Whittell Forest and Wildlife Area

Whittell Forest and Wildlife Area

A living lab of forested mountain land in the Sierra Nevada, dedicated for research, instruction and community outreach.

About the Whittell Forest and Wildlife Area

The Whittell Forest and Wildlife Area was gifted to the University in 1959 by George Whittell, with the intent of maintaining a living lab of forested mountain land in a primitive state for research, instruction and community outreach.

A scenic view of a forested area of Whittell Forest and Wildlife Area with granite rocks and trees

The 2,650 acre property is located in the Carson Range on the east slope of the Sierra Nevada about 30 kilometers south of Reno, Nevada. It is five kilometers west of Washoe Lake and four kilometers northeast of Lake Tahoe. Most of the property is in Little Valley at an elevation of 1,900-2,500 meters. Vegetation is dominated by Jeffrey pine forests with an understory of bitterbrush and manzanita, lodgepole pine forests and subalpine meadows.

The climate is semi-arid with a long summer dry season, with most precipitation falling during the winter in the form of snow. Franktown Creek drains the valley into Washoe Lake to the east. Access is through a locked gate and via a steep dirt road, accessible by four-wheel drive vehicles from May to November most years, and by foot, skis or snowshoes during the remainder of the year.

In order to maintain its primitive nature, and thereby further the University’s research and teaching efforts, the operation of private recreational vehicles within the forest is prohibited.

Forest use and management

Sarah Bisbing, assistant professor in the College of Agriculture, Biotechnology and Natural Resources’ Department of Natural Resources and Environmental Science, serves as the Director of the Whittell Forest and Wildlife Area, a part-time role that administratively reports to Research & Innovation. Bisbing has in-depth knowledge of forest ecology, forest management, fire ecology, and related sciences to support forestry activities, field research and a learn-by-doing approach to education. She also has extensive experience with cultural and regulatory requirements relative to field stations and remote sites.

The Whittell Forest Advisory Committee, chaired by Bisbing, assists in the establishment of procedures and practices to make the forest available as a core academic and research facility. The advisory committee currently includes members from the community, the Nevada Department of Forestry, the USDA and the University of Nevada, Reno.

Bisbing envisions the land being utilized by a broad range of disciplines across campus and throughout the Nevada System of Higher Education for experiential learning opportunities, research and creative activities. The unique mountain system offers the region and the State of Nevada a vital habitat to study natural resources, contribute to environmentally based education and perpetuate the vision of protecting land.

