MAMMALIAN SPECIES No. 532, pp. 1-5, 3 figs Arborimus longicaudus. By John P. Hayes Published 27 December 1996 by the American Society of Mammalogists Arborimus longicaudus True, 1890 Red Tree Vole Phenacomys longicauns True, 1890:308, Type locality “Marsh ‘ell, Coos Ca, Oregon, Phenacomys silicolus A. B. Howell, 192198. Type locality “5 tiles southeast of Tillanook, Tillamook Go., Oregon.” Phenacomys silvicola Miller 1923400, A renaming of Phenaco- ‘mys slricolus Howell, Arborimus langicaudas Johnson, 196827, Elevation of the sub ‘genus Arborimus to generic rank CONTEXT AND CONTENT. Osler Rodentia, Suborder Myomoepha, Superfamily Muroidea, Family Muridae, Subfamily Ae- vieolinae, (Master and Carleton, 1993). Masser and Carleton (1993) Tecogrived thre extant species inthe genus Arborimus, and Jon ‘Son and George (1991) recognized wwe subspecies ofA. longicau- ‘tus A. 1 longicaudus True, 1890:303, see above. ALL. slracola A. B, Howell, 192198, see above, DIAGNOSIS. Arborimus fongicaudus can be distinguished foe all ether sympatric arvicoline ments by the combination of {reddish dorsnn lacking median stripe, s venter washed with redish-orange hair, and a well-haged til thal is not dstinely bicolored and is >50 mm long and >S0% of length of head and body (Maser and Storm, 1970: Vers and Carraway, 1984). Skulls lof the genus Arborimus can be separated from those of other a icoline genera in the ange ofA. longicaudus by reentrant angles ofthe lower molars extending about two-thirds the way toward the labial side on the lingual side and about one-third the way towael the lingual sie om the labial side. The skull of 4, dongicaudus can bre distinguished from A. albipes by the combination of & pro ‘nounced longitudinal depression between the orbits tt is bordered by a strong rie, strongly recurved incisor, and wide inciive foramina (Maser and Storm, 1970). In contrast A. pomo, A. lon sicaudus is generally larger overall, has nasals that extend farther fosterirly relative tothe maillaris, lacks the abductor digit se und muscle and a full complement of lombricales muscles, and differs in chromosome umber and morphaligy ahnson and George, 1991). GENERAL CHARACTERS. Arborimus longicauius is a small (mass 25 10 50 g} miceotine rodent (Fig. 1) The dorsal pelage is uniformly cinnamon (Hall, 1981) or height rusty brown (Tie, 1300) with many hes tipped in black in. lomgicaudas, but light (cayal) brown sparsely tipped with biack in ALL. sivicola (Hal, 1961; Howell 1921; True, 1890} the underparts are whitish in bath subspecies (Hall, IAT). The til is long and wel-haited, varying fom black 10 brown and not distinlly bicolored (Maser and Storm, 1970). Juveniles ate duller. with less ted and mow brown in dorsal coloration and black til (Maser and Storm, 1970, Melanistic forms of 4. silecola can occur There are ix plantar tubercles om the hind fot and five on the forefoot (Hooper and Hart, 1962: Taylor, 1913). The claw on the fist digit ofthe front foot modified int a nail (Keser, 1986). There are four inguinal marnmary glans (Howell, 126: Johnsen, 1973; Taylor, 1915). The ears are 10-12 som long in adult, with sean! fine hair Jobson, 1973), The skull ofA. Zongicaudus is heavy and moserately angular. ‘The nasal hones extend beyond the incor (Maser and Stor, 1970; Fig. 2). The dental formula i 1 1, © 0, p (VD, m 303, total 16, The molariform teeth are rootless in juveniles and become rooted as the animal matures (Hooper and Hart, 1962: Howell, 1926), Rooted molars generally are considered to be an ancestral character forthe subfamily (Carleton, 1985). Roots of the lower incisors extend slighty posterior and Labial M, (Hooper and Har 1962}. The incisors are sharply recurved (Maset and Storm, 1970) The cervical vertebrae are short ae close tether, the