Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Do measures of plant intake and digestibility from captive feeding trials align with foraging patterns of free-ranging snowshoe hares?

Ethan Ellsworth A B , Aaron J. Wirsing C F , Lisa A. Shipley D and Dennis L. Murray E
+ Author Affiliations
- Author Affiliations

A Department of Fish and Wildlife Resources, University of Idaho, Moscow, ID 83843, USA.

B Present address: Bureau of Land Management, Mount Lewis Field Office, Battle Mountain, NV 89820, USA.

C School of Environmental and Forest Sciences, Box 352100, University of Washington, Seattle, WA 98195-2100, USA.

D School of the Environment, Washington State University, Pullman, WA 99164-6410, USA.

E Department of Biology, Trent University, Peterborough, ON K97 7B8, Canada.

F Corresponding author. Email: wirsinga@uw.edu

Wildlife Research 40(5) 349-357 https://doi.org/10.1071/WR13106
Submitted: 7 June 2013  Accepted: 17 June 2013   Published: 10 July 2013

Abstract

Context: Measures of intake and digestibility from captive feeding experiments are often used to evaluate the nutritional value of plant species to herbivores; however, there is question about how well plant-quality rankings from these trials predict foraging patterns of free-ranging animals. Studies addressing the alignment of results from feeding trials and herbivory in the field using captive and free-roaming conspecifics are needed.

Aims: Our goal was to compare the feeding patterns of snowshoe hares in captive intake and digestion trials with those of free-living conspecifics in the species’ south-western range.

Methods: We conducted in vivo intake and digestion trials using captive hares to determine quality and consumption levels of the predominant conifer species in our study system. In the field, we quantified browsing intensity and over-winter depletion patterns of these conifers. We then compared voluntary intake and nutritional quality measured in captivity to consumption in the field.

Key results: Digestible energy (DE, kJ g–1) of conifers ranged from 11.0 (Pinus contorta) to 13.8 (Pseudotsuga menziesii) among six conifers, and digestible protein (DP, g protein per 100 g feed) from 1.2 (Thuja plicata) to 2.7 (P. contorta). During digestion trials, single-species intake was correlated with the content of digestible protein (DP) and digestible energy (DE). Hares maintained energy balance when fed two single-species diets (Pinus contorta, Pseudotsuga menziesii) and a mixed-species diet. Conifer species on which hares were able to maintain body mass (Pinus contorta, Picea engelmannii, Pseudotsuga menziesii) also tended to be the most heavily exploited by free-living hares. DP content of browse species predicted both browsing intensity and overwinter depletion of conifer species.

Conclusions: Voluntary intake and nutritional quality of browse, especially DP, successfully predicted foraging patterns of free-ranging conspecifics.

Implications: Intake and digestion trials can be a useful tool for better understanding patterns of herbivory in the field, and winter habitat quality for populations in this region is likely to be influenced by access to the most energy- and protein-rich conifers.

Additional keywords: conifer, diet, digestion, energy, Lepus americanus, nutrition, Pinus contorta, protein, Pseudotsuga menziesii.


References

Baraza, E., Zamora, R., and Hódar, J. A. (2006). Conditional outcomes in plant–herbivore interactions: neighbours matter. Oikos 113, 148–156.
Conditional outcomes in plant–herbivore interactions: neighbours matter.Crossref | GoogleScholarGoogle Scholar |

Barbosa, P., Hines, J., Kaplan, I., Martinson, H., Szczepaniec, A., and Szendrei, Z. (2009). Associational resistance and associational susceptibility: having right or wrong neighbors. Annual Review of Ecology Evolution and Systematics 40, 1–20.
Associational resistance and associational susceptibility: having right or wrong neighbors.Crossref | GoogleScholarGoogle Scholar |

Bergman, M., Iason, G. R., and Hester, A. J. (2005). Feeding patterns by roe deer and rabbits on pine, willow and birch in relation to spatial arrangement. Oikos 109, 513–520.
Feeding patterns by roe deer and rabbits on pine, willow and birch in relation to spatial arrangement.Crossref | GoogleScholarGoogle Scholar |

Bookhout, T. A. (1965). Feeding coactions between snowshoe hares and white-tailed deer in northern Michigan. Transaction of the North American Wildlife and Natural Resources Conference 30, 321–335.