Journal publications related to the forest

  • Publications relating to bark beetles

    Gary Blomquist and Claus Tittiger labs

    • S.J. Seybold, D.R. Quilici, J.A. Tillman, D. Vanderwel, D.L. Wood and G.J. Blomquist. De novo biosynthesis of the aggregation pheromone components ipsenol and ipsdienol by the pine bark beetle, Ips paraconfusus Lanier and Ips pini (Say)(Coleoptera: Scolytidae). Proc. Natl. Acad. Sci. 92:8393-8397 (1995).
    • P. Ivarsson, G.J. Blomquist, S.J. Seybold. In vitro production of pheromone intermediates in the bark beetles Ips pini (Say) and I. paraconfusus Lanier (Coleoptera:Scolytidae). Naturwissenschaften 84:454-457 (1997).
    • P. Ivarsson, C. Tittiger, C. Blomquist, C.E. Borgeson, S.J. Seybold, G.J. Blomquist and H.-E. Hogberg. Pheromone precursor synthesis is localized in the metathorax of Ips paraconfusus Lanier (Coleoptera: Scolytidae). Naturwissenschaften 85:507-511(1998).
    • J.A. Tillman, G.L. Holbrook, P. Dallara, C. Schal, D.L. Wood, G.J. Blomquist, and S.J. Seybold. Endocrine regulation of de novo aggregation pheromone biosynthesis in the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae). Insect Biochem. Molec. Biol. 28:705-715 (1998).
    • C. Tittiger, G.J. Blomquist, P. Ivarsson, C.E. Borgeson and S.J. Seybold. Juvenile hormone regulation of HMG-R gene expression in the bark beetle Ips paraconfusus (Coleoptera: Scolytidae): implications for male aggregation pheromone biosynthesis. Cell. Mol. Life Sci. 55:121-127 (1999).
    • C. Tittiger, C. O’Keeffe, C.S. Bengoa, L. Barkawi, S.J. Seybold and G.J. Blomquist. Isolation and endocrine regulation of HMG-CoA synthase cDNA from the male Jeffrey pine beetle, Dendroctonus jeffreyi. Insect Biochem. Molec. Biol. 30:1203-1211 (2000).
    • G. M. Hall, C. Tittiger, G. Andrews, G. Mastick, M. Kuenzli, X. Luo, S.J. Seybold and G.J. Blomquist. Male pine engraver Beetles, Ips pini, synthesize the monoterpenoid pheromone ipsdienol de novo in midgut tissue. Naturwissenschaften 89:79-83 (2002).
    • G.M. Hall, C. Tittiger, G.J. Blomquist, G. Andrews, G. Mastick, L.A. Barkawi, Cody S. Bengoa, and S. J. Seybold. Male Jeffrey Pine Beetles, Dendroctonus jeffreyi, synthesize the pheromone component frontalin in anterior midgut tissue. Insect Biochem. Mol. Biol. 32:1525-1532 (2002).
    • J.B. Nardi, A. Gilg Young, E. Ujhelyi, C. Tittiger, M.J. Lehane and G.J. Blomquist.Specialization of midgut cells for synthesis of male isoprenoid pheromone in two scolytid beetles, Dendroctonus jeffreyi and Ips pini. Tissue and Cell. 226:221-231 (2002).
    • C. Tittiger, C. Bengoa, L. Barkawi, G.J. Blomquist and S.J. Seybold, Structure and juvenile hormonemediated regulation of the HMG-CoA reductase gene from the Jeffrey pine beetle, Dendroctonus jeffreyi. Molecular and Cellular Endocrinology 199:11-21 (2003).
    • L.S. Barkawi, W. Francke, G.J. Blomquist and S.J. Seybold. Frontalin: De novo biosynthesis of an aggregation pheromone component by Dendroctonus spp. bark beetles (Coleoptera: Scolytidae). Insect Biochem. Molec. Biol. 33:773-788 (2003).
    • C.I. Keeling, G.J. Blomquist and C. Tittiger. Coordinated mevalonate gene expression upon feeding in the anterior midgut of the pine engraver beetle, Ips pini (Say) (Coleoptera: Scolytidae). Naturwissenschaften 91:324-329 (2004).
    • J.A. Tillman, F. Lu, L.M. Goodard, Z. Donaldson, S.C. Dwinell, C. Tittiger, G.M. Hall, A. J. torer, G.J. Blomquist and S.J. Seybold. Juvenile hormone regulates de novo aggregation pheromone biosynthesis in the pine bark beetles Ips spp. (Coleoptera: Scolytidae) through transcriptional control of HMG-CoA reductase. J. Chem. Ecol. 30:2459-2494 (2004).
    • A.B. Gilg, J.C. Bearfield, C. Tittiger, W.H. Welch and G.J. Blomquist. Isolation and functional expression of an animal geranyl diphosphate synthase and its role in bark beetle pheromone biosynthesis. Proc. Natl. Acad. Sci. 102:9760-9765 (2005).
    • C.I. Keeling, S. Young, G.J. Blomquist and C. Tittiger. Juvenile hormone-regulated gene expression in the pheromone-producing midgut of the pine engraver beetle, Ips pini. Insect Mol. Biol. 15:207- 216 (2006).
  • Publications on soils and nutrient cycling from Little Valley

