5. Woodland Jumping Mouse Napaeozapus insignis French: Zapode des bois / German: \Waldhipfmaus / Spanish: Raton saltador de bosque Taxonomy. Zapus insignis G. S. Miller, 1891, Restigouche River, New Brunswick, Canada. The earliest record of Napaeozapus is from the mid-Pleistocene at Cumberland Cave, Maryland. During the furthest advance of the Wisconsin glaciation, ¢.22,000 years ago, present distribution of Napaeozapus was under ice except for areas south of northern Pennsylvania, USA. During the post-glacial period, suitable habitat for Napaeozapus occurred from Connecticut to West Virginia and Ohio, and south to northern Georgia. Napaeozapus has been found at fourlate Pleistocene sites: Bedford Co. and Bootlegger Sink, York Co., Pennsylvania; Natural Chimneys, Augusta Co., Virgina; and Robinson Cave, north-central Tennessee. The primitive zapodine stage of Megasminthus occurred by the Miocene. Most subspecies of Napaeozapus are described based on morphology alone and should probably be synonymized. Five subspecies recognized. Subspecies and Distribution. N.i.insignisG.S.Miller,1891—ECanada(fromNewBrunswick,PrinceEdwardIsland,andNovaScotia,SWthroughoutE&SQuebectoSEOntario)andSthroughNEUSAtoNE&CPennsylvania,NNewJersey,andEWestVirginia. N.i.abietorumPreble,1899—SECanada(SCOntarioandC&SWQuebec). N.i.frutectanusH.H.T.Jackson,1919—SCCanada(extremeSEManitobaandextremeSWOntario)andNCUSA(NMinnesota,NWisconsin,andNMichigan). N.i.roanensisPreble,1899—EUSA(EOhio,WPennsylvania,NE,S&WCWestVirginia,CVirginia,extremeSEKentucky,andalongtheSouthCarolina—NorthCarolinaborder,toextremeNGeorgiaandNWSouthCarolina). N. i. saguenayensis R. M. Anderson, 1942 — E & SE Canada (from Hamilton Inlet, Labrador, to the Strait of Belle Isle and SW along the N shore of the Saint Lawrence River to Lac Saint-Jean in S Quebec). Descriptive notes. Head-body 89-98 mm, tail 115-160 mm, hindfoot 28-34 mm; weight 17-26 g. Tail, hindlegs, and hindfeet of Woodland Jumping Mice are characteristically elongated. Front feet are small. It is brightly colored compared with other zapodids. Fur is light brown, with distinctive yellow or reddish tint and dark guard hairs scattered throughout. Dark brown dorsal band runs from forehead to rump. Underparts are white. Tail is very long, dark above and white on underside, and it has white tip. Females are slightly larger than males. Four pairs of teats are present: one pectoral, two abdominal, and one inguinal. Incisors are yellow or orange with deep grooves on front of upper incisors. Zygoma has jugal plates extending dorsally along maxillary ramus and articulating with lachrymal bone. Infraorbital is large and oval. Nasal bones project considerably beyond incisors. In contrast with species of Zapus, there are three molars, and premolars are absent. Molars are rooted and flat-crowned and exhibit complicated pattern of re-entrant folds, islands, and valleys. Dental formulaisI1l/1,C0/0,PM0/0,M 3/3 (x2) = 16. Habitat. Cool moist environments, almost always in wooded areas. Species of Zapus primarily occur in grasslands but can inhabit woods with adequate ground cover, particularly where Woodland Jumping Mice do not occur. Woodland Jumping Mice generally inhabit spruce-fir (Picea-Abies), hemlock (7suga, all Pinaceae), and hardwood forests of the north-eastern USA and south-eastern Canada. They also live in bogs and sphagnum swamps. At Pinkham Notch, Carroll Co., New Hampshire, they attain greatest densities in grass (Poaceae), sedge (Cyperaceae), and alder (Alnus, Betulaceae) lined borders of small streams where sand, gravel, and forest duff make it easy to construct burrows. Spruce-fir associations are common in the north and at higher elevations, whereas hemlock forests are more common in southern parts ofits distribution. Along with hemlock, northern hardwood forests are mainly of white pine (Pinus strobus, Pinaceae), beech (Fagus, Fagaceae), yellow birch (Betula allenghaniensis, Betulaceae), basswood (Tilia americana, Tiliaceae), and various species of maple (Acer, Sapindaceae). Further south, basswood, buckeye (Aesculus, Sapindaceae), tulip (Liriodendron), magnolia (Magnolia, both Magnoliaceae), and