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    MAMMALIAN SPECIES no. 595, pp. 1-5. 3 fs. Arborimus pomo. By Michael D. Adam and John P. Hayes Published 4 December 1998 by the American Society of Mammalogists Arborimus Johnson, 1968 Phenacomys True, 18903308. Type locality “Marshfield [= Coos Bayh Coos Co., Oregon Arborimus Johnson, 196827. Elevation of the subgenus Arborimus to generic rank CONTENT AND CONTEXT, Order Rodentia, Suborder My- eur, Superfan Murodes, Fay Murdaey Subfamily Ar Vicoline, Tribe Mieini (Musser and Carleton, 1993) Following isa fey tothe thre extant species of Arborimus recogniged by Musser and Caslston (1950) 1 Redsh orange to cinnamon on dorsum tall thick, well Tare and ot stony bicloed: incisive foramina wide: Upper incisors strongly recurved: «pronounced long Anal depresion between ais 2 Dark brown om dorsum; tail sends, scanty ined and ‘sinelly bicolor incisive framina tron, upper ‘aos ot strongly recurved; Ingtadival depression be tween ote nat pronounced “Arbormus alipes 2 Large overall nase extend further posterioy relative o ‘the malasen lacks an abetor digit secund muscle tan Tack ful complement of lunbrieales muses tp ically exhibits a diploid number (2n) = 52, with 22 pairs a acrocentic or subtslocentie atonomes, 2 fall subcacetriceulosomen «lange sab Xv anda stall metwentne Y ehromowomes some ind ‘dal exhiit a diploid umber (2m) = 48 in which cocensc aie veplace the two lange metacente pais Gohinson and George, 199}; occuring in west forests af Oregon veces AL longicwuds Ssller overall nasis do nit extend free postesiony re ave tothe maslaris, presence of an abductor digit Secunti muscle and all complement of lumbrcales tousles; exit a diploid number 2a) = 40 or 2 th Spars of lage metaventie autosmes, 1 ple of lange Submetacene abtosomes, 1 pais of small arocente fn subtelocentie autores, 2 pir of soll submc Centric autasomes, and a small suselocenic Y cho- frosome: individuals with 42 chromosomes difler fom thie wih 40 fn hating one lee pao lage metacenrc utosomes and two aldlanal pais ofacrocentie or aub- acetic autsomes Jhon and George, 199) occu Fig in conta ors of ortwest Califia. A. pomo Arborimus pomo Johnson and George, 1991 Pomo Tree Vole Phenacomys longicaudus True, 1890:308. Type locality Marsh- field, Coos Co. Oregon. Arborimus longicaudus Tahason, 196827. Elevation of the sue ‘genus Arborimus to generic rank, Arborimus pomo Johnson and George, 1991:12. Type locality 0.8 ‘km north of Jenner, Sonoma County, California, Revision of California populations of A. longieaudus CONTENT AND CONTENT. Onler Rodentia, Suborder My- ‘omerpha, Superfamily Muroidea, Family Muridac, Sublamaly A ‘eolinae, Tribe Microtini (Musser and Careton, 1998). Musser and Carleton (1993) recognized three extant species in the genus Ar- Forimus. No subspecies of Arborimus pomo are recognized Jon- son and George, 1991). DIAGNOSIS. 4. pomo (Fig. 1) and A. Longicaudus can be Alistinguished from similar rodents by having a brownish-red dor- sum ped hing light gray venter often washed with teddish-orange, ail a well-bairee ta ht ix >50 mm Tong and 50% af length of head and body (Maser and Storm, 1970: Vert and Carrawey, 1084). A” pomo differs from A. longicaudus. by Smaller overall body size, nasals that do not extend posteriorly be- yond the maxillares, presence of the abductor digit secu mas- le and a complete set of lumbricales muscles, and differences in ‘umber and morphology of chromosomes (Johnson and George, 1991), Skulls of Arborimus ean be distinguished from thowe of other arvicoline rodents by a combination of two characters: presence of three projections on the lingual side ofthe lat upper molar; re= fentrant angles of the lower