    Dale Johnson and Wally Miller labs

    • Burcar, S., W.W. Miller, S.W. Tyler, and R.R. Blank. 1997. Moist- and dry-season nitrogen transport in Sierra Nevada soils. Soil Sci. Soc. Amer. J. 61: 1774-1780.
    • Burcar, S., W.W. Miller, S.W. Tyler, and D.W. Johnson. 1994. Seasonal preferential flow in two Sierra Nevada soils under forested and meadow cover. Soil Sci. Soc. Amer. J. 58: 1555-1561.
    • Johnson, D.W. 1995. Soil properties beneath Ceanothus and pine stand in the eastern Sierra Nevada. Soil Sci. Soc. Amer. J. 59: 918-924.
    • Johnson, D.W., W. Cheng, and I.C. Burke. 2000. Biotic and Abiotic Nitrogen Retention in a Variety of Forest Soils. Soil Sci. Soc. Amer. J. 64: 1503-1514.
    • Johnson, D. W., Glass, D. W., Murphy, J. D., Stein, C. M., Miller, W. W. 2010. Hot Spots and Hot Moments: Another Look at Nutrient Variability in Sierra Nevada Forest Soils. Biogeochemistry, 101, 93-103.
    • Johnson, D. W., W.W. Miller, R.B. Susfalk, R.A. Dahlgren, J.D. Murphy, and D.W. Glass. 2009. Biogeochemical Cycling in Forest Soils of the Eastern Sierra Nevada Mountains, USA. Forest Ecology and Management 258: 2249-2260.
    • Johnson, D. W., J.D. Murphy, R.F. Walker, D. Glass, and W.W. Miller. 2007. Wildfire Effects on Forest Carbon and Nutrient Budgets.. Ecological Engineering. 31: 183-192.
    • Johnson, D.W., R. B. Susfalk, and R.A. Dahlgren. 1997. Nutrient fluxes in forests of the eastern Sierra Nevada Mountains, USA. Global Biogeochem. Cycles 11: 673-681.
    • Johnson, D.W., R.B. Susfalk, R.A. Dahlgren, T.G. Caldwell and W.W. Miller. 2001. Nutrient fluxes in a snow-dominated, semi-arid forest: Spatial and temporal patterns. Biogeochemistry 55:219-245.
    • Johnson, D.W., R.B. Susfalk, R.A. Dahlgren, and J.M. Klopatek. 1998. Fire is more important than water for nitrogen fluxes in semi-arid forests. Environ. Sci. Pol. 1: 79-86.
    • Johnson, D.W., R.B. Susfalk, H.L. Gholz, and P.J. Hanson. 2001. Simulated effects of temperature and precipitation change in several forest ecosystems. J. Hydrology 235: 183-204.
    • Johnson, D.W., R.B. Susfalk, T.G. Caldwell, J.R. Murphy, W.W. Miller, and R.F. Walker. 2004. Fire Effects on Carbon and Nitrogen Budgets in Forests. Water, Air, and Soil Pollut. Focus 4: 263- 275.
    • Johnson, D.W., J.D. Murphy, R.B. Susfalk, T.G. Caldwell, W.W. Miller, R.F. Walker, and R.F. Powers. 2005. The effects of wildfire, salvage logging, and post-fire N fixation on the nutrient budgets of a Sierran forest. For. Ecol. Manag. 220: 155-165.
    • Miller, W. W., Johnson, D. W., Karam, S. L., Walker, R. F., Weisberg, P. J. 2010. A synthesis of Sierran forest biomass management studies and potential effects on water quality. Forests, 1, 131-153.
    • Stark, N.M., 1973. Nutrient cycling in a jeffrey pine ecosystem. University of Montana, Missoula, MT Stein, C. M., Johnson, D. W., Miller, W. W., Powers, R. F., Young, D. A., Glass, D. S. 2010. Snowbrush (Ceanothus velutinus Dougl) Effects on Soil Leaching and Water Quality in a Sierran Ecosystem. Ecohydrology, 3, 79-87.
    • Susfalk, R.B., and D.W. Johnson. 2002. Ion exchange resin based soil solution lysimeters and snowmelt collectors. Comm. Soil Sci. Plant Anal.33: 1261-1275.
    • Johnson, D.W., R.B. Susfalk, R.A. Dahlgren, T.G. Caldwell and W.W. Miller. 2001. Nutrient fluxes in a snow-dominated, semi-arid forest: Spatial and temporal patterns. Biogeochemistry 55:219-245.
    • Susfalk, R.B., and D.W. Johnson. 2002. Ion exchange resin based soil solution lysimeters and snowmelt collectors. Comm. Soil Sci. Plant Anal.33: 1261-1275.
    • Johnson, B.G., D.W. Johnson, W.W. Miller, E.M. Carroll-Moore, and D.I. Board. 2011. The effects of slash pile burning on soil and water macronutrients. Soil Science 176: 413-425.
    • Johnson, B.G., D.W. Johnson, W.W. Miller, and D.I. Board. 2012. The effects of ash influx on burned and unburned soil water-extractable nutrients using a mechanical vacuum extractor. Soil Sci, 177: 338-344).
  • Publications on rodent foraging and seed dispersal