Boonstra, R., Krebs, C. J., and Stenseth, N. C. (1998). Population cycles in small mammals: the problem of explaining the low phase. Ecology 79, 1479–1488.
Population cycles in small mammals: the problem of explaining the low phase.Crossref | GoogleScholarGoogle Scholar |

Bryant, J. P., and Kuropat, P. J. (1980). Selection of winter forage by subarctic browsing vertebrates: the role of plant chemistry. Annual Review of Ecology and Systematics 11, 261–285.
Selection of winter forage by subarctic browsing vertebrates: the role of plant chemistry.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL3MXhsVWkuw%3D%3D&md5=006dccda9a7c1c5340885d665aa652eaCAS |

Bryant, J. P., Wieland, G. D., Clausen, T. P., and Kuropat, P. J. (1985). Interactions of snowshoe hare and feltleaf willow in Alaska. Ecology 66, 1564–1573.
Interactions of snowshoe hare and feltleaf willow in Alaska.Crossref | GoogleScholarGoogle Scholar |

Bryant, J. P., Swihart, R. K., Reichardt, P. B., and Newton, L. (1994). Biogeography of woody plant chemical defense against snowshoe hare browsing: comparison of Alaska and eastern North America. Oikos 70, 385–395.
Biogeography of woody plant chemical defense against snowshoe hare browsing: comparison of Alaska and eastern North America.Crossref | GoogleScholarGoogle Scholar |

Bryant, J. P., Clausen, T. P., Swihart, R. K., Landhausser, S. M., Stevens, M. T., Hawkins, C. D. B., Carriere, S., Kirilenko, A. P., Veitch, A. M., Popko, R. A., Cleland, D. T., Williams, J. H., Jakubas, W. J., Carlson, M. R., Bodony, K. L., Cebrian, M., Paragi, T. F., Picone, P. M., Moore, J. E., Packee, E. C., and Malone, T. (2009). Fire drives transcontinental variation in tree birch defense against browsing by snowshoe hares. American Naturalist 174, 13–23.
Fire drives transcontinental variation in tree birch defense against browsing by snowshoe hares.Crossref | GoogleScholarGoogle Scholar | 19422319PubMed |

Burnham, K. P., and Anderson, D. R. (2002). ‘Model Selection and Multimodel Inference: a Practical Information-theoretic Approach.’ 2nd edn. (Springer: New York.)

Coltrane, J. A., Farley, S., Barboza, P. S., Kohl, F., Sinnott, R., and Barnes, B. M. (2011). Seasonal body composition, water turnover, and field metabolic rates in porcupines (Erethizon dorsatum) in Alaska. Journal of Mammalogy 92, 601–610.
Seasonal body composition, water turnover, and field metabolic rates in porcupines (Erethizon dorsatum) in Alaska.Crossref | GoogleScholarGoogle Scholar |

Dearing, M. D., Foley, W. J., and McLean, S. (2005). The influence of plant secondary metabolites on the nutritional ecology of herbivorous terrestrial vertebrates. Annual Review of Ecology Evolution and Systematics 36, 169–189.
The influence of plant secondary metabolites on the nutritional ecology of herbivorous terrestrial vertebrates.Crossref | GoogleScholarGoogle Scholar |

Duncan, A. J., and Gordon, I. J. (1999). Habitat selection according to the ability to eat, digest and detoxify foods. The Proceedings of the Nutrition Society 58, 799–805.
Habitat selection according to the ability to eat, digest and detoxify foods.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BD3c3nsVyntQ%3D%3D&md5=ca3ef2a1b237412c6b4984a926ce0402CAS | 10817146PubMed |

Duncan, A. J., Ginane, C., Elston, D. A., Kunaver, A., and Gordon, I. J. (2006). How do herbivores trade-off the positive and negative consequences of diet selection decisions? Animal Behaviour 71, 93–99.
How do herbivores trade-off the positive and negative consequences of diet selection decisions?Crossref | GoogleScholarGoogle Scholar |

Ellsworth, E. E. (2009). Snowshoe hare nutrition in a conifer forest: effects of winter food on energy use, activity, and demography in a low-density population. Ph.D. Dissertation, University of Idaho, Moscow, ID.

Foley, W. J., and Moore, B. D. (2005). Plant secondary metabolites and vertebrate herbivores – from physiological regulation to ecosystem function. Current Opinion in Plant Biology 8, 430–435.
Plant secondary metabolites and vertebrate herbivores – from physiological regulation to ecosystem function.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2MXlsFGgt78%3D&md5=ade96eae5a3567d34b14c6af8351b336CAS | 15939665PubMed |

Freeland, W. J., and Janzen, D. H. (1974). Strategies in herbivory by mammals: the role of plant secondary compounds. American Naturalist 108, 269–289.
Strategies in herbivory by mammals: the role of plant secondary compounds.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE2cXkvVynsrs%3D&md5=3c7bec74e9124abbb7fcd1ad0cdf9f71CAS |

Goering, H. K., and Van Soest, P. (1970). ‘Forage Analyses.’ (United States Department of Agriculture: Washington, DC.)