molars exter two-thirds of the wat the labial side onthe lingual side and about one-third of the way to the lingual side on the labial side (Maser and Storm, 1970) Skulls of A. pomo (Fig 2) and A. longicaudus ean be dilerentiated from A. albipes by having a pronounced longitudinal deprestic between orbits bordered by a strong ridge on each side, wide cisive foramina, and strongly recurved upper incisors (Maser and Storm, 1970) GENERAL CHARACTERS. Pelage of A. pomo is reddish brow on the dorsum and gray with rsty-browa tinged hairs onthe venter. Tail is thick, well-haised, dusky-brown above, somewhat paler helow (Johnson and George, 1991). Body mass is 20-15 Ears are siall (10-12 mm) and almost haisless. Fxtemally, pmo is almost identical to A. longicands, although A. pomo Slightly smaller in overall hdy size Johnson and George, 1991). Females are usually larger than males. Four inguinal mammary {lands are present (Howell, 1926; Johnson, 1973; Taylor, 1915) Six wellleveloped tubercles are on the hind feet and five are on the forefeet (Hooper and Hart, 1962; Taylor, 1915}, and the claw ‘on digit Ion the front foo is modified to a nail (Kesner, 1926). Minigtum and maximum mean extemal measurements (in mm) of ‘A. pom tat five geographic samples fr 56 males avd Wo Keo {raphie samples for 121 females (Johnson and George, 1991) are fs follows: length of heal and body, male 95-98, female 101-108; tail length, male 61-68, female 66-73: and hind foot, male 19-20, female 19" Lumbar transverse processes of A. pomo are large and flaring, and cervical vertebrae are short an close together Johnson, 1973) Acromian process of the seapula is narrow, sacral foramina are large, and Sterm is wide Uohnson, 1973). ‘Skull of A. pomo is heavy and moderately angular. Dental for mula is 1/1, € 00, p 00, m 33, total 16. Molar tooth are rootless in juveniles and become rooted with age (Hooper and Har, 1962; Howell, 1926). Fie 1 Adult Arborimus pomo from Humboldt County, Cali: fornia. Photograph by Lowell Diller. Smm Fic. 2. Dorel, venta, and lateral views of eranium and lateral view of mandible of Arborimus pomo from 0.5 mi S of Occidental, Sonoma Co., California (Oregon State University Department of Fisheries and Wildlife mammal collection no. 6438). Drawing by G.Bracher. Greatest Tength of the skull i= 22.5 mun, Minimuny and maximum mean values (in mm) of 4 cranial mea surements for 121 females from five geographic samples and 56 tales from two geographic samples Johnson and George, 1991) fare as follows: greatest length of skull, male 24.03-24.15, female 24.49.2492: zygomatic breadth, male 13.63-13.70, female 13.09 L.18; leat interorbital width, male 3.28-$.38, female 3.12-3.47, mastoid breadth, male 836-869, female 8.50-8.77: length of na sals, male 6.29-6.43, female 6.42-6.72: depth of brainease, male 7.08-7.19, female 7-13-7.26: widhh of itesparietl, male &14— 831, female 815-844; length of interpaietal, male 3.31-3.54, female 3528.72: length of diastem, mile 645-6.76, female 667-6383; nosal-maxilary distance, male 086-0.89, fomale 0.82 LLL: length of palatal foramen, male 4.40.61, female 447-471; length of maxillary tothiow, wale 541-543, female 5.39-64% M2-M2 width, male 5.38-5.52, female 538-5.58: and width of upper M2, male 1.22123, female 1-20-1283, DISTRIBUTION. Arborimus pomo is restricted to coniferous forests in northwestern California from the Klamath Mountains to Sonoma County, nowt of San Franeisew Fig. 3: Hull etal, 1992; MAMMALIAN SPECIES 593 Rie. a Wen). Disubution of Arborimus pom (medified from Hi Johnson and George, 1991; Meiselman and Doyle, 1996; Moray, 1995), Johnson and George (1991) hypothesized that the Klamath Mountains area geographical barier between A. pomo