of lope vss eta dplodontie may prevent the eficen tepping of sole such ex utes inthe talry inert may sou in par forte pr ne concern sites Hoe tl, 1963:188), Breas othe peor tin concentrating able ofthe kine, the concentration of ex reted sles inthe urine was almost sential to tat in ples thereby requring hgh sake of mater by roustin hesers (Nu agrer and Pier, 1963). This ay xooun or the {Estribton of plodontnto the “wet west slopes of the Northwest (Nangee and Pfeifer, 1905:298). Rectal temperature varied fom 344°C after 3 period of inact, to 36°37 ding sore pros of inact, fo 37. 5~ SHC daring normal activi, end to 38°38.5°C during exended {orod acy Fier, 1965) Te iver of mountain besvers ask, ‘avanti blr parte and the hepteytes haves raat plementation eompored of melanin (Newun and Syder, 1978) "The molars of 4. rae ae characterized by Bt and ped Ungual enamel crest om the och srface that lacks detitl sraovesRenaberger. 1970), The lua eae hs «cores ‘urface, an rode and interprimatcsuberances equally ren tent chemical reser Mon Toth wear mowntin beers Caued by contact rather than fod abrasion. The ater edge of the tet wears faster than the oeckal race but the Bagul lamina of cementum preven the formation of large Hake pis characteristic of tet of ther berivorou rodents (Renaerger, iste) “The cochlea of Aplodontia rf has three an oe al rr, ot which «tart port ef the upper vara may be closed, and the teala tympani conntited inthe upper one and oneal ture (Gerseich et al 1973}. Mountain beavers ave an uneallarge Sorat eeclear nucle eharaceriaed by ony 4 small anon of Inna orgsiatin,heviymyenated Sry in the cence, st'The even large cochlear gra fd Saracteried by nerons 4-5 wm indmete tat hae “cel that occupies about bal ofthe rome ofthe miclews”(Merseich fal, 1918-335). Ths speiliatons ae ved to alow oun tain beavers to eter aw changes in ir presare win their brows (Meraeich eal, 1973) The nadory eles ate mach Le thow found in Side except that the new hss coal prominence between tearing surface, he tps hae rane [een aperture inthe bute between the anterior and porter cure td the males lacks seit apap tae ca and ‘cephalic processes (Cockerell, 191¢ "The quadangiarhaped seapla is formed by lage thin plate it's thick angulated verteral border, «high spine fr ache oft rbonbot ante seratan thd leo scapole ‘mineles, nd large metaroon (Hil 1987; Levin, 1949) The tong ard carved cli bas spa hap in te mile ede compre toward the en (hewn, 1949) The jen permite cesar moti because oft ose cape: The hu ‘er hs mulpe large, rough attachment srfces end oches sith a dep sles end medial Hp (Hill 1937 the ead of the Toner may oe and sem age for the rough rsmatie tein proximal tows The un bas # lng wed beary Slcranon that provide conierabe mechanical advtntage to the forearm tren trl Kel ecenting from he sarsnon tte proces ad served in two planes (Lem, 1949). The ran has fo paeodsylid proces (Hil, 1937}, Th femur has an incomplete trochanter ridge uno third tocbarer. "Te sacral yertirac tong fed andthe mbar tranavere procenes sr reduced” ry. 1981185) Inthe heed and neck region, mountain beavers lack some segments ofthe mphineter al profandsmsl, there na reduetion it'tumber of sient and son ofthe roster marl, ‘conrclar mele, and muscles on the drm of the head, sn 1 singleton of the nes, -ygoateails, masala, rotund, and bucnatree mois (Ningener, 1970). Redsctin Sf the cervcoarcua is svcd ih treason nthe sae of the pina. The syahyotecs muscle originates at the sgl carulage ntead ofthe Jaguar process atin pocket gophers (Geo MAMMALIAN SPECIES 431 snyidae—Hlil, 1957), The jugulhyoious muscle, which does