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° . FOODS OF WHITE-TAILED DEER IN THE UPPER GREAT LAKES REGION--A REVIEW

Lynn L. Rogers, Wildlife Biologist, North Central Forest Experiment Station, , St. Paul, Minnesota Jack J. Mooty, Wildlife Biologist, Minnesota Department of Natural Resources Grand Rapids, Minnesota - and Deanna Dawson,i Wildlife Biologist, North .Central Forest Experiment Station, " _ ' St. Paul, Minnesota

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This report summarizes available information on the Research methods to obtain information on food food habits of white-tailed deer (Odocoileus virgini- habits of deer in the UGLR have included analyzing anus) in the northern forests of Minnesota, Wisconsin, stomach contents, examining feeding sites, and ob- and Michigan (fig. 1). This area, referred to here as the serving tame deer on a leash. None of these methods Upper Great Lakes Region (UGLR), is near the north- gives a complete picture of food habits, and each has a ern edge ofthe white-tail's present range (Taylor 1956). different bias as will be discussed below. However, Deer were not abundant in the mature, pine-domi- each method gives an approximation of the diet. nated, "virgin" forests of the UGLR. However, the Results of studies conducted in seven areas of the young hardWood forests created by large-scale logging UGLR during 1936-1976 are presented in tables 1-12 and extensive forest fires allowed deer to expand their (Appendix). Differences in reported diets reflect local range northward (Swift 1946, Stenlund 1958, Erickson differences in vegetation and differences in methods et al. 1961). Today, deer are important both aestheti- of study. Omitted from the tables are results of seven cally and economically in the UGLR. They are viewed studies (Hammerstrom and Blake 1939, Swift 1946, and photographed as an important adjunct to the Dahlbergand Guettinger 1956,KreftingandHansen Re_on's tourist industry, and hunters spend over 200 1963, Orke 1966, McCaffery and Kohn 1971, and milliondollars a year on deer hunting (USDI 1977). Hennings 1977) that did not permit percentage Deer are also the main prey of the timber wolf (Canis •: . lupus) in the UGLR. This Region has the only sizable population of wolves in the contiguous United States (Mech and Karns 1977). The young forest that brought deer to the UGLR is now reaching maturity, and deer populations are de- ' clining (Byelich 1965, Stone 1966, Mooty 1971, Mech and Kams 1977). State governments in this Region are now spending nearly 2 million dollars a year on deer _H_o_ habitat improvement. However, optimal habitat man- agement is difficult because the year-round food habits of deer in the UGLR have not been documented thor-

0n!y recently (Kohn and Mooty 197!, McCaifery et al. _,_.,_,_ o1974,ughly.MootyStudy1976of gr, oBwing-auerseas1977on).foodThishabreivitsewwasempha-begun _ _.,,_,.. _ /. sizes the need for further information and is a step toward disseminating available information on deer Figure 1.--Northern forest area of the midwestern food habits. United States.