and A. lon sicaudus. However, May (1995) proposed that specimens fom rorthwest California (Del Norte Cousty) be assigned to A. Tong aud. Precise limits of the range of A. pomo are unknown, In particular the northern boundary of 4. pomo and tho southern Doundary of 4. tongicaualus are uncertain (Hayes, 1996), in part duc tothe limited morphologe and genetic data for individuals in {his aren, Arborimus pomo most frequently orcure in forest con sisting predominantly of Douglassir (Peeudotsug menses) that tmay include ether te species such at westem hemlock (Touga heterophylla), westem red cedar (Thuja plcata), Sitka spruce (Picea sichenss), grand fi (Abies grandis), and redwood (Sequoia sempertirens—Maser, 1965a; Meiselman and Doyle, 1996; Walker, 1930). Fossils of 4. pomo are not known FORM AND FUNCTION. Arborimus pomo is similar in mor phology to A. longicaudus. Due to the recency of the taxonomic Feelamsfcation of A. pomo (Johnaon and George, 1991), much of the current literature deseribing the morphology of red tree roles has not distinguished between A. pomo and A. longicaudus. Origin ‘of specimens teed in several studies of morphology (Dearden, 1958; Kesner, 1980, 198) were unknown or ot provided; Hayes (1996) reviewed these studies ‘The cecum of A. pomo differs from that of other microtines by the presenee of a postecal spiral consisting of 3-4 closely applied coils. Long cecal villi (up to LI mim) are present im A. pomo both in the cecum proper and in the ampulla eaealis cli. Ceeal vill: lifer fom those in the leur vill in having larger size, more elab- orate stricture, glandular inpocketings, an abeence of goblet cells, and taller epithelial cells (Voge and Bern, 1949). The cecum of Tongicaudus may have similar characteristics Arborimus pomo possesses the full complement of lambrieales ‘museles to digits three through five plus a ship to digit two (Kesner, 1986), In ation, A. pomo possesses the adductor digit secund, which is absent in AL longicaudus. Kesner (1986) provided no Tocality information for specimens examined, and assignment to A MAMMALIAN SPECIES 593 ‘pomo and A. longicaudus is hased on comments in Johnson and George (199i. ONTOGENY AND REPRODUCTION. The estrous cycle in ‘A. poms is ieregulat and variable. Duration of vaginal eomifieation of individuals Kept in captivity ranged from 1 10 21 days, wth a ‘mean of 5.9 days. Periods between comifiation ranged from 1 12 62 days. Mean proportion of time that four females had vaginal comification was 13%, ranging from 5 te 17%. Poro tree voles exhibit postpartum estas and effective postpartum mating i the wild (Hamilton, 1962) ‘Ovulation in captive induced by copulation, Prior to copulation, individuals typically exhiit a brief initial contact through a frontal approach. This is generally followed by both as- ‘suming a naso-anal position and the pal spinning ina smal ciel. Subsequently the male generally overtakes and mounts the female. While mounted, males vibrate the hindquarters ml main in this position for <4 seconds (Hamilton, tivity a male will mount a female 6-133 times consecutively permitted by the female. Time between copulations varies frm loss than one to several seconds. Fallowing eoplation the male fre= ‘quently sts and licks his genital aea (Hamilton, 19€2). Arborimus pomo sometimes exhibits delayed implantation. Gestation ranges from 27 to 48 days, averaging 31 days (Hamilton, 1962). Under laboratory conditions, females give birth duting daylight hours. Litter size is typically 2, hut vanes from I wo 3. Average Weight of 35 newboms was inversely proportional to hier size and ‘aried from 198 to 2.8 g. At hinh eyes ae covered by akin, and ye ait frm between 18h and 3 days. Prior to opening