not cain mio oreo rer rgnates onthe gle process Sd ise onthe sigbyl cartlage A few sel er oft eviglnin mole ter on the hyd apparetin. The wide Ho. Goons muscle extends ateraly fon the fod tothe tongue. The Syms orgnates fom the spbhyl cartage, stead of the jie proces gn geomyi anderson the jguahyod stead ihe toga al 1937) ine rpm of te pst te ge ee as unsuly massive tes thom, ecptenepar, singe Aoi eater scape, aed ster vrata mince tach et imme veapua (Levis, 1949), The aanorcpolarn sce ners nthe acromion. Te wicep, and aime te cone and dorsal epicohlear muscles allow forte pomerfl extension ofthe free. ‘he exor muse ofthe pectoral de, the coracbrachia, Heep sd brechs, are ont reduced in st. he spnater and po. mute are well develops however, the rate of the eee eee ae eee seatieceare tt erences a Soourr meme ae ee rea Bitter Sees ie eee eee eee ee Sl ti a Se ete ac ee eae e pee oeore ees artery (Hill, 1935). Inari sets (fen, 1971) The pas dorsal pervocelars has Gemenici ems gua diireeet fete tee aaa beep eet rs mond eche se ie ee eT Aplodonita females havea typical duplex ters, an st Short mesulpins that connects the oviduct ap ovary but dacs not fechas sealant ear eae sees eae gage meenaMAMMALIAN SPECIES 431 hem stage, flowed by nonvsclar, but functional, cho Tone places Ao, asco nonabt, basco havi tir pleenta located inthe mesometiam flowing the fnbbod sage (army. 1953). Splodonia males have penis ce. 45 mi lng: pied lip cl capo. end tbat lourtiral land e. 5 tn tog; sd cmial vee lost within the vente fscio othe provate gla (Pleler, 1986). The senal vous ave appre Snip in beeing nda when the vlc ae 1 rm lng at ‘permatona areca, 100 jan long ad “usualy bea are ql cepts wen ota! fromthe ves deferens or epi” Pier, 1350059) Searnal varaon in the plage i alg the won mnerost is abwaye proven and there singh somal mole Stangl and Genes, 1883), Guard fais are 17.5 by 0084 mm mn summer, 21.8 by 0.081 mmm wir, and cca in ero scton, Pega {nd ating fre have patches of cera ck ais rou the sigles (Peifer, 1935) In the sel regions othe shdomen, the It faces are larger than flies hctedckewkere onthe a Sen an pgs nr dpi veg srammary lan the, idee than srotning ton. Exe Simvelved in these aia ie chenges wien eres rlieratin of the urine lands (Per, 1993) ONTOGENY AND REPRODUCTION. Onset of duce sty inthe mountain beaver evidenced among, mlx ty rapid increas in se ofthe tne, al prostate nd bulbourehral fiance, and bythe tenes taining a"wemacrotl” poten (4. rf Esno uve seroma Hubtard, 1922) in November or December (Gtr, 1992; Lovejoy e aby, 1978; Phir, 1956:632, Bg 2) ‘a. (1978924) reported that in captivity Both sexes produce ‘cloudy yelowishanlly wre” cts season shen peed in a handing cage (Black and Hooren, 1973) At Temas exhibits hypervoptied ripples a realy ewolen ail smear showing 4 clowtof cored epi ‘elt (feo, 1986.25). The preovastry ovary tay oman flicles as large an 3mm in ameter (Peer, 1956) Yering female ovate (Corpora ea are formed) but oot breed (02 spermato found i reproductive tact thin, orton f consid: rel wo be sponteneno Peer, 1958) “éplodencia raf has a tngle annual estou period with con- sidersleintopopation synchrony among breeding female (Pf fer 1958), Onset of enrs seminal varies with lose a xtinated dies of prrrtion(n'= 1-5 lel) wore au ely a6 Febrany in Maria Coy California, and ar ite se 8 May th Teflerson Co, Washington (Pfeifer, 1958), Date of parturion vas not rlated strongly with ute a i was 20 March for Placer Ga, Calor 22 March for Lane Co, Oregon: end 11 Mazeh=-3 Api for Ki G2, Washington (ey 1588) Duran of gation na com fred 2880 days by Seheflor (1929) and