. breakdowns of diet or presentation by month. How- bunchberry, wintergreen, strawberry, and barren ever, common and scientific names of foods men- strawberry (McCaffery and Kohn 1971). As spring tioned in those studies are included in Appendix II progresses, new green grass, emerging forbs, and new which lists reported-deer foods in the UGLR. leaves of trees and shrubs become available. Woody twigs and evergreen forbs are forsaken, and new green growth forms over 90 percent of the diet (table 4 and 5) SEASONAL FOODS OF DEER (Pierce 1975). This vegetation is both nutritious and IN THE UPPER GREAT LAKES easilydigestible (Verme and Ullrey 1972). At this REGION time, deer frequently feed in treeless areas such as old fields, roadsides, and powerline rights-of-way which "green up" earlier than the surrounding forests Foods ofdeercan be categorized into such groups as (McCaffery and Creed 1969). Woody browse, conifer needles, evergreen forbs, non- In late spring and early summer, deer, like moose Usevergreene of theseforbs,groupsdeciduousby deer followsleaves, afruit,seasonalfungi,trendetc. (Alces alces), are sometimes observed eating aquatic throughout the UGLR despite local differences in vegetation. Hennings (1977) observed deer in north- vegetatiom _ eastern Minnesota selectively eating (in decreasing order of use) burreeds, filamentous algae, ribbonleaf In Winter, woody browse usually is the main food pondweed, water horsetail, arrowhead, sedges, available, and it forms the bulk of the diet (tables t-4, marsh cinquefoil, white pond lily, yellow pond lily, 11-12). This-food is low in nutrient quality and purple watershield, and St. John's-Tort. Water mil- digestibility, and.deer lose weight on it even when it foil and wild rice also have been reported as deer is available in unlimited quantities (Ullrey et al. foods in the UGLR (Dahlberg and Guettinger 1956, 1964, 1967, 1968, Verme and Ullrey 1972, Grigal et Irwin 1974). Aquatic vegetation is fairly nutritious al. 1979). Aprolonged diet of woody browse causes (Linnet al. 1973) and may be important in supplying whimalnutritionch are commonand starvationin the UGLR(Mautz in1978),late bothwinterof annual requirements of sodium and other nutrients (Botkin et al. 1973, Jordan et al. 1974, Hanson and (Bartlett 1938, Erickson et al. 1961, Stenlund 1970 Karns 1980). Jones 1977). Aquatic feeding reaches its peak during periods of reduced water levels in early summer Although cedar is a preferred winter food (Dahl- (Townsend and Smith 1933, Behrend 1966, Skinner berg and Guettinger 1956), needles of balsam fir, and Telfer 1974, Hennings 1977). The portion of the spruce, and pine apparently make an even poorer diet diet that aquatic vegetation comprises during peri- pristhanedmost58 woodyto 60 percentbrowse species.of the rumenThese needlescontentscorn-of ods of peak use is unknown but is probably low (Hennings 1977, Joyal and Scherrer 1978). starved deer autopsied in late winter during popula- In summer, deer eat the leaves of nonevergreen tion highs of past decades (Aldous and Smith 1938, terrestrial plants, mushrooms, and fruit (Kohn and Dahlberg and Guettinger 1956). However, these needles made up only a small portion of the winter Mooty 1971, McCaffery et al. 1974). Aspen forests, diet of healthier deer studied more recently (tables 1- especially poorly stocked stands or those under 25 years of age, are important summer habitats (McCaf- 3). Among the 'latter deer, the greatest use of balsam fery and Creed 1969, Mooty 1971, McCaffery 1976, _ fir, pine, and spruce needles was by deer that eventu- Bauer 1977, Gullion 1977). Leaves from aspen suck- ally were killed by wolves. Sixteen percent of the 'stomach contents of 32 wolf-killed deer were needles ers less than one year old are a preferred food of these species (Wetzel 1972). (McCaffery 1976). In northern Wisconsin, where After the rigors of a northern winter and a nutri- first-year aspen suckers are abundant due to fre- ' tionaliy marginal diet, fat reserves are depleted and quent clear-cutting of aspen forests, aspen leaves deer become nutritionally stressed (Verme 1969, form a larger portion of the summer diet than does any other item (McCaffery et al. 1974). Leaves from Mautz 1978, Karns 1980). In early spring, metabolic older aspen are less preferred, however 1. Conse- rates increase, does are in the last months of preg- quently, in northern Minnesota where aspen clear- nancy, and a high quality diet is essential to both survival and reproductive success (Julander and cutting is less frequent aspen leaves form a smaller part of the summer diet (Kohn 1970, Pierce 1975). Robinette 1950, L0nghurst et hi. 1952, Verme 1963, Leaves other than those of aspen also are important 1969, Silver et al. 1969). As snow melts on south-facing slopes and-around the bases of trees in the uplands, deer begin supplementing their browse diets with leaves of small plants that remain green/overwinter: 1personal communication with L. Verme, 1980.