thir eves, Young receive all of their nourishment from nursing. Absolute ‘eight gain increases at about the same rate until eyes open, OF 1? animals raise in captivity, eyes opened 18-23 days afer birth, with «| mean of 20.8 days. Body weight of individuals at time of eye opening ranged from 7.5 to 10.4. After opening their eyes, agp ent lags and in wiht epey aio, ‘Amount of time spent nursing decreases from the time young begin eating slid food until they are weaned. Time of weening vanes among liters. Litters in eaplivity were weaned ax early as 25 days and as late as 46 days (Hamilton, 1962) ECOLOGY AND BEHAVIOR, Pomo tee voles may occur in higher densities in older forests than in younger forests (Huff et al. 1992; Meiselman and Doyle, 1996), bt port this hypothesis is minimal. However, microclins {nol forests, such as thermal stability nd protection from precip- itation (Franklin et al, 1981), may provide suitable habitat con- fers, although occasionally the inner bark of twigs is eaten. Dou last needles are most commonly consumed, bu other onifer spe- cies, including grand fir and Sitka spruce. are also eaten (Benson land ‘Borll, 1931). Captive Pono tree voles eat foods other than ‘eaten of conifers, but have survived in captivity only when spe plied with fresh, moist newdles of conifer (Benson and Borel, 1981; Hamilton, 1962), Free water is obtained from dew that ac ‘cumulates on needles (Taylor, 1915). Water may limit the eastern Alistbution ofA. pomo (Hamilton, 1962) andthe abundant forage ‘and moist forest eoditions of the humid coastal belt of Califoraia provide optimal habitat fr Pomo tree voles. “Arborimus pomo feed on Douglas-fi needles in a meticulous manner. A needle is bitten off at the base and held in one oF both forefet, Sides ofthe needle and resin ducts are split off and the remainder eaten (Henson and Bocll, 1931; Hamilton, 1962; Taylor, 1915). Occasionally young needles are consumed entzely (Benson and Borel, 1931). [Nests of A. pomo ae typically located in Douglas trees, sl- though grand fi, Sitka spruce, wester hemlock, redwood, and Pa cific: madeone (Arbutus mencizsd) are lao used (Meiselman and Dosle, 1996; Vriete, 1980, Zentner, 1977). Nests occur 15-35 m above ground (Taylor, 1915) and are often Teeated near the lowest branches with green leaves (Meiselman and Doyle, 1996; Views, 1980). Location of nesta varies depending on size of tee. In smaller trees, nests are typically locted near the top next tthe bole, bat in lager tees, nests ean be farther out on branches and lange tees ‘may have more than one nest (Benson and Borell, 1931; Howell 1926; Taylor, 1915). Location of a nest probably depends on lo: ion of structural supports and proximity to green leaves, Nes ‘te most frequently located on the south side of « tree Howell, 1926; Meiselman and Doyle, 1996) Pomo wee voles probably initiate construction of nests using a numberof discanied twigs and needles established in a structurally secure location. Size of nest increases with accumulation of i ‘carded resin ducts, branchlets, and fees. Nests ean reach over one meter in diameter depending on lengh of oecupation, materials ‘sed for construction, and location inthe tee. Several generations ‘of voles may use the sume nest. Exterally, nests of A pomo are somewhat spherical in shape, have a flattened top, and consist of ‘mall diameter twigs and branchlets. One or more entrance holes fare often located nea the bottom of the nest. Intermally, nests have ‘series of tunnels leading to individual chambers, which tend to he near the top of the nest even as the nest incresses in size wi ‘age (Howell, 1926). Inside, the nest is typically lined with resin ‘Facts, moss or lichen, and hits of