Pfefer 1058) hut 6-8 senks by Hubbard (1922) nA. rufa, itere of two and tree ate common, four ls so age AB Lory a Bik 74 ong YR, Pile. 1956, Seber, 1929, 1980, Seton, 1929) Lier se, Iced on embry coun, wa roported ar even feraee with two cryon and ine with thee Sheer, 1929) Ir Washington, mean Attra (n= 12) was 28 sed courts of eopors tn ad 2: based on counts of embryos (Peer, 1950). Based on counts of pigmented sites of inplantation, 25 females had mean of 2.5, Soumgl tree others, with an average of 27 corpora ven, had an {verage of 2.7 plemented sits of mpastaton (Peter, 1958) “Yong ae torn eter bea rst oi the Breech presentation (Cramtiet sot Rilenbnur, 1956). A: hit, Soong are fink nae, Netplen and nd, but ale to emi squeaks and iin 20 mn 10 tue (Cramblet and Rienbour, 1999) Vibraas ae vil, it Auditory cals and eyes are closed; the ear formed but not tendo the foot tons are well developed ant clawed, the bind es mere sub athe al pices (Crambletand Ridenour, 1956; Lovejoy et al., 1978). Sixteen hours after birth one of three Young weighed 25'5g and wan TO mim long 25 afer bith the Three young weighed $5.0, 26°, and 90.5 and were 85,90, ac 87 eum long espectvely (Cambie ane Rierour, 1956). These tires indvtal conoued fo grow an kngth, bu gained itn imax after the rd day; all ded onthe Gh day air ith, Two Otber indus weighed 18.0 and 21.0 gant averged 66 mm tong 2 day ster tey dif or qa st 3 day (Lovey eal, 19TB) a 5 daye the akin was dary pigmented on the 5 dorsum and pates et 7 days, the skin was wrinkled, especially on ‘the hind lgst at 9 days, a sparse pelage of ie hats covered most of the body and the pinnae were free and erect, at 10 days, the tye sits were formed, but both eyes and ears remained closed at 1H days, the pelage was complete with dorsal hairs dark and sleek, ‘bur ventral hairs were white: at 15 days, the young began to craw solely by use of the front legs at 21 days, the ears were open, and the hair was dark and mace brownish: at 28 day, the lowe incisors crapted; at 33 days, the upper incisors erupted and one individual teas obeerved to erateh its face with a hind foot at 35 days, the ‘head was elevated and crawling by use ofall four feet was observed: fat 38 days, young began to eat solid food: at 45 daye, one young had open eyes but the eyes of the other didnot open unt 9 days later (Lovejoy etal, 1978). Average mass for the two Young was 27.0 g et 7 days and increased exponentially to 364.5 gat 60 days (Lovejoy et aly 1978:327, fig. 3 ‘The two captive young accepted natural foods in week 7, but ‘continued 10 muri in week 8; thereafter, both commenced to lose veeight and both died in week 10 (Lovejoy et al, 1978). Maternel ermales lactate for =2 months (Plier, 1958), nthe wild, young mountain beavers probably are weaned by 6-8 weeks while sill inthe nest; thus, they likely feed on plans tarred to the nest by the maternal female (Lovejoy and Black, 1974). In Benton Co., Oregon, the earliest capture of « young-f. thesyear was on 3 June: it weighed 280 g, thus was considered to be about 7-B weeks old (Lovejoy and Black, 1974). Mean rates of nin in mas among repeatedly lvetrapped juveniles declined during June and July, platesued in August, end remained so throughout the juvenile period (Lovejoy and Black, 1974). As yearlings, some individuals doubled their mass, attaining an average mass >740 g within a year of becoming independent (Lovejoy and Black, 1974). Deng tp fennel damn feuadieed teste ahr Es canis eon pislrrecree tints sh amp ihe neia Scr ee feet aston oe ieee eorea reas Sore senate nt eee pean err carte habeas mes ee eae mount weavers (Camp, C518) Aig