. 2 foodsinaspenstands.Vegetationstudieins Minne- varyingwithspecies,subspecieps,henology,andsite sota(Ohmann and Ream 1971),Wisconsin(McCaf- factors(Nagyand Regelin1977).The palatabilityof fery 1976),and Michigan(Bauer1977)showedthat plantsdecreaseassconcentrationsofcertainsecond- principasul mmer deer foodssuchas maple,birch, arycompoundsincrease(Nagy and Regelin1977). willow,juneberry,hazel,cherry,honeysucklebush-, A diversedietmay alsobenefitdeerin another honeysuckleros, e,large-leafaster,and strawberry way.Eatingcertainplantsmay aidinthedigestionof areprevalentin aspenstands, others.For example,some plantsare toolow in In thefall,nonevergreenleavesbecome increas- nitrogen,phosphorus,magnesium,orsulfurforade- inglyscarcewith the exceptionoflarge-leaafster quaterumen digestion,but theseplantsmay be whichpersistsinfrost-sheltestreadndsintolatefall. utilizeifdtheyarecombinedwithothersthatprovide When nonevergreenleavesand othersummer foods the deficientelements(Church1977:138,Hanson become scarce,deer turn to grasses,sedges,and and Jones1977:254,256). evergreenforbsuntilthesebecomecoveredby snow (table1s0and 11).Then deeronceagainbegineating dormant woody browsewhich dominatesthe diet untilthesnow meltsandtheabovecycleisrepeated. The seasonalfeedingpatternindicatesthatdeer PROBLEMS OF DEER IN generallypreferyoungnonevergreenleavesbu_twill NORTHERN FORESTED eat(in_decreasingorderofpreferencema)turenon- VERSUS SOUTHERN evergreenleaves,evergreenforbs,cedarleaves,de- AGRICULTURAL DEER ciduotmwoodybrowse,andconifenreedles.Preferred RANGES IN THE foodsalsomay includearborealfruticoslie chens, GREAT LAKES REGION acorns,fruit,and certainfungiaccordingtostudies conductedin otherregions(Cushwa etal.1970, Harl0w and Hooper 1972,Skinnerand Telfer1974, Crawfordetal.1975).Additionastudyl isneededto assesstheimportanceofthesefoodsintheUGLR. Diversityapparentlyisimportantinthedeerdiet (Verme and Ullrey1972).Dahlbergand Guettinger Deeratthenorthernedgeoftheirrangearelimited (1956)foundthatcaptivewinteringdeermaintained innumber mainlyby problemsofoverwintemrortal- weightbetteron a varietyofsecond-choicewoody ityand nutrition-relaterdeproductivefailures. browsespeciesthan theydidon a dietofstraight Theseproblemsareeasedby improvementsinthe cedar,afirst-choiwintce erfood.MiguelleandJordan quantityand qualityofyear-roundfoodsupplysuch (1980)found thatcaptivemoose chosea diverse asoccurafterextensivlogginge and burning(Erick- summer dietevenwhen moutain-ash,a highlypre- sonetal.1961).However,problemsincreasine years ferred food, was made available ad libitum. Jordan when access to food patches is restricted by unusually (1967.) found deer preference for a species to be deep or long-lasting snow (Moen 1976, 1978, Nelson highest where that species was scarcest, and Mech 1980). Deer on the George Reserve in • ' Perhaps one of the reasons ruminants choose a southern Michigan seldom dig through more than 3 diversediet isto av0idingestingtoomuchofanyoneof inches (7.5 cm) of snow to reach food (Coblentz 1970) the many plant compounds that inhibit digestion and deer in northcentral Minnesota have not been (Nagy et al. 1964, Longhurst et al. 1968, Levin 1976). observed to dig through more than 12 inches (30 cm) Deer have long been known to select the most nutri- of snow (Mooty and Rogers, personal observations). tious forage available (Swift 1948, Weir and Torell Travel becomes difficult when deer sink beyond their !959), and it now is apparent deer also detect and chests (about 20 inches (50 cm)) (Formozov 1946, avoid compounds that inhibit the action of rumen Edwards 1956, Gilbert et al. 1970). Snow in the micro-organisms (Nagy et al. 1964, Longhurst et al. northern forests of Minnesota, Wisconsin, and Michi- 1968, Nagy and Regelin 1977). These secondary gan becomes more than 20 inches deep nearly every compounds, as they are called, have little or no impact winter (Environmental Science Services Adminis- on rumen micro-organisms at low concentrations tration 1968), burying low-growing plants beyond (Nagy and Regelin 1977), but at higher concentra- reach and hampering travel to the extent that even tions, their inhibitory action accelerates (Nagy and woody browse becomes difficult to obtain in quantity. Tengerdy 1968). A wide variety of secondary corn- By comparison, snow in the southern portions of pounds are foundin plants, with the amount and kind these States usually is not as deep or long-lasting.