tans verse pracesars ofthe lumbar vertebrae are large and faring. The Inoil bone has short lateral processes and no spine ora reduced Gone. The aeromion proces of the scapula is narrow (Johnson, 1973) ‘Gm average females are lrg than males in inost external and cranial messurements. Cranial or extemal measurements do tot differ distinctly between subspecies (Johnson an George, 1091), Jehnson (1973) gives ranges for distema (6.1 to 75 man 35), minim or maximum values for seven measurements, and qualitative descriptions of 22 morphological characteristics, Mini rn aed masini mean values (un) of 14 eranial measurements for 7 females grouped into seven geographic samples ad 50 males roped into four geographic samples (Jshnson and George, 1991) fre as follows (values for ales listed fis) greatest length of skull 24.42 and 25.38, 21.81 and 26.06; nygomatie breadth, 13.80 and 14.50, Th] and 14.73; least interorbilal width, 337 and 3.48, 3.22 and 3.59; mastoid breadth, 843 and 8.69, BAT and 8.86: length of nasal, 6.26 and 6.94, 6.50 and 6.85; length of maxillary toathrow, 5:56 and 5.86, 5.50 and 5.Xh depth of rainease, 6.93 and 7.16, 7.02 and 7.23: nasal-masilary distance, 44 an 0.56, (0140 and 0.66; wich of interparietal, 8.17 aed 8.61, 7.83 and 2.02; Fic. An adult Arborimus longicauus fom Benton Coun- ‘y, Oregan. Photograph by Stephen DeStefanoFic. 2. Dorsal, ventral, and Ineral views of erenium ad at ral view of mandible of Arborimus longicaudus trom 7 mi W ‘Alpine, Alsea Falls Park, Benton Co. Oregon female, Oregon State University Department of Fisheries and Wildlife, mammal collec: tion no, 3384). Drawing by G. Bracher. Tength of inteparietal, 3.16 and 3.40, 3.20 and 3.72; M2-M2 width, 5.49 and 5.78, S41 and 5,84; lngth of diastoma, 0.484 and 7.14, 690 and 7.5% width of upper M2, 1.28 and 131, 121 and 1.35; and length of palatal foramen, 4.18 and 450, 4.24 and 4.69. Mean external measurements (om) and extremes (im paren= theses) for six adult male and five adult female A. longteaudus (Hall, 1981) are as fellows (values for males listed frst: total Teng, 166 (158176), 182 (170-187: length of tail, 67 (60-72), 73 (66-83), and Tongth of hind foot, 20 19-21), 21 21-22). Mew surements for one adult male and one adul female Af silisicola (Hall, 1961), respectively are (tal length, 193, 191; length of ti ‘il; and length of hind ft, 20, 22 DISTRIBUTION. Arhorimus longieaudus is restricted 10 conifers forests west of the crest of the Cascade Mountains in Oregon and northem California Johnson and George, 1901 Maser, 1065; Mureay, 1995; Fig. 3). Precise limite and extent of disti= bution in northern California, scutheen Oregon, extreme northwest em Oregon, and parts of the Oregon Cascades ae uncertain, John Fic. 8. Distribution of Arborimus longieaudus: 1, Ak lon= icaudus: 2, ALL silsieota son and George (1991) regard all California tee vole populations fs A. pomo. Murray (1996) classifies red tree voles in noathera fornia as A. longicaudus on the basis of genetic similarity Johnson (1973) speclted tht the genus Arborimus evolved in the Pacific Northwest during the Plicene, but there are no know fos sils of A. longicaudus. FORM AND FUNCTION. In red tee voles, the palatal pore tion of the palatines is Tong: the basisphenoi is short, wide, and relatively shallows and the aucktory bullae are small and thin= walled (Hooper and Hast, 1962). The anterior longitudinal ridge of the diasteral palate is constricted and depressed i ts mile ates, with the medial azea exposed and the incisive papilla not covered (Quay, 1954, "The abductor pollicis brevis, abductor digit quit and flexor digit quin brevis are present in the mans of A. fongicaudus but fabsent from the manus of some ther micrlines. The adductor digit secund was absent from specimens relered to a8. silsicola {presumably A. I. sileicola) but present in specimens referred to 35, A. longicandus (Kesner. 