bark. Feces oceur throughout the nes, although some chambers are more frequently used for latrines than others. Green needles, cut branchlets, resin ducts, and fresh ces typically occur on top ofan active nest. Disearded resin ducts and cut green twigs ae sometimes foun atthe hase of tees with {elive ness, Although A. pomo probably constructs mos ofits own rests, i sometimes uses nesta of other arboreal species, inluding those constructed by northern flying squimels (Glaucomys sabri= ‘nus), Douglas squires (Tamiasciuras douglasit), weslem gay Squirrels (Sciurus griseus), wootrats (Neotoma fuseipes), det Imice (Peromyscus maniculatus) or bits (Benson and. Borell, 1981; Howell, 1926, Taylor, 1913. During wet periods, feces, discarded twigs, and resin ducts in the lave pri ofthe nest may become ist ad emg Sl lesan Females occupy separate nesta, except when the female is sexually receptive. Ness of females may be larger th thore of males and subadult feales, although this patter is ine consistent (Howell, 1926; Tayler, 1915). A. pomo is primarily ar- boreal and nocturnal (Howell, 1926) Most activity ofA. pomo may be restricted to trees in which it lives, although Pome tree voles sometimes occur onthe ground (Raphael, 1988} the frequency ad function of non-arboreal ccivity are unknawn. Male A. pomo may tse subterranean neste (Howell, 1926), but this has not been con firmed since orginally reported, "Abandoned nests oflen occur in the immediate vicinity of active neste. Reatons for abaavonment are unknown, but A. longicaudus 8 Oregon deserted a nest when disturbed (Maser, 19652). Neste of Av pomo are often distributed in a clumped manner, leading so peal tat. pomg is lose coll onl, 1977), Whether clumped disteibution sdspersl abilities, parasitism, use Is or presence af favorable conditions ina specie area is unknown, Several 4. pomo individuals were infested with the mite Atr- cholaclapscalifornicus (Benson and Berell, 1981), Other ectopar ‘sites, primarily mites ticks, and fess, common an oF in nests of A. fongicaudas from Oregon (Hayes, 1996) may be associated with ‘A. pomo. Unidentified nematodes, probably in the Order Filaridea, infested one individual A. pomo (Benson and Borell, 1981, A primary predator ofA. pomo is probably the nother spotted cowl (Sirtx occidentalis caurina-—Miller, 1983), A. longicaudus comprise nearly 50% of the prey items consumed by spotted owls in Oregon (Forsman eta, 1984); spotted owls may also consume large numbers of A. pomo in Califorsia. Other birds, documented ‘5 potential predators of A longicaudas in Oregon, include great homed owls (Bubo virginianus-—Maser, 19656), long-eared owls (Asio otus—Reynolds, 1970), and northern sawewhet owls (Aego- Tis acadicus-Forsman and Maser, 1970), may als prey upon A ‘pomo in California. Mammals, including raccoons (Procyon foto), ‘martens (Martes americana), and fishers (M. pennant, probably prey on A. longicaudus (Maser, 1965a}, and tay also prey on A pore. “Arborimus pomo is difficult 1 capture, Stulies conducted in foria have had Tmited success capturing A. pomo (Ralph et al. 1991: Raphael. 1983).'The most successful method appears to be catching voles by band as a nest ie pulled apart (Carey etal, 1991) Logging a populations ofA. pomo (Benson and Borel, 1931; Hu tes may result in declines and extinctions of some etal, 1982; 4 ‘Zeotner, 1977) Pemo tee voles may have limited dispersal eapa- bilities (Towell, 1926) and may be vulnerable to bitat loss or fragmentation (ull et al, 1992), GENETICS. Arhorimus pomo has a diploid number of 40 or 42 with 5 pairs of large metacentrie autosomes, 1 pair of lange submetacentic autosomes, 11 paits of small acrocenti or subtel- of large metacentic autosomes and