mountain laurel (Kalmia latifolia, Ericaceae) are often found. Particular species oftree is not important, but adequate ground coveris typically a determining factor. Jumping mice often occur along water, but not because of the wateritself, but usually because of adequate ground cover present in those areas. In north-western New York, Woodland Jumping Mice are most commonly found in moist dense woods, usually in conifers (spruce, fir, and hemlock) or in deciduous woods. Five of 74 individuals there were found in open situations near woods, but Woodland Jumping Mice and species of Zapus were often found together in woods. J. O. Whitaker, Jr. in 1963 used a stratified random trapping scheme via a random numbers table to study MeadowJumping Mice (Zapus hudsonius) in the Ithaca East Quadrangle, Tompkins Co., New York. Seventy-six 100 x 25 m plots were trapped, and 26 Woodland Jumping Mice were captured. All but three were captured in woods, swamp, or wet woods. Two were captured in brush, and one was taken in a field. Sixteen occurred in areas with good ground cover, only one in an area with fair cover, but six were captured in rocky wooded banks with many burrows. The individual taken in the field was ¢.40 m from woods and probably came from there, as indicated by the food in its stomach. There was no significant relationship between occurrence of mice and distance to water. Fifty-one Woodland Jumping Mice were captured elsewhere, and of those, one was in dry open woods, four were on woody stony banks, eight were in wet open woods, and the remaining 38 were in open woods with abundant ground COVET. Food and Feeding. The most abundant food (33-3% by volume) found in 103 Woodland Jumping Mice from New York was a subterranean fungus (Endogonaceae), also reported to be eaten in North Carolina, Tennessee, and New Hampshire. Spores of Elaphomyces, Hymenogaster, and Melanogaster were also found. Some have questioned if fungi might have been eaten incidentally along with other foods. That does not seem to be the case because some stomachs contained 100% fungi. Spores on the chest of some mice indicate that they probably find fungi using olfaction and then dig it up. Other important food items in New York were unidentified seeds, caterpillars, beetles, touch-me-not (Mimosa, Fabaceae), and other seeds, fruits, and invertebrates. Stomachs from 18 mice collected in the earlier part of the century by the US Biological Survey contained 33-9% by volume miscellaneous vegetation, 33-5% Endogonaceae, 10-3% strawberry, 5-3%, unidentified animal material, 5-2% beetle, and 0-8% moths. Known predators include skunks (Mephitidae), weasels (Mustela), American Mink (Neovison vison), Bobcats (Lynx rufus), domestic cat, screech owls (Megascops), timber rattlesnakes (Crotalus horridus), and copperheads (Agkistrodon contortrix). Several internal parasites have been observed in the Woodland Jumping Mouse. Protozoans of the genus Hexamita are often observed in Zapus and were found in two cecal smears examined from Woodland Jumping Mice. Cestodes (tapeworms) known to occur are Hymenolepis bennetti and Cladotaenia globifera. Nematodes (roundworms) include Rictularia sp. and Citellinoides zapodis. Mites, other than chiggers, include Glycyphagidae: Glycyphagus hypudaer, G. newyorkensis, and G. zapus; Laelapidae: Androlaelaps fahrenholz, Echinonyssus isabellinus, Eulaelaps stabularis, Haemogamasus alaskensis, Laelalps kochi, and Listrophorus mexicanus; Macronyssidae: Ornithonyssus bacoti; Myobiidae: Protomyobia brevisetosa and Radfordia ewingi; Myocoptidae: Gliricoptes glirinus, and Pygmephoridae: Pygmephorus horridus and P. mahunkai. Chiggers that have been found include Euschoengastia diversa, E. peromysci, E. rubra, E. setosa, Eutrombicula alfreddugest, Leptotrombidum peromysci, Miyatrombicula esoensis, Neotrombicula harper, N. microti, N. richmondi, and N. whartona. Ticks include Dermacentor variabilis and Ixodes muris. Botflies include Cuterebra sp. and C. fontinella. Fleas include Ctenocephalides pseudagyrtes, Corrodopsylla curvata, Epitedia sp., Megabothris asio, M. quirini, Orchopeas leucopus, Peromyscopsylla