1996). According to Johnson and Ceorge (1001) specimens refereed to as A. longicnudus by Kesner (1986) fare actually A. pomo, but no locality information is provided by Kesner (1986). Specimens refered to as A. longicaudus were the only microtine specimens examined with a fall complement of Tumbricales museles to digits I through V with a slip to Tl In specimens referred to as A. sileicola, a complete complement of Iulvicsles muscles was not present, OF the micrtines examined, the palmaris transversus muscle was anigue to the genus Arbor ‘mus. The palnaris tansversus is @ thin muscle originating from the radial aspect of the metacarpophslangel joint of digit V and ineerting into the ulnar aspect of the metacerpophalangea joint capsule of digit Il. Kesner (1986) concluded that modification of the palmaris transversus, presence ofa nail on digit I and retention of several muscles los in other species ate adaptations for food handling and for morement on terminal branches of trees: reasonsMAMMALIAN SPECIES 582 {or diferences between specimens ceferred to as A. longicuadus and A, silvicola were not explained. In general, the masticatory musculatuce of A. longicaudus is Iypieal af most micrtine rodents. The re re vole differs from the {ypieal mierotine plan in having reduced posteodorsal insertion of the superficial masseter and a eoresponing lateral process om the angular process ofthe mandible, and in lacking dstinet mesial portion of the posterior belly uf the diapastte muscle (Kesner 180). ‘The penis is heavily bowed in lateral view, corresponding to the shape'of the baculum, The glans ofA. longicaudus han exten or surface. The dorsal labe ofthe glans i 4 single spine-tipped process on exch side telatvely shore wi (oper and Hart, 1982) “The base ofthe baculum has a highly ireyular proximal ede (Hooper and Hart, 1962) thats triangular (Hamilton, 1946) to semilunar (Dearden, 1958) or rounded (Hooper and Hart, 1962) ‘The shaft is bow-shaped with = spatulate, almost tiled head (Hooper and Ham 1962). Theee distal igi processes are carti= laginous (Dearden, 1958) or cnsfied (Hamilton. 1940). The distal algal processes are ankylosed in some individuals (Hamilton, 1946), but not in others (Dearden, 1958). More complete ossifea- tion and ankylosis may be more typical of alder a ind Hat, 1962). The hac frm a single spec length, with a shat length of 2.75 mm and a median process 0.35 mm long (Harlin, 1946) Arborimus longicaudus possesses sebaceous, sudoriferous, ‘and mucous glands ofthe oral lips and oral angle (Quay, 1965), thu lacks posterolateral sebaceous glands (Quay. 1968). Number of tarsal glands in the eyelids from four individuals ranged fer 15 10 19, with an average of 17 (Quay. 1954), ONTOGENY AND REPRODUCTION, Red te sles ex- hibit postpartum estrus and effective posipartim mating in the wi ‘Brown, 1964). Reproductive biology of A Zongiomuds is probably Similar to that reported for A. pomo by Harmilton (1962}, but no tailed studies af the species have been conducted ECOLOGY. Red ee voles cccur in second-row forests {Brom 196: Can av Bry, 186 omet, 190 ner 1950) 1y be mare numerous in mature of oligrowth stands (Corn ‘By 19008 Cibo an Alc, 1981) Soe coe the Species to be closely associated with the old-growth seral sage (Huff et a. 1992: Ruggiero etal. 1991), ed ire sales most frequent are found in forests dominated bby Douglashr (Pseudotuga mensiew) Int slso occur in stands of grand fr (Abies grandis, Sitka spruce (Picea siichenss) and ester hemlock (Tsuga heeeropkyla: Jewet, 1920; Maser, 19650: Walker, 1930). Red tee voles occur at elevations of seu evel 1300'm (Huff eta, 1992). [Ness ypically ate arboreal and ate