Griffin, P. C., and Mills, L. S. (2009). Sinks without borders: snowshoe hare dynamics in a complex landscape. Oikos 118, 1487–1498.
Sinks without borders: snowshoe hare dynamics in a complex landscape.Crossref | GoogleScholarGoogle Scholar |

Hik, D. S. (1995). Does risk of predation influence population dynamics? Evidence from the cyclic decline of snowshoe hares. Wildlife Research 22, 115–129.
Does risk of predation influence population dynamics? Evidence from the cyclic decline of snowshoe hares.Crossref | GoogleScholarGoogle Scholar |

Hodges, K. E. (2000). Ecology of snowshoe hares in southern boreal and montane forests. In ‘Ecology and Conservation of Lynx in the United States’. (Eds L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, G. M. Koehler, C. J. Krebs, K. S., McKelvey and J. R. Squires.) pp. 163–207. General Technical Report RMRS-GTR-30WWW. Agriculture, Forest Service, Rocky Mountain Research Station, Fort Collins, CO.

Hodges, K. E., and Sinclair, A. R. E. (2003). Does predation risk cause snowshoe hares to modify their diets? Canadian Journal of Zoology 81, 1973–1985.
Does predation risk cause snowshoe hares to modify their diets?Crossref | GoogleScholarGoogle Scholar |

Hodges, K. E., Mills, L. S., and Murphy, K. M. (2009). Distribution and abundance of snowshoe hares in Yellowstone National Park. Journal of Mammalogy 90, 870–878.
Distribution and abundance of snowshoe hares in Yellowstone National Park.Crossref | GoogleScholarGoogle Scholar |

Keith, L. B. (1990). Dynamics of snowshoe hare populations. In ‘Current Mammalogy’. (Ed. H. H. Genoways.) pp. 119–195. (Plenum Press: New York.)

Krebs, C. J., Boonstra, R., Boutin, S., and Sinclair, A. R. E. (2001). What drives the 10-year cycle of snowshoe hare? Bioscience 51, 25–35.
What drives the 10-year cycle of snowshoe hare?Crossref | GoogleScholarGoogle Scholar |

Lima, S. L., and Dill, L. M. (1990). Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology 68, 619–640.
Behavioral decisions made under the risk of predation: a review and prospectus.Crossref | GoogleScholarGoogle Scholar |

Lopez-Perez, E. (2006). Natural selenium and planted forages: effects on mule deer and elk in Washington. Ph.D. Dissertation, Washington State University, Pullman, WA.

Miller, A. M., McArthur, C., and Smethhurst, P. J. (2007). Effects of within-patch characteristics on the vulnerability of a plant to herbivory. Oikos 116, 41–52.
Effects of within-patch characteristics on the vulnerability of a plant to herbivory.Crossref | GoogleScholarGoogle Scholar |

Murray, D. L. (2000). A geographic analysis of snowshoe hare population demography. Canadian Journal of Zoology 78, 1207–1217.
A geographic analysis of snowshoe hare population demography.Crossref | GoogleScholarGoogle Scholar |

Murray, D. L. (2003). Snowshoe hare and other hares. In ‘Wild Mammals of North America Vol. II’. (Eds G. A. Feldhamer and B. Thompson.) pp. 147–175. (Johns Hopkins University Press: Baltimore, MD.)

Mysterud, A., and Ostbye, E. (2006). Effect of climate and density on individual and population growth of roe deer Capreolus capreolus at northern latitudes: the Lier valley, Norway. Wildlife Biology 12, 321–329.
Effect of climate and density on individual and population growth of roe deer Capreolus capreolus at northern latitudes: the Lier valley, Norway.Crossref | GoogleScholarGoogle Scholar |

Nams, V. O., Folkard, N. F. G., and Smith, J. N. M. (1996). Nitrogen fertilization stimulates herbivory by snowshoe hares in the boreal forest. Canadian Journal of Zoology 74, 196–199.
Nitrogen fertilization stimulates herbivory by snowshoe hares in the boreal forest.Crossref | GoogleScholarGoogle Scholar |