    Stephen Vander Wall lab

    • Vander Wall, S. B. 1991. Mechanisms of cache recovery by yellow pine chipmunks. Animal Behaviour 41:851-863.
    • Vander Wall, S. B. 1992. The role of animals in dispersing a "wind-dispersed" pines. Ecology 73:614- 621.
    • Vander Wall, S. B. 1992. Establishment of Jeffrey pine seedlings from animal caches. Western Journal of Applied Forestry 7:14-20.
    • Vander Wall, S. B. 1993. A model of caching depth: Implications for scatter hoarders and plant dispersal. American Naturalist 141:217-232.
    • Vander Wall, S. B. 1993. Seed water content and the vulnerability of buried seeds to foraging rodents. American Midland Naturalist 129:272-281.
    • Vander Wall, S. B. 1993. Salivary water loss to seeds by yellow pine chipmunks and Merriam's kangaroo rats. Ecology 74:1307-1312.
    • Vander Wall, S. B. 1993. Cache site selection by chipmunks (Tamias spp.) and its influence on the effectiveness of seed dispersal in Jeffrey pine (Pinus jeffreyi). Oecologia 96:246-252.
    • Vander Wall, S. B. 1994. Removal of wind-dispersed pine seeds by ground-foraging vertebrates. Oikos 69:125-132.
    • Vander Wall, S. B. 1994. Seed fate pathways of antelope bitterbrush: Dispersal by seed-caching yellow pine chipmunks. Ecology 75:1911-1926.
    • Vander Wall, S. B. 1995. Influence of substrate water on the ability of rodents to find buried seeds. Journal of Mammalogy 76:851-856.
    • Vander Wall, S. B. 1995. Salivary water loss during seed husking in deer mice and Great Basin pocket mice. Physiological Zoology 68:878-886.
    • Vander Wall, S. B. 1995. Dynamics of yellow pine chipmunk (Tamias amoenus) seed caches: Underground traffic in bitterbrush seeds. Ecoscience 2:261-266.
    • Vander Wall, S. B. 1995. Sequential patterns of scatter hoarding in yellow pine chipmunks. American Midland Naturalist 133:312-321.
    • Vander Wall, S. B. 1995. The effects of seed value on the caching behavior of yellow pine chipmunks. Oikos 74:533-537.
    • Vander Wall, S. B., and E. Peterson. 1996. Associative learning and the use of cache markers by yellow pine chipmunks (Tamias amoenus). Southwestern Naturalist 41:88-90.
    • Vander Wall, S. B. 1998. Foraging success of granivorous rodents: effects of variation in seed and soil water content on olfaction. Ecology 79:233-241.
    • Vander Wall, S. B., and J. W. Joyner. 1998. Recaching of Jeffrey pine (Pinus jeffreyi) seeds by yellow pine chipmunks (Tamias amoenus): potential effects on plant reproductive success. Canadian Journal of Zoology 76:154-162.
    • Vander Wall, S. B., and J. W. Joyner. 1998. Secondary wind dispersal of winged pine seeds. American Midland Naturalist 139:365-373.
    • Vander Wall, S. B., and W. Longland. 1998. Cheek pouch capacities and loading rates of deer mice (Peromyscus maniculatus). Great Basin Naturalist 59:278-280.
    • Vander Wall, S. B. 2000. The influence of environmental conditions on cache recovery and cache pilferage by yellow pine chipmunks (Tamias amoenus) and deer mice (Peromyscus maniculatus). Behavioral Ecology 11:544-549.
    • Vander Wall, S. B., T. C. Thayer, J. S. Hodge, M. J. Beck, J. K. Roth. 2001. Scatter-hoarding behavior of deer mice (Peromyscus maniculatus). Western North American Naturalist 61:109-113.
    • Vander Wall, S. B. 2002. Secondary dispersal of Jeffrey pine seeds by rodent scatter hoarders: the roles of pilfering, recaching, and a variable environment. Pages 193-208 In Levey, D. J., Silva, W. R., and M. Galetti. Seed Dispersal and Frugivory: Ecology, Evolution and Conservation. CABI Publishing, Wallingford, Oxfordshire, UK.
    • Vander Wall, S. B. 2002. Masting in pines alters the use of cached seeds by rodents and causes increased seed survival. Ecology 83:3508-3516.
    • Vander Wall, S. B. 2003. Effect of seed size of wind-dispersed pines (Pinus) on secondary seed dispersal and the caching behavior of rodents. Oikos 82:25-34.
    • Vander Wall, S. B., M. J. Beck, J. S. Briggs, J. K. Roth, T. C. Thayer, J. L. Hollander, and J. M. Armstrong. 2003. Interspecific variation in the olfactory abilities of granivorous rodents. Journal of Mammalogy 84:159-168.
    • Vander Wall, S. B. 2003. How rodents smell buried seeds: a model based on the behavior of pesticides in soil. Journal of Mammalogy. 84:1089-1099.
    • Vander Wall, S. B., and S. H. Jenkins. 2003. Reciprocal pilfering and the evolution of food-hoarding behavior. Behavioral Ecology 14:656-667.
    • Briggs, J. S., and S. B. Vander Wall. 2004. Substrate type affects caching and pilferage of pine seeds by chipmunks. Behavioral Ecology 15:666-672.
    • Thayer, T. C., and S. B. Vander Wall. 2005. Interactions between Steller’s jays and yellow pine chipmunk over scatter-hoarded sugar pine seeds. Journal of Animal Ecology 74:365-374.
    • Vander Wall, S. B., K. M. Kuhn, and M. J. Beck. 2005. Seed removal, seed predation, and secondary dispersal. Ecology 86:801-806.