° StudiesinagriculturalareasinMinnesotashowed numbers declinedto some extentthroughoutthe thatsnow-freepatchesenabledeertoeatnonwoody UGLR (Mech and Karns 1977),but more recently, - foodslongerthanisusualintheUGLR (Moen 1966). deernumbersovermuch oftheUGLR haverecovered The highernutritionapll aneintheseagricultural considerablyduetoa returntomore normalwinters areashelpsdeertowithstandcoldweather(Moen and tighterestrictionson deerhunting(Mechand 1966).They spendmuch oftheiractiveand resting Karns1977,4).However,innortheasternMinnesota, timeneartheirfoodsupplieins openfieldsand are where lossestotimberwolvesarehigh(Mech and notaslikelytouseheavycoverinwinterasaredeer Karns 1977),no increaseindeernumbers was de- farther north (Verme 1965, Moen 1966, Wetze11972). tected through 1978 (Floyd et al. 1979). Deer in agricultural areas also show faster growth rates and higher reproductive rates than those in the forested north, yearling bucks in southern Minne- sota.average 130 pounds (59 kg) field-dressed, while RESEARCH NEEDS IN those in northeastern Minnesota average only 106 THE UPPER GREAT pounds (48 kg). 2Between 29 and 52 percent of female LAKES REGION fawns in southern Minnesota, Wisconsin, and Michi- gan become pregnant compared with only 3 to 11 Additional information is needed on deer diets in percent in the northern portions of those states the snowfree seasons. Most previous studies in the (Verme and Ullrey 1972, Harder 1980). ' UGLR were conducted in winter because it is then Deer in agricultural areas are able to achieve very that food is scarcest and starvation greatest. How- high densities where hunting and other human- ever, it is now known that deer in the UGLR are related mortality factors are controlled. A confined "semi-hibernators"; in winter, their metabolic rates herd at the Twin Cities Army Ammunition Plant decrease, they are less active, and they require less (TCAAP) achieved a density of 200 deer per square food (Silver et al. 1969, Thompson et al. 1973, Moen mile (77/km2) 3, which is several times higher than 1978). Studies of captive deer in northern Michigan densities recorded anywhere in the UGLR (Olson showed some deer can survive overwinter even if 1938, Erickson et al. 1961). Deer at the TCAAP they have no food for several weeks (Ozoga and usually survived overwinter by digging through Verme 1970). It is now apparent that overwinter shallow snow for herbaceous material 3. In years survival depends notonly on winterconditionsbuton when access to herbaceous material was prevented the energy reserves deer accumulate before winter byan icecrust, fawns starved 3. Herbaceous material begins (Mautz 1978). is seldom available during winter in northern Min- Knowledge of spring diet might prove useful to nesota forests, managers because the spring diet of pregnant does is Deer habitat in northern Lake and Cook Counties, especially crucial to the survival of their fawns in the northeastern corner of Minnesota, is (Verme 1962). Studies of captive deer showed that .deteriorating due to succession of aspen forest to fawn mortality was less than 33 percent when moth- balsam fir and spruce (Erickson et al. 1961, Urich ers that were undernourished in winter were we11- 1973, Mech and Karns 1977). Nevertheless, the area nourished in spring. However, when mothers were still appears capable, from a food standpoint, of undernourished in both winter and spring, fawn • supporting at least 10 deer per square mile (4]km2) 3. mortality rose to 90 percent (Verme 1962). Growth of However, during the late 1960s and early 1970s, a fawns depends largely on summer and fall nutrition, series 0f winters with unusually deep or long-lasting and the fall weight of female fawns largely snow caused starvation and reproductive failure determines whether or not the fawns will breed , among those