Pease, J. L., Vowles, R. H., and Keith, L. B. (1979). Interaction of snowshoe hares and woody vegetation. The Journal of Wildlife Management 43, 43–60.
Interaction of snowshoe hares and woody vegetation.Crossref | GoogleScholarGoogle Scholar |

Rafferty, C. M., Lamont, B. B., and Hanley, M. E. (2010). Herbivore feeding preferences in captive and wild populations. Austral Ecology 35, 257–263.
Herbivore feeding preferences in captive and wild populations.Crossref | GoogleScholarGoogle Scholar |

Reichardt, P. B., Bryant, J. P., Clausen, T. P., and Wieland, G. D. (1984). Defense of winter-dormant Alaska paper birch against snowhsoe hare. Oecologia 65, 58–69.
Defense of winter-dormant Alaska paper birch against snowhsoe hare.Crossref | GoogleScholarGoogle Scholar |

Robbins, C. T. (1993). ‘Wildlife Feeding and Nutrition.’ (Academic Press: San Diego, CA.)

Robbins, C. T., Hanley, T. A., Hagerman, A. E., Hjeljord, O., Baker, D. L., and Schwartz, C. C. (1987a). Role of tannins in defending plants against ruminants: reduction in protein availability. Ecology 68, 98–107.
Role of tannins in defending plants against ruminants: reduction in protein availability.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL2sXhtlGmtrg%3D&md5=08f07bb596fe4345b5989091e5f4f3e7CAS |

Robbins, C. T., Mole, S., Hagerman, A. E., and Hanley, T. A. (1987b). Role of tannins in defending plants against ruminants: reduction in dry matter digestion? Ecology 68, 1606–1615.
Role of tannins in defending plants against ruminants: reduction in dry matter digestion?Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL1cXhtFSqu70%3D&md5=c23e2c6d3665a1e0b23544e710234df9CAS |

Rodgers, A. R., and Sinclair, A. R. E. (1997). Diet choice and nutrition of captive snowshoe hares (Lepus americanus): interactions of energy, protein, and plant secondary compounds. Ecoscience 4, 163–169.

Schmitz, O. J. (2008). Herbivory from individuals to ecosystems. Annual Review of Ecology Evolution and Systematics 39, 133–152.
Herbivory from individuals to ecosystems.Crossref | GoogleScholarGoogle Scholar |

Schmitz, O. J., Hik, D. S., and Sinclair, A. R. E. (1992). Plant chemical defense and twig selection by snowshoe hares. Oikos 65, 295–300.
Plant chemical defense and twig selection by snowshoe hares.Crossref | GoogleScholarGoogle Scholar |

Seccombe-Hett, P., and Turkington, R. (2008). Summer diet selection of snowshoe hares: a test of nutritional hypotheses. Oikos 117, 1874–1884.
Summer diet selection of snowshoe hares: a test of nutritional hypotheses.Crossref | GoogleScholarGoogle Scholar |

Sheriff, M. J., Speakman, J. R., Kuchel, J. R., Boutin, S., and Humphries, M. M. (2009). The cold shoulder: free-ranging snowshoe hares maintain a low cost of living in cold climates. Canadian Journal of Zoology 87, 956–964.
The cold shoulder: free-ranging snowshoe hares maintain a low cost of living in cold climates.Crossref | GoogleScholarGoogle Scholar |

Shipley, L. A., Blomquist, S., and Danell, K. (1998). Diet choices made by free-ranging moose in northern Sweden in relation to plant distribution, chemistry and morphology. Canadian Journal of Zoology 76, 1722–1733.
Diet choices made by free-ranging moose in northern Sweden in relation to plant distribution, chemistry and morphology.Crossref | GoogleScholarGoogle Scholar |

Shipley, L. A., Davila, T. B., Thines, N. J., and Elias, B. A. (2006). Nutritional requirements and diet choices of the pygmy rabbit (brachylagus idahoensis): a sagebrush specialist. Journal of Chemical Ecology 32, 2455–2474.
Nutritional requirements and diet choices of the pygmy rabbit (brachylagus idahoensis): a sagebrush specialist.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28Xht1Wgu7fO&md5=656bfdadcfbc0536c80848c0b0898103CAS | 17082988PubMed |

Sinclair, A. R. E., and Smith, J. N. M. (1984). Do plant secondary compounds determine feeding preferences of snowshoe hares? Oecologia 61, 403–410.
Do plant secondary compounds determine feeding preferences of snowshoe hares?Crossref | GoogleScholarGoogle Scholar |