mumerom abet sane pee ee eg nee alae! he Haat requrenents or 4 raf do not ee to hate ss Fo cumin ego eet facie ane mira ee a er Ee oe ‘polh of excavate earth ae formed at only few openings ne oT earaaae car meal cgtnte Wig ny atari emt eames (a eg coe meester eect euiteatnat ryan tien eee coum erect cisauamits bitomemevaten ISathipaencieis caheucae rare deodorant cians foe tania del geen na Ep bow anmueageraie ce aciciperad trand oe ah manana Wisma madtace cornet agree aia a ies ae ge oe eteoenathataseRes One siyc Sacer ne ah Gini aaa lan nena sag fein game ec serenade ena6 simated dense were 83 and 1.3/ba on 7- and ba reas respectively (Motobu, 1978). However, « population density a high 6 15-20/ha was repored (Hoven, 1977) desertion of the ‘methods on which the eatinate was bsed was ot provided. Lovejoy {Ed Blck (19794, 19798) eapred 181 nds in 5,168 tap tights ona 5 Schasrea in west Oregon ding» 20-mont pri: 10 ofthe mountain Beavers hey captured din tape, To ater suthor etnsted denies of mountin beavers monthly by dct on ona hn area (5.5 raping grid plus 40-mwie Tundary stp) at 4.1-3.4 animale ha By ang 880. boundary strip ean distance between eapres) ‘0 areas sampled by Camp (1918) and Voth (1968) estimates by those authore wore nearly ‘ental withthe mimi estimate of Lovejoy and Buck (19790) Neverthe, Lovejoy and Black (1979) conslered thei extinaes to be conservative nd prevented an estate bythe Lincoln Index for T month that cased tem to sugges tat the dectenumeration ‘uehod praliced underestimates othe population tacy sample. ‘rapped juve (2 ='72) wax exci (ee eh) wae in va mae 1B1) wae 11.5 in favor of adults, icing earings (Lovejoy apd Back, 19796). Lovejoy and Black (1979: 436) comidered the deren among sexes to bea “tre ratio ad ot an aac of tapping but provided no evidence in spor of thei contention, OF 45 animals marked by Lovejoy and ack (19796) during he Gest month of wads, 29 (64%) were known to alive I year later and 11 (24%) were known tbe alive 2 Years later Sore of thee idvidsls mere sexually mature when Bet captioned yeu i who ut aptired: Boel on e estimated longevity 5-6 years (Lavo and Black, 19795), During election of 75 A. rufa in Washington, Oregon, and Calera install legal tape atin runways, nine Splogele frail, four Mustela fenotas one M, ermine, ove ME. son, Seht Neotoma fseipes oe N. cinerea, two Tamiascarus doug: ‘one Zapu prince. four Sylelngus backmams, one Microts Califoricss one. nchardson, one Thomomys btvae, oe Peo ‘myseus manieafatus sod one Dicamprodonensais were ea uted (eifer 1985). Other were of mountain beever burrows inelud Neurotrichas gibi, Scapanus orarus, Lepus americas, Clete omomscalforneas, Pkenacomya albipes (Maver ly 1981) ‘Mophiis mephits (Camp, 1918). Procyon lotor (Lord, 1866), Tasiden tax, and Mores pennant (Lat, 1878). Temi of A rufa occured in only 12 (8.47) of 143 fal eoppings of Felis rues calected zoughout te Coast Range of ‘Oregon consating only 5.3% of thease of prey eaten Nusa fn Maser, 1975). However, a the Cascade Monat uf Oregon, A raf occurred in 87 (14.1%) of 404 fecal droppings of Fu {oweil'andArshony, 19880) and in 70 (8.9%) of O44 fecal, Aeoppings of Can latrons (Towel and Anthory, 1988). Where carnivore poulators were nat expltel inthe Coast Range of ‘Oregon, remains of moantain beavers oocurred in 24 (79.0%) of 309 fecal droppings of Claus, adn 182 (73.7%) of 247 fecal Aeoppings fF rs (Winner and delet, 1906)-In Washington, 20°85 ofthe composition of feces of C.Tatrans was renaiteaf “4. raf in May-tly. tony 5% 9 Augist-Septerber (Brigham, 1954). Mosntain beavers occree in one (67%) stomach and