deer and made escape from timber (Harder 1980). wolves more difficult (Mech and Karns 1977). These Information on diet during the snowfree months is natural factors, together with hunting by man, caused difficult to obtain because present techniques are not a severe decline in the deer population during the well suited for determining diet during that period. early1970s(MechandKarns1977).Deernumbersfell One of the most common techniques, examining to fewer than two per square mile (1/kin2) (Floyd et al. feeding sites, depends upon deer leaving part of each 1979) in some areas that formerly had been among the food plant behind so observers can see that some has best deer hunting areas in the state (Olson 1938). Deer

2Unpublished data by Ludwig and Karns on file. 3personal communication with Karns, 1978. 4personal comunication with Win. Creed, 1980.

" 4 been eaten. This is a suitable method for approximat- seasons would reveal any day-night differences in ing use ofwoody browse and large herbaceous plants, habitat use and any seasonal differences in daily " However, any feeding on small plants that are nipped consumption rates. The validity of this approach off Close to the ground, and any feeding on items that might be tested, in part, by determining whether the deer eat whole (mushrooms, berries, acorns, nuts, tame deer survive, reproduce, and raise young as dried leaves, and lichens) is difficult to detect by this successfully as wild ones under conditions of severe method. Moreover, unless radio-collared deer are winters and predation by timber wolves. The number used;feeding sites are difficult to find in the forest, so of deer required for such study would depend upon those examined •tend to be in open or muddy areas the extent to which individuals differed from one where deer or their tracks are more visible (Peek another in their food preferences. 1975). According to Wallmo et al. (1973), inability to Land managers need more information on year- distribute the feeding site sample in proper relation round diet in order to determine which timber man- to the distribution of deer feeding leads to underesti- agement practices produce the best deer habitats. mation0fuse of shrubsandforbsand overestimation Information on year-round diet also is needed to of Use of grasses. Aquatic feeding is missed entirely, determine the effects of such natural factors as fire, Other problems with this method are determining drought, plant disease, and defoliating insects on howmuch of each species was eaten and whether deer habitat. These factors can change both the kinds browsing occurred days or months before the exami- and the nutritional values of the plants available to nations (Peek 1975). deer (McEwen and Dietz 1965, Halls and Epps 1969, Rumen analyses also are fraught with problems. Urich 1973, Lyon et al. 1978, Ohmann and Grigal Usually only a small portion of the rumen material is 1979). Information on year-round diet will lead to a identifiable, with the identifiable items often being better understanding of the relations between deer the least digestible ones (Bergerud and Russell 1964). and their habitat and will enable land managers to Moreover, rumen contents usually are collected from maximize the benefits from deer habitat manage- deer killed along roads or in large open areas. Conse- ment funds. : quently, results are biased toward roadside grasses andother foods that grow in the open. In areas of low deer density, obtaining adequate samples of rumens LITERATURE CITED in all seasons is another major difficulty. Recently, Observations of tame, harnessed deer Aldous, S. E., and C. F. Smith. 