two additional pairs af acro- ‘entre or subtelocentric autosomes Johnson and George, 1991 Johnson and George (1991) reported Laryotypie analyses or four A longicaudus % A. pomo ybeid individuals bred in captivity: F, individuals had 6 large metacentries, 1 large submetacentre, 35. scrocentrie of subielocentic chronwcomes, and 4 small metacen- tui or submetacentrie chromosomes, ‘Murray (1995) summarized mDNA variability among seven mi- cerotine species in the genera Arborimus, Cethrionomys, Microtus, land Phenacomy, including 21 specimens of Arborimus from four counties clustered together with A. pomo, thus defining more pre- ‘ely the distribution ofthe two species (Murray, 1995) REMARKS. Taxonomy of Arborimus remains controversial Some argued that it should be included inthe geaus Phenacomys (Mall, 1981; Jones otal, 1992: Verts and Carraway, 1995). Hayes (1996) reengnized Arhorimus based on recent karyotypic, mtDNA, fad marphologie evidence olson, 1973: Johason and Maser, 1982; Murray, 1995), Sislarly, the findings of Johnson and George (1991) and Murray (1995) support the specific status of A. pom. Johnson and George (1991) hypothesized that both species are a lopatric and reproductive izlated. Reproductive success af cap tive-bred males from Oregon and females from California was sig- riicantly lower than io reciprocal crosses, suggesting 2 barier > ‘epmduetion between the two populations. Unknown linits of the Alistrbution of A. poma, along with uncemainty of locality records of specimens identified as A. ongioaudus leaves isch of the cur. ‘eat information subject to debate. "T. Lawlor provided helpfl comments on an earlier draft of this ‘paper. Support for preparation of this paper was provided by the Coastal Oregon Productivity Enhancement Progra LITERATURE CITED Benson, $B, Ant A. E, BORELL. 1931, Notes on the if history a ibe re ee rouse, Phenacomys longcaudu. Joural of Ma i306, Cay, ABB. L BRwris, aNo J. W. Wer 1991, Methods “ot measusng population of aaral rodents United States Department of Aypcutue, Foret Service, General Technical Report, PNW-G1T273, 24 pp. Deana, LC, 1958, ‘The baculum in Zagurus and related mi- ‘rte. Jounal of Mammal, 39:581-553, Fonsitey, Ey AND C, Mase ISTO, Sawn onl preys on red tre mice, Murnlet, 3110 Forsiten, BD. EC. Mrsiom, ax HM, Wicit, 1988, Die tribution and blog of the spotted owl in Oregon. Wildlife Monogrphe, 8721-64 Fravgtined Fy £7 Al 1981, Ecological characteristics of olde ‘powth Doiglafi foess, United Sates Department of Agi Eultur, Forest Servie, General Technieal Report PNW-118, 8 Haus) ET 1961, ‘The mammals of Non Ameria. Second ed. Hob Wiley & Sons, New York, 2601-1181. + 90. Hutto, Ws I 1962.” Reproductive adaptations ofthe re tee taouse. Jaa of Manmalogy, 43:180-50, Haves, JP. 1996. Asborimu longus. Mammalian Species, ‘Sia, Horta. E, axo B.S, Haye, 1962, A synopis of recent North "Ameticn mlreine dents, Miscellancon Publications ot ‘Museum of Zoology, Universi of Michigan, 2031-08 MAMMALIAN SPECIES 593 Howe, A. BL 1926, Volos of the genus Phenacomys. North American Fauna, 48:1-64. Hurry, MoH, RS. Hormacsta, aNp KB. Atuny. 1992. Hab- ‘tat management for red tree voles in Douglas-fir forest. Uni ced Slates Department of Agriculture, Forest Service, General ‘Technical Report, PNW-CTR-302:1-16. Jousor, M. L. 1968, Application af blood protein electopho- ‘etic studies to problems in mammalian taxonomy. Systematic Zoology, 1723-80, "1073. Characters ofthe heather vole, Phenacomys, and the red tee vale, Arborimas. Journal of Mammalogy, 54:239- 2a JONSON, M. Ly, AND S. B. Geowet. 1991. Species limits within the Arborimas longicaudus species-complex (Marinas: Ro-

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