two (9.3%) colon tract of F. concolor from te wertera Cascade Moun. ‘ans of Oregon (Towel and Maser, 1985), Skunks and weasels tht te mountain beaver herow systems may prey on nstings and javenes (oven, 1977) Agua chrysacts i known to rey on “A raf Secvbeen, 1978) and "owls ae to be regard os enemies” cam, 1918:555), Ecioparasts cecorded from A. raft include Siphonepter: fleas: Eptedin yordant (Hubbard, 1940a), E. seapant, Cerae- Phyllis {nonopsyias] eiliats protims, Opsodasys een, Or ‘pss [Diamar] montana, Poratyphlocera oegnens (Levis oP, 1988), Dasypsylls gllinalae perpinnates, Dolichpsylla ‘ylomus (Hbsrd, 19400}. Hystrichopsyila schefer (nchales ‘mammoth nd hubbardi—Levis cal, 1988), and Trichopsyloides Gregonensis (Hubbard, 1945), Acarina: mien Alphalnelaps (Ca laps} aplodontinze Ulison, 19450: Radford, 1951: Whitaker et 161910), Patrinysus Uchoronyers]huBbordi, Hoemogamassis ‘eid, Haomogamasss sp. (apprenly 4 new specie). plan fochinus boreal Microlabidopus americana, dplodontopus la ths, Buryparastus sp. (Whitaker etal, 1979), Hirstilonystus oe Cidentais (Steantmana and Morlan, 1958), Neotrombicula plodontiae (Brenan, 1946; Wrenn and Maser, 1901}. ¥. avi ‘ola, Aplodontophila pacifica, Euschoengastia brennan. and Eus -MANMALIAN SPECIES 431 eg mr oC seagate at pe i ie nda ee i ee dae naytretccit bili caatan goer sare Sey aunremne dees ribs pis ontlans el a ga thet ie Ce sae re wong ncn sgelcreaelaes a mene ba Senate avs ety ei mine nega ante sierra ae ret eatin th Cee Aa ape es oar ance gabe eat Vitec st arene istics or aa tect he eel St ite tari ll a ec el een ems eck etait Steen ees eoneta Se ee ae oy mae ee Gens Mn is wa Sm Goren Fase ae Sa paar canoe ieee pene eta ta ee Pia eeemuan maces aedete Ears font i Ra eet ee ea eneee tevwephyla eter 1229) and suneto er sce pas te mmm ew reforestation oy ease exemve damage and conan ow (lack sane alan ghtcianiomenat cae BS sala 2 aly a lg Fc rea raat anc ae at Iai crate aS rie hae ica merge aaa eh pe ceen cine temas Es staat i te fe enc a ae Ese cs ct eae nee Saeco shi ynuieelateponaennie Se cl thenar aie ea aera encod sates Atm “sub oa hao to scent eae eerie a a slMAMMALIAN SPECIES 431 Arie vegetation nearby orn the entrances to mountain beaver dens fd sated or implied that the material was hay being cured by ‘mountain beavers (Hooven, 1977; Lum, 1878; Mercian, 1896; Seton, 1929). Such activity commences ip August (Hamilton, 1939). “Moet authors indicate thatthe material moved into te bureew and eaten, whereas Camp (1918) and Sehefer (1929) claimed that ‘tis not eaten but used at nesting material. Both funtion ae ike the wilted material may be eaten and the dry material wed inthe ‘est (Maser etal, 1981). Hooven (1977) indicated that body mass ff mountain beavers thet survived an application of herbicide followed by a contelled hurn to remove vegetation di not decline, ican that they were feeding on stored material. Voth (1008) hypothesized ‘hat mountain beavers harvested dried vegetation for nesting rm terial, but cut, partly cured, and ste the partially cured mater to tdjost their water intake. An alternative hypothesis war tht the ‘alting and pling of vegetation was eelated tothe security of moan: ‘tin beaver food supplies during periods when foraping was unsafe (Voth, 1968) In northeastern Caornia, food caches accumulated by twoor more mountain beavers were 58.3% Delphinium glauca, 20.7% nus tenuifolia, 18.2% Corydalis caseana, 1.3% Vera ‘rum californicum, anil 1%e other species (O'Brien, 1981). Obvi ‘ously, mountain avers were selective of plants stored ae D.glaue. lum ad A. tenuiflia composed only & and 5% of the vegetative