    • Vander Wall, S. B., E. C. H. Hager, and K. M. Kuhn. 2005. Pilfering of stored seeds and the relative costs of scatter hoarding versus larder hoarding in yellow pine chipmunks. Western North American Naturalist 65:248-257.
    • Roth, J. K., and S. B. Vander Wall. 2005. Dispersal and establishment of Sierra chinquapin (Castanopsis sempervirens) by seed-caching rodents. Ecology 86:2428-2439.
    • Vander Wall, S. B., K. M. Kuhn, and J. Gworek. 2005. Two phase seed dispersal: linking the effects of frugivorous birds and seed-caching rodents. Oecologia 145:282-287.
    • Vander Wall, S. B. (with contributions from 14 colleagues). 2005. Whittell Forest Fuel Reduction and Ecosystem Enhancement Plan. Approved and implemented by the Whittell Board of Control. 86p.
    • Vander Wall, S. B., J. S. Briggs, S. H. Jenkins, K. M. Kuhn, T. C. Thayer and M. J. Beck. 2006. Do food-hoarding animals have a recovery advantage? determining recovery of stored food. Animal Behaviour 72:189-197.
    • Gworek, J. R., S. B. Vander Wall, and P. F. Brussard. 2007. Changes in biotic interactions and climate determine recruitment of Jeffrey pine along an elevation gradient. Forest Ecology and Management 239:57-68.
    • Richardson, T. W., and S. B. Vander Wall. 2007. Yellow pine chipmunks cannot climb quaking aspens: implications for avian nest site selection. Western North American Naturalist 67:251-257.
    • Kuhn, K. M., and S. B. Vander Wall. 2007. Black bears (Ursus americanus) harvest Jeffrey pine (Pinus jeffreyi) seeds from tree canopies. Western North American Naturalist 67:384-388.
    • Vander Wall, S. B. 2008. On the relative contributions of wind versus animals to seed dispersal of four Sierra Nevada pines. Ecology 89:1837-1849.
    • Vander Wall, S. B., C. J. Downs, M. S. Enders, and B. A. Waitman. 2008. Do yellow-pine chipmunks prefer to recover their own caches? Western North American Naturalist 68:319-325.
    • Kuhn, K. M., and S. B. Vander Wall. 2008. Linking summer foraging to winter survival in yellow pine chipmunks (Tamias amoenus). Oecologia 157:349-360.
    • Briggs, J. S., S. B. Vander Wall, and S. H. Jenkins. 2009. Forest rodents provide directed dispersal of Jeffrey pine seeds. Ecology 90:675-687.
    • Vander Wall, S. B., M. S. Enders and B. A. Waitman. 2009. Asymmetrical cache pilfering between yellow pine chipmunks and golden-mantled ground squirrels. Animal Behaviour 78:555-561.
    • Kuhn, K. M., and S. B. Vander Wall. 2009. Formation and contents of yellow-pine chipmunk (Tamias amoenus) winter larders. Western North American Naturalist 69:309-318.
    • Downs, C. J., and S. B. Vander Wall. 2009. High relative humidity increases pilfering success of yellow pine chipmunks. Journal of Mammalogy 90:796-802.
    • Legras, E. C., S. B. Vander Wall and D. I. Board. 2010. The role of germination microsite in the establishment of sugar pine and Jeffrey pine seedlings. Forest Ecology and Management 260:806-813.
    • Vander Wall, S. B. and S. H. Jenkins. 2011. Plant-animal interactions and climate: why do yellow pine chipmunks (Tamias amoenus) and eastern chipmunks (Tamias striatus) have such different effects on plants? Ecoscience 18:130-137.
    • Vander Wall, S. B. and Beck, M. J. 2012. Frugivory versus food-hoarding dispersal syndromes. Botanical Review 78:10-31.
    • Enders, M. S. and S. B. Vander Wall. 2012. Black bears are effective dispersers of seeds, with a little help from their friends. Oikos 121:589-596.
    • Hollander, J. L., S. B. Vander Wall, and W. S. Longland. 2012. A comparison of the detectability of wildland and cultivated seeds by foraging rodents. Western North American Naturalist 73:339-347.
    • Barga, S. C. and S. B. Vander Wall. 2013. Dispersal of an herbaceous perennial, Paeonia brownii, by scatter-hoarding rodent. Ecoscience 20:172-181.
    • Moore, C. M., and S. B. Vander Wall. 2015. Scatter-hoarding rodents disperse seeds to safe sites in a fire-prone ecosystem. Plant Ecology 216:1137-1153.
    • Vander Wall, S. B., and C. M. Moore. 2016. Interaction diversity of North American seed-dispersal mutualisms. Global Ecology and Biogeography 25:1377-1386.
    • Vander Wall, S. C. Barga, and A. E. Seaman. 2017. The geographic distribution of seed-dispersal mutualisms in North America. Evolutionary Ecology 31:725-740.
    • Dittel, J. W., R. Perea, and S. B. Vander Wall. 2017. Pilfering dynamics in a seed-caching rodent community. Behavioral Ecology and Sociobiology 71:1-8.
    • Dittel, J. W. and S. B. Vander Wall. 2018. The effects of rodent abundance and richness on cache pilfering. Integrative Zoology 13:328-335.
    • Dittel, J. W., C. M. Moore, and S. B. Vander Wall. 2018. The mismatch in distributions of vertebrates and the plants that they disperse. Ecography 42:565-575.
    • Vander Wall, S. B. Seed harvest by a scatter-hoarding rodent. Journal of Mammalogy in press
  • Research on beavers and other rodents