catatina, and Stenoponia americana. One individual louse, Neohaematopinus sp., has been reported. Breeding. Woodland Jumping Mice breed in May-August. Females mate and have young soon after emergence from hibernation in late May and June. Earliest breeding records found in the central part of the distribution were 8-9 May, with embryos only 1-2 mm long. Most births occur in June because more females are pregnant or show signs of having young than at any other time. Nevertheless, not all females reproduce in May-June, and most that do not are probably individuals born late in the previous year. Reproduction occurs at a low rate during a second breeding peak in July. Many females captured in August show evidence of already having had one litter. It is clear that many females produce two litters per year, but three litters in one year would be unusual. Very few,if any, bear young in the year of their birth. Eighty litters examined by R. E. Wrigley in 1972 averaged 4-6 embryos (range 2-7), 45 placental scar counts averaged 4-3 (range 2-7), and litter size was 1-8 young. Gestation is c.18 days but is 2-3 days longer if a female is lactating when mating occurs. There is only one litter per year in Nova Scotia. Early dates on which firstjuveniles were caught in Nova Scotia were 27 July, 6 August, 16 August, and 23 August. In a five-year study carried out by P. F. Connor in 1966 in north-western New York, the earliest reproductive female was captured on 26 May, but she had already had one litter. At Pinkham Notch, New Hampshire, breeding season extended from late May to August. Woodland Jumping Mice nest in underground burrows or brush, or underfallen logs. Young are born naked and hairless except for vibrissae. Four pairs of mammary glands are visible in both sexes. At ten days of age, dorsal pigmentation appears, and ear pinnae unfold, although external auditory meatus remains closed. Fine hair appears on dorsum of head, body, and limbs at twelve days. At this time, vibrissae are ¢.6 mm, and young are able to stand up and take a few steps. White hair appears on venter at 14 days old. At 19 days, dorsal pelage is well developed, except dorsal band is not yet evident. Lower incisors protrude 0-2-0-3 mm, and claws are well developed. Dark dorsal band is well formed at 24 days. Eyes and external auditory meatus open at 26 days, and young begin moving by short hops. At 28 days, lower incisors protrude 2 mm, and upper incisors protrude 1 mm. At this stage, young are able to hop lengths of more than 30 cm and are nearly mature and independent. Activity patterns. Woodland Jumping Mice are usually nocturnal but may be active in late morning and evening, especially on cloudy or rainy days. They typically walk or crawl through ground cover when moving slowly but hop when greater speed is desired. When startled, they are capable ofjumping at least 2 m at heights of nearly 1 m. After several longer leaps, they will usually stop and hide under cover and remain motionless unless pursued. Woodland Jumping Mice can climb bushes, probably to get berries, but there is no evidence that they climb trees. They can swim probably even below the surface. They are much less inclined to form definite runways than many other species of small mammals, but they will use runways of other small rodents and larger mammals such as North American Beavers (Castor canadensis), deer (Odocoileus), and Moose (Alces alces). They will dig their own burrows, although they will use those of other mammals. Nests of leaves or leaves and grass have been located in brush piles. Young are “vociferous,” and adults produce soft chucking sounds. They use tails to drum. Associates are chipmunks (7amias), White-footed Deermice (Peromyscus leucopus), Cinereus Shrews (Sorex cinereus), Smoky Shrews (Sorex fumeus), Northern Short-tailed Shrews (Blarina brevicauda), and Southern Red-backed Voles (Myodes gapperr). North American Red Squirrels (7 Tamiasciurus hudsonicus), Meadow Jumping Mice, House Mice (Mus musculus), Meadow Voles (Microtus pennsylvanicus), Hairy-tailed Moles (Parascalops breweri), Star-nosed Moles (Condylura cristata), Woodland Voles (Microtus