cover, respectively, but unnamed species in caches composed 30% ofthe vegetative cover (O'Brien, 1981) "As a marking technique, 15-35 mg/kg rhodamine B adnin- Introd asa single dose by oral gavage produced bands that fuoresced orange under ulraviokt ight mn lng in il within 2-4 days fand 1-2 mm long in vibrissae and guard hairs within 3-7 days. Freediving mountain beavers were marked similsely by ingesting pple baits containing 213 mg/kg of the dye. Marks lasted 19-26 ‘works in hair and 13-21 weeks in claws of caged enimals and 17 28 weeks in haic but only 6 weeks in claws of fresiving animals (Lindsey, 1983}. A°50% dextrose slution administered by oral rectal gavage reduced mortality in mountuin beavers resulting from shock of trapping and handling: no quantitative data on rate of sucess was provided (Dodge and Canpbell, 1965). Radio trans- Imiters mognted in "beeak-open collars which clouely encicle the animals neck” were used successfully (Dodge and Church, 1965: 118} BEHAVIOR, The mountain beaver is largely fosoril i rarely venutes more than few meters from its DurroW entrance Despite statements to the contrary (Grinnell and Storer, 1924: Schefler. 1929), uf can cmb res Toges, 1960; Seton, 1929). it ases the stampe of previously cut branches uch in the manner ‘of rungs in ladder (© ascend or descend headfirst. It sometines forages by pruning twigs from trees =6 m above the ground Ingles, 1960) rufa ao enters the water and swims wll Anthony, 1916: Ingles, 195), ‘More commonly, to exploit a stable chimp of vegetation, ‘rufa digs «new tunnel to the vsinity and makes short sales to eat ems or fronds which i transports to its burrow where it feeds (Camp, 1918). While eating. rufa grasps food with th cele pollex on the forepaws as tats onthe til with hind feet, fpoctoved ventrally anc soe facing forward (Cacyp, 1918). While ‘tecating, 4. rufa squat with tll and parally ever anus po tioned upward: hard pelts are taken into the mouth and," fn upward jerk of the head.” tossed into Itrine (Kindschy and Lereton, 1961:2), whereas oft pelts are eeingested (gles, 1961). Soft pellets are approximately four times the volume of hard pelt. In the large intestine, soft pellets are intermixed wth hard pellet, and stomachs of sme mountain beavers taken in the wil possess “unmistakable remains of reingested soft ples (nals, 1961). "Although moet burrowing activity odGurs in eure and above ground activity nearly ceases in winter, mourtain beavers do not Fiernate (gles, 1959). In surnner they have Bve-ax bouts of {oraging each 24h period. Nevertheless, they are 50-00% more active a right than during daylight hours (Ingles 1959), prom ‘eports of ther chiely nocturnal activity (Grinnell and Stor ‘Scheffer, 1920), Lawrence snd Sherman (1963) described « device barrows, but it wae notable ge width for individuals eaptured seven or more times at ive o more tap) sites averaged (= SE) 0.32 "= 0.05 (range, 0.09-0.70: 4™= 18) ha for adult ales and 0.17 = 0.05 (range. 0.05-0.40: n = 7) ba for 7 adult fomsles. Sila data for juveniles were 0.13 + 0.3 (range, 0.04-0.24: n = 7) he (Lovejoy and Black, 1979n), Home-range areas calevlated by the minimim-area method for 10 adults of both Sexes radiotracked for 3-10 months averagod 0.12 (range, 0.03- 0.20) ha (Martin, 1971}; both maximo and minimum calculated ress were for individuals mentored for >12 months. Maximum ‘movements from the nest for these individuals were 42,6 and 36.6 sm, respectively. Two of 11 subeduls made extensive ear move: ‘ments considered by Mactn (1971) to be dispersal: a female moved 500 m:a male moved 188, stayed 8 days atthe ste, then moved fan addtional 191 m and rerauined 46 days until contact was lost (Mert, 1972). “The mountain beaver & at a social animal sltbough in some Dhobitats animals may be evonded together such that thoi tunnels “tainly. honeycomb the gronnd (Scheffer, 19294). Indvidoals fight when placed together in captivity, even in O.1sha enclosures . 