found in Douglas-fi tees, Dut ahey can occur in a variety of species of ree, including grand fir, Sitka spruce, or westcn hemlock (Gilleberg aul Carey, D1: ewet, 1920; Maser, 1965a: Walker, 1980): one nest was located «a big-leat maple Gleer macrophlum: Maser, 19634). Nests may be constructed by A. longicaucls, but areas the majority af nests used by ted tre voles are hilt inital by other species (Maser, 19650). TR tee vole nests may be constructed on the base of a bids rest of may have heen absndoned by other mammals, including vwestem grey squitrels (Sciurus griseus). Douglas squirrels (Tam= faseiurus douglas. wooseats (Neotoma fuscipe), an eer mi (Peromyscus maniculatus; Maser, 19650}. Neste built by Arbor ‘mus longicaudus have heen simultaneousiy aeeupied by red tree soles anid Peromyscus maniculatus Brown, 1964) and by sever species of invertebrates (Maser. 1965a). A nest built by Seiurus ariseus was occupied simulianeausly by S, griseus and A. ongi= ands Mas, 15a) i constructed by A. longicaudus olen are in contact wi the wee tank ut abo oar ches ana few he tae ies, oF in dead tops of trees (Cilesbeggaral Carey. 1991; 1965a: Taylor 1915: Wight, 1925). Neate ae located a a fare generally near a source of ive neces, and are mast abundant in the lower canopy. Nests are cispreportionately common, in large-diameter tees, but are found in Trees a smal 9 10 cm in diameter at breast height. Often several nests are found in 3 Single tee (Gillesberg and Carey, 1991; Maser, 19652), 3 ‘Nest construetion begins with platform of twigs with needles sed for food The platform is se asa lateine, and feces and rine accumulate on the nest, Oser time, additional forage = brought to the nest, and the nest compacts with use. Red tee voles move dlisearded resin duets tothe sides of the platform, forming a cup like depression. Additional resin duets are positioned aver the top ofthe depression, forming an encloned nest charrr. Feeding take place both inside and on top of the nest. The nest continues to tow a the inhabitant adds new twigs and deposits feces. Feces ‘are found throughout the nest, except in the sleeping chambers ana Tunnels. The cermpacted feces anchor the nest to the tee (Maser, 19650) (OF 117 ced wee vole nets, $15 of the volume consisted of resin duets, 19% of lichen. 13% of focos, and 99% each of needles and fine twigs (illesberg and Carey, 1991), Nests are sometimes Tipped with freshly ext twigs with green needles Jewett, 1930), Nests vary in size an may be up to 1m in diameter (Taylor, 1915). Nest site height and form clo not consistent differ between males and females. Mean height, ith, and length fem) of wi neste oc ‘cupid by males were 25, 28, and 32, respectively: those of 21 nests occupied by females were 28, 37, and 44. Neste vary in siructute ut generally havea tne leading fom the internal nest, ‘chamber to the bottom of the nest, one or two tunnels lead the top af the nest, and a system of internal tunnels (Maser Male and female 4. fongicaudus tend to occupy separate nests, although oreasionally an edult pair wll occupy @ nest during Ineeding (Maser 1965 Waker, 1928). Newts may he used by ml tiple generations of re tree voles, There is no evidence of uili= zation f terrestrial nests by A. longizaudus (Maser, 19654) The northern spotted owl (Strix occidentalis caurina) is prob- ably the principal predator of red tree voles (Porsman, 1976; Maser, 1965), which comprive nearly 50% of the prey of spated onl some locations (Foesman etal. 1984). Other avian predators in ude great horned owls (Bubo irgimanus: Maser, 1905e), long ‘are onls [Aso atus; Reynolds, 1970) and northern saw-ehet owe (egotius acadicus: Forsman and Maser, 1970). Racooons (Pro- «yon Toto). marens (Martes americana), a fishers (M. pennanti) fate among the