    Stephen Jenkins lab

    • Busher, P. E., R. J. Warner, and S. H. Jenkins. 1983. Population density, colony composition, and local movements in two Sierra Nevada beaver populations. Journal of Mammalogy 64:314-318.
    • Mewaldt, W. T., and S. H. Jenkins. 1986. Genetic variation of woodrats (Neotoma cinerea) and deer mice (Peromyscus maniculatus) on montane habitat islands in the Great Basin. Great Basin Naturalist 46:577-580.
    • Basey, J. M., S. H. Jenkins, and P. E. Busher. 1988. Optimal central-place foraging by beavers: tree-size selection in relation to defensive chemicals of quaking aspen. Oecologia 76:278-282.
    • Eshelman, B. D., and S. H. Jenkins. 1989. Food selection by Belding's ground squirrels in relation to plant nutritional features. Journal of Mammalogy 70:846-852.
    • Jenkins, S. H., and P. W. Bollinger. 1989. An experimental test of diet selection by the pocket gopher Thomomys monticola. Journal of Mammalogy 70:406-412.
    • Basey, J. M., S. H. Jenkins, and G. C. Miller. 1990. Food selection by beavers in relation to inducible defenses of Populus tremuloides. Oikos 59:57-62.
    • Duncan, R. D., and S. H. Jenkins. 1998. Use of visual cues in foraging by a diurnal herbivore, the Belding’s ground squirrel. Canadian Journal of Zoology 76:1766-1770.
  • Miscellaneous publications
    • Stark, N. M., 1973. Nutrient cycling in a jeffrey pine ecosystem. MS Thesis. University of Montana, Missoula, MT.
    • Vreeland, P. and H. Vreeland. 1975. Fire and ice: the brilliant snow plant lights the forest as winter’s drifts recede. Mentzelia 1:14-16.
    • Vreeland, P., H. Vreeland, and E. F. Kleiner. 1976. Sarcodes sanguinea Torr., a mycorrhizal species. American Midland Naturalist 96:507-512.
    • Vreeland, P., E. F. Kleiner, and H. Vreeland. 1981. Mycorrhizal symbiosis of Sarcodes sanguinea. Environmental and Experimental Botany 21:15-25.
    • Goodrich, J. and J. Berger, 1994. Winter recreation and hibernating black bears. Biological Conservation 67:105-110.
    • Hefty, K. L., and K. M. Stewart. 2018. Novel location data reveal spatiotemporal strategies used by a central-place forager. Journal of Mammalogy 99:333-340.
    • Hefty, K. L., and K. M. Stewart. 2019. Flexible resource use strategies of a central-place forager