1964). Although home ranges may overlap, nests and bur roms are defended, with invidaals exhibiting considerable fekisy tothe nest site (Martin, 1971). However, Lovejoy and Black (10970: 400) indicated chat individuals “irequented one oF two nest sites In captivity, mountain beavers reportedly become docile quickly avis, 1941). ‘The mountain beaver has limited ability t0 see and hear, but tactile and olfactory senses are well developed (Carp, 1918; Scbef fer, 1929), Seton (1929) suggested thatthe strong musky odor of the mountain beaver might be associated with inraspeciie com> ‘munication. The vocal repertoire of 4. rufa controvesiah early naturalist attributed whistles and booming sounds tothe mountain beaver (hence, the vernacular name "beomer"), but Scheffer (1929), tributed these sounde to birds and other mammals App the only sounds produced by 4. rufa are a soft whi sound when in pain (Kinsey and Lasts, 1961), 4g produced with the teeth (Camp, TOT8), and hig ‘when fighting (Godin, 1968) oF when apparently frustrated (Kindschy {and Larsson, 1961). GENETICS. In both A,r. californica and A. phaca, 20 = 46 with st pits of metacenric and 16 pairs of submetacentie sutosomes the Y-chromosome is sulmetacentric (MeMilin and Sut ton, 1972). In Ar. pacfca there are five pars of metacentic and 17 pair of submetatentrie autowmes the Yechromowome i met centic (Carrasco and Humphrey, 1968). The karyotype is consid ‘ered wo be advanced becaune it contained no acrocentrieauosomes (MMi and Suton, 1972) Hemoglobins of 4. rufa migrate anodlly Jonson and Wicks 1959), 4 piebald condition on the underparts and sles is comme (Anderson and Russel, 1957: Couch, 1926: Maser ot al, 1981) ‘Mbino Jewett, 1985) and melanie (Fler, 1965; T. E. Lavlor, in it) individuals occur. REMARKS. The generic name Aplodontia was derived rm the Greek haploor meaning single or snp, tnd endear meaning touth, The spetic name rufa was deve fom the La rues dnething reddish Uneger, 1948)" “Aplodoita raft as been refered toby a wide variety of ‘eraciar nmes inching mtn boomer, geal fro, rod iar, giant mole seve show chowa hte Swale ia ei wend ot 1510, 15'1904, 30 vrais on te paling of plodonta were applied in plications (Coes and Alle, 1874: Tap, 1918) ranging rom Hapladon, Hopton. Hepooien, Hopldes nd Hapluonta, to dptuodonti, Aplodonte, ipleodentias and plodonta Because of Ks prime ratre and unas characteris, Aplodnta raf vary was refered to the fan Prenseuantin (mich inched Castor, Mus, an Tartan among other secon Sctrina’'of the family ‘Murhiae, onder Gives, fanly Canicarn (which included arvectines and georyit among others) the Poe Aston (pocket gperst.sbtanly Caraer Ihe poreapies presmaly Exetel tly Sound fou 1829 to 1877 (Taylor, 1918} However, acorn to Ces an ale (187), itjeborg (1866) rated Hlptodon to ail rank Haplodontide), thereby removing fram the aun Castrinae ofthe fay Siti: be omidered tnesextporeipnes intone afi. Subsequently Gil (1872) repre Hapldan from other Ges talents by rng i rar to of pei Haploderce) 1. K. foes ea BP. Lovejoy and. Lawlor commented onan erie deft of the mamsenpt This i Techical Paper No. 9490, Oregon Agricul Experinent StatonLITERATURE CITED Color end dental ab- Kansas Actde ANDERSON, S, AND R. 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