potential mammalian predators in some areas (Maser, 1965.2 Maser et al, 1981), “Arborimus longicaudus and its sibling species, A. pomo, ae 1 on conifer needles for forage. e voles is needles of Douglas-fir, but needles of other conifer and bark are sometimes consumed (Mase 19652: Walker, 1930; Wight, 1925). Captive individuals can be maintained exclusively on Douglas-fir needles and water (Brown, 1964). Harton (1962) has speculated that conditions in the Pa cif Northwest (extensive conifers forests with wet conditions and Imodeate temperatures) sult in a unique esvieonment that permits ‘exploitation of this feeding niche Read tree voles probably obtain water from foliage or from fos ‘or dew that collects foliage (Maser, 19654). Limited availability ff water may restrict their eastem distribution (Hamilton, 1962, Maser. 19652). rtoparesitie mites found on red tre voles inluded Andro Jnclaps Jahenholsi, A. glasgow, Glyeyphagns hypudae, Har- imogamius ambulans, He pontiges, Hirtionysaus xp, Radfordia arborimus, and a previously undescribed species of Uistephorid tite (Maser, 19652: Whitaker and Maser, 1979) The listeophorid was described as Quasliirphorus microticolas by Fain ct al (1970. Radjordia arborimus was fest described onthe basis af collection trom A. fongicaudus (Fain and Whitaker, 1973). Ticks Toure on red tee vole o in their nests include Feodes spinipalps, 1 angustus, and larval Haemaphysalis sp. A Rea (Megabothris ‘quirini) was collected from a single individual, several unidentified fheas were collected from red tree vole nest, ane an unidentified cestode war taken from aa adult fernale (Maser, 1%65a). Logging activites are implicated in declines and extinctions of some red tee vole populations (Com and Bury. 1986: Huff et al, 1992: Maser tal, 1981). Red tre voles mey have limited Alispersl capabilities andl early seral stage forets may be a barrier to dispersal ofA. ongicaudus (Carey, 1989). longicaudus may be vulnerable to loss or fragmentation of old-growth Douglas-fir forests (Half etal, 1992), “Arborimus longicaudus is dificult to eaptute. The only reli- able method of eapturing red tree voles is t0 disturb a est and Ihand-captare the animal as it les (Carey etal, 1991), There is ‘one reco of re tee vole capture in a sha tap in a tee (Wight,1 1925), Attempts to catch red te voles in Sherman live traps oF pitfall trape set in tees have been unsuccessful (Cillesberg and Carey, 1991). Pitfall taps in the ground catch red tre voles infe= quent (Com and Bury, 1986, 1968, 1991: Gilbert and Allwine, 199, BEHAVIOR. Arlorimus longicandus foes on Douglas-fir roads in steceotypical fashion. Ate biting a mature needle from {tgs thoy eplit off und discard the sides ofthe needle, including resin ducts and eat the remainder, including the medullary portion (Wenson art Borell, 1931; Hamilton, 1962: Taylor, 1915). Red tee ‘oles sometimes consume young noes entirely. including the res- in duets (Benson and Borel, 1931; Maser, 196: ‘When distred, red tree voles run ont on a limb, an down the tree headfirst, or jump to the ground Jewett, 1920; Maser, 1965). Red tee voles generally are arboreal, but occasionally ex hibit terrestrial activity, evidenced by their capture in pitfall taps (Corn ane) Bury. 1986, 1985, 1991: Gilbert and Allwine, 1901 Much of their activity takes place close to source of live needles (Maser, 19652). Red tee voles apparently are noetumal Maser fal, 1981), but no detailed studies of aetisly patter, home range Sse, or home range utilization have been published GENETICS. Arborimus longicaudus typically exhibits «dip- oid number (2a) ~ 52, with 22 pairs of acrocentric or sublelocen- luie aitesomes, three pairs of small submetacentic autosones, large submetacentsir Xy and a small metacentie Y chramosome (Hau