1938. Food habits of have been used to determine summer food habits. Minnesota deer as determined by stomach analy- Where comparisons have been made, food selections sis. Transactions of the North American Wildlife by,tame deer did not differ from those of wild deer in Conference 3:756-767. similar habitats (Healy 1967, Longhurst et al. 1968, Bartlett, I. H. 1938. White-tail--presenting Michi- Wallmo and Neff1970). However, in all studies using gan's deer problem. Michigan Department of Con- tame white-tailed deer, the researcher selected the servation Game Division Bulletin, 64 p. feedinghabitat. None of the available methods can be used to determine seasonal changes in amounts Bauer, W. A. 1977. Forage preferences of tame deer ' Consumed per day. in the aspen type of northern Michigan. 85 p.M.S. Clearly, there is a need to develop improved tech- Thesis, Michigan Technological University. niques for determining food habits--especially for Houghton, Michigan. the snowfree months. An observation by Watts Behrend, D. F. 1966. Correlation of white-tailed deer (1964) that an escaped tame deer remained tame activity, distribution and behavior with climatic 4 ' after 2 weeks of freedom suggests that harnesses are and other factors. New York State Conservation unnecessary and that attempts should be made to Department, Job Completion Report, Project study tame radio-marked deer living unfettered and W-105-R-7, 78 p. full time on natural ranges. Such deer could become Bergerud, A. T., and L. Russell. 1964. Evaluation of fully familiar with the habitats and foods available in rumen food analyses for Newfoundland Caribou. their homeranges and could select them without Journal of Wildlife Managment 28(4):809-814. being hindered by researchers. Feeding data ob- Botkin, D. B., P. A. Jordan, A. S. Dominski, H. S. tained by observing these deer would not be biased in Lowendorf, and G. E. Hutchinson. 1973. Sodium the ways inherent in the other methods mentioned, dynamics in a northern ecosystem. Proceedings of Around-the-clock observations of these deer in all theNationalAcademyofScience70(10):2745-2748.

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..Cushwa, C.T:, R. L. Downing, R. F. Harlow, a_d D.F. movements and winter foods of white-tailed deer in Urbston. 1970. The importance of woody twig ends central Wisconsin. Journal of Mammalogy 20:315- to deer in the Southeast. U.S. Department of Agri- 325. culture Forest Service, Research Paper SE-67, 12 p. U.S. Department of Agriculture Forest Service, Hanson, H. C., and R. L. Jones. 1977. The biogeoche- Southeastern Forest Experiment Station, Ashe- Naturalmistry ofHistoryblue, snow,Survey.and Ross'Urbana,geese.Illinois.320 p. Illinois ville, North Carolina. Dahlberg, B. L., and R. C. Guettinger. 1956. The Harder, J. D. 1980. Reproduction of white-tailed deer . white-tailed deer in Wisconsin. Wisconsin Conser- in the north central United States. P. 23-35. In vation Department Technical Wildlife Bulletin 14. Ruth L. Hine and Susan Nehls, eds. White-tailed deer population management in the north-central 282 p. Madison, Wisconsin. States. Proceedings 1979 Symposium North Cen- Edwards, R. Y. i956. Snow depths and ungulate tral Section Wildlife Society. 116 p. abundance in the mountains of western Canada. Journal of Wildlife Management 20(2)'159-168 Harlow, R. F., and R. G. Hooper. 1971. Forages eaten • by deer in the southeast. Southeast Association of Environmental Science Services Administration. Game and Fish Commissioners Annual Confer- " 1968. Climatic atlas of the United States. 80 p. U.S. ence Proceedings 25"18-46. Department of Commerce. Healy, W. M. 1967. Forage preferences of captive Er.ickson, A. B., V. E. Gunvalson, M. H. Stenlund, D. deer while free- ranging in the Allegheny National W. Burcalow, and L. H. Blankenship. 1961. The Forest. 93 p. M.S. Thesis. Pennsylvania State •white-tailed deer in Minnesota. Minnesota Depart- University, University Park, Pennsylvania. •: ' ment of Conservation Technical Bulletin 5, 64 p. Irwin, L