Sinclair, A. R. E., Krebs, C. J., and Smith, J. N. M. (1982). Diet quality and food limitation in herbivores: the case of the snowshoe hare. Canadian Journal of Zoology 60, 889–897.
Diet quality and food limitation in herbivores: the case of the snowshoe hare.Crossref | GoogleScholarGoogle Scholar |

Sinclair, A. R. E., Krebs, C. J., Smith, J. N. M., and Boutin, S. (1988a). Population biology of snowshoe hares. III. Nutrition, plant secondary compounds and food limitation. Journal of Animal Ecology 57, 787–806.
Population biology of snowshoe hares. III. Nutrition, plant secondary compounds and food limitation.Crossref | GoogleScholarGoogle Scholar |

Sinclair, A. R. E., Jogia, M. K., and Andersen, R. J. (1988b). Camphor from juvenile white spruce as an antifeedant for snowshoe hares. Journal of Chemical Ecology 14, 1505–1514.
Camphor from juvenile white spruce as an antifeedant for snowshoe hares.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL1cXkslSltbk%3D&md5=f69f3ea1c0c403d11e137ad29266dafeCAS |

Smith, J. N. M., Krebs, C. J., Sinclair, A. R. E., and Boonstra, R. (1988). Population biology of snowshoe hares. II. Interactions with winter food plants. Journal of Animal Ecology 57, 269–286.
Population biology of snowshoe hares. II. Interactions with winter food plants.Crossref | GoogleScholarGoogle Scholar |

Sponheimer, M., Robinson, T., Roeder, B., Hammer, J., Ayliffe, L., Passey, B., Cerling, T., Dearing, D., and Ehleringer, J. (2003). Digestion and passage rates of grass hays by llamas, alpacas, goats, rabbits, and horses. Small Ruminant Research 48, 149–154.
Digestion and passage rates of grass hays by llamas, alpacas, goats, rabbits, and horses.Crossref | GoogleScholarGoogle Scholar |

Sullivan, T. P., Sullivan, D. S., Lindgren, P. M. F., and Ransome, D. B. (2006). Influence of repeated fertilization on forest ecosystems: relative habitat use by snowshoe hares (Lepus americanus). Canadian Journal of Forest Research 36, 2080–2089.
Influence of repeated fertilization on forest ecosystems: relative habitat use by snowshoe hares (Lepus americanus).Crossref | GoogleScholarGoogle Scholar |

Swihart, R. K., and Bryant, J. P. (2001). Importance of biogeography and ontogeny of woody plants in winter herbivory by mammals. Journal of Mammalogy 82, 1–21.
Importance of biogeography and ontogeny of woody plants in winter herbivory by mammals.Crossref | GoogleScholarGoogle Scholar |

Van Soest, P. J. (1994). ‘Nutritional Ecology of the Ruminant,’ 2nd edn. (Cornell University Press: Ithaca, NY.)

Whittaker, M. E., and Thomas, V. G. (1983). Seasonal levels of fat and protein reserves of snowshoe hares in Ontario. Canadian Journal of Zoology 61, 1339–1345.
Seasonal levels of fat and protein reserves of snowshoe hares in Ontario.Crossref | GoogleScholarGoogle Scholar |

Wirsing, A. J., and Murray, D. L. (2002). Patterns of woody browse consumption by snowshoe hares in the northwestern United States. Ecoscience 9, 440–449.

Wirsing, A. J., and Murray, D. L. (2007). Food supplementation experiments revisited: verifying that supplemental food is used by it intended recipients. Canadian Journal of Zoology 85, 679–685.
Food supplementation experiments revisited: verifying that supplemental food is used by it intended recipients.Crossref | GoogleScholarGoogle Scholar |

Wirsing, A. J., Steury, T. D., and Murray, D. L. (2002). A demographic analysis of a southern snowshoe hare population in a fragmented habitat: evaluating the refugium model. Canadian Journal of Zoology 80, 169–177.
A demographic analysis of a southern snowshoe hare population in a fragmented habitat: evaluating the refugium model.Crossref | GoogleScholarGoogle Scholar |

Wolff, J. O. (1980). The role of habitat patchiness in the population dynamics of snowshoe hares (Lepus americanus). Ecological Monographs 50, 111–130.
The role of habitat patchiness in the population dynamics of snowshoe hares (Lepus americanus).Crossref | GoogleScholarGoogle Scholar |

Zahratka, J. L., and Shenk, T. M. (2008). Population estimates of snowshoe hares in the southern Rocky Mountains. The Journal of Wildlife Management 72, 906–912.