and Benisschke, 1974: Johnson and George, 1001). OF six specimens from Oregon, one from the southernmost locality had & n= 48, because of the substitution of two large metacentie pairs {or four acrorentrie pairs (Jolson anid George, 1991) Electrophoretic’ mobiley of albumin iw A. longicaudus was faster than that of Phenacoms intermedius. In addition, the glab- tlie allele found in A. fongicaudur had a faster eleetophoretic i than that of P intermedius (Johnson, 1968) ucray (1995) analyzed restriction fragment lengths polymor- hisms from amplified portions of miDNA extracted from 12 A. pomo, 94. langicaudus, 3 Microtusealifornicus, M, ongicaudus, nl Clethrionomys gapperi, 2 C. rutidus, and A. albipes an Phenacomys intermedius. Distinet nthers and southern clades of lave voles were observed, consistent withthe hypothesis that Tongicanduc and A. pomo are distinct species. Red tree vole spec= Jens from Del Nore County, California elustered with specimens from Oregon. suggesting thatthe distibution of A. longicaudus extends into California. albips, A. ongicaudus, and 4, pomo ‘lastred together but Phenacomys intermedius clustered with MI ferotus, forming a sister group to Arborinus, Those data suggest that Avborimas isa distinct genus, and shoul not be considered tw he w subgenus of Phenacomss REMARKS. Tho taxonomy of A. longicaudus remains con troversial, At the generie level, some place Arborimus inthe genus Phenacomys (Hall, 158 Jones eal, 1992: MeAllister and Holf- rman, 1988: Verts and Carraway, 1905). Hall (1981) provisionally ‘accorded subgeneric rank to Arborimus, but Hooper and Hart (1962) argue that diferences among species of Phenacomys do not ‘even warantsubgeneric designation of Arborimas. Johnson (1968, 1973), Jabnson and Maser (1982). and Murray (1005) argue that the subgenus should be elevated to genus. I recognize the generic rank of Arborime primarily onthe basis of gence analyses (Mar fay, 1995) and supporting. moephological data, Johnson, 1973: Johnson and Maser, 1982), Similarities hetween blod proteins sug- Jost subspecific status ofA. silvicola Johnson, 1968), but no dis- tinct morpometeic or kanyaypie iiferences between the subspe- cies have heen detected in Oregon (Johnson and George, 1991). Jahnson and George (1991) and Murray (1995) provide com= pelling morphometric and genet data indiating that southern tee sole populations constitute a sibling species. Arborimas pom Pre ise init ofthe eanges ofthe eo species are not known ln several works, the origin of specimens used is unknown: as a result, some tod. pomo rather than a. longicaudus iflerences in findings among studies, such as differences in bac- ‘lum monphology reported! by Hamilton (1946) andl Dearden (1958), tay be the result of differences beiween the species. The ex Uthich A. longicandus andl A. pomo difler for several behavior ecological, and morphological characteristics is not known, A his: torical account of eaely etforts to collet red tree voles i proved by Jobanek (1988), MAMMALIAN SPECIES 532 Brian Biswell and Alison Moldenke provided helpful com- rents on an earlier draft ofthis paper Support for preparation of this paper wan provided by the Coustal Oregon Productivity En Ihancement Program, This is Paper 3000 of the Forest Research Laboratory, Oren State University, Corvallis, Oregon. LITERATURE CITED Bryson, S. BAND ALE. Bong. 1981. Notes onthe life history ofthe red tee mouse, Phenacomys longicaudus. Journal of Manmalogy, 12226-233 Brows, LN. 1964. Broeding records and notes on Phenacomys ‘lotcola in Oregon. Joural of Marmmalogy. 4:647-648. AB, 1989. Wildlife associated with old-growth in the Pacific Norwest. Natural Areas Joumal, 9:151- Camry, AB. Be LBisweis, ayo J. W. Wirt, 1991. 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