pinetorum), Southern Bog Lemmings (Synaptomys cooperi), and North American Deermice (Peromyscus maniculatus) are less common associates. In one study, survival of adults was 16-1-41-2% during two years. Sixteen percent of males and 28:6% of females that were first captured as juveniles survived until their third active season, but none survived until their fourth active season. Ten males and ten females caught as adults survived another season (three seasons total), but three males and three females were caught in three more seasons and survived four consecutive seasons. One female was caught as an adult in 1980 and again in 1984 (but not between), surviving for at least five seasons. Males emerged 15-33 days ahead of females. In preparation for hibernation, Woodland Jumping Mice accumulate large amounts offat, often one-third of their body weights. Sufficient energy from fat must be available to maintain mice for at least six months. Weight gain occurs over c.2 weeks for any individual but probably over c.6 weeks beginning in late August and early September for the population. There is some evidence that photoperiod is at least partly a cue for hibernation. Latest time in which Woodland Jumping Mice are active in central New York is late October. Most of the population present this late in the season is composed of younger individuals that will hibernate after they have put on adequate fat. It is likely that those not adding adequate fat die before spring emergence. During hibernation, an individual curls into a ball with its nose in its abdomen, tail curled around its body, forelegs along its chest, and hindlegs along its face. Respiration rate of hibernating individuals is c.12 breaths/minute; when sleeping,it is ¢.200 breaths/ minute. Emergence in spring takes place mostly from mid-April through mid-May; males emerge before females. Dates of emergence varied from 4 to 13 May for males and 28 May to 7 June for females. Dates of disappearance, presumably to hibernate were about the same for males and females; however, juveniles of both sexes tended to disappear earlier than adults. Sex ratios were 44 males to 62 females (1:1-4) for adults but 54 males to 45 females (1:0-83) for juveniles. Movements, Home range and Social organization. Home ranges of Woodland Jumping Mice are 0-4-2-6 ha for females and 0-4-3-6 ha for males, and they broadly overlap. W. H. Burt in 1954 calculated home ranges of 0-4-0-81 ha. Populations of 7-4-59-3 ind/ha have been reported, but the latter is probably too high. Woodland Jumping Mice were c.21% of the small mammal community in central New York. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Woodland Jumping Mouse is widely distributed, presumably has large populations, and occurs in a number of protected areas. Population trends are stable, and it is unlikely to be declining fast enough to qualify forlisting in a more threatened category. Bibliography. Abbott & Parsons (1961), Benton & Krug (1956), Blair (1941), Brennan & Wharton (1950), Brower & Cade (1966), Burt (1954), Collins & Cameron (1984), Connor (1960, 1966), Dikmans (1939), Dodds et al. (1969), Fain & Whitaker (1973), Fain et al. (1985), Farner (1946), Farrell (1956), Freeman (1959, 1960), Geary (1959), Gidley & Gazin (1938), Goodwin (1924, 1929, 1935), Greenfield (1938), Guilday (1962), Guilday, Hamilton & McCrady (1966), Guilday, Martin & McCrady (1964), Hamilton (1941), Harper (1929), Holland (1949), Holland & Benton (1968), Jameson (1949), Jones & Thomas (1982), Jordan (1929), Layne & Hamilton (1954), Linzey & Linzey (1968), Manville (1949), McCrady & Schmidt (1963), Morris (1948), Neumann & Cade (1964), Orrock et al. (2003), Ovaska & Herman (1988), Platte (1968), Preble (1956), Priddy (1949), Reeves, Durden et al. (2007), Sheldon (1934, 1938), Siegmund (1964), Snyder (1924), Strandtmann (1949), Whitaker (1962, 1963a), Whitaker & French (1982), Whitaker et al. (1975), Wilson & Reeder (2005), Wrenn (1974), Wright (1979), Wrigley (1972)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Zapodidae, pp. 50-61 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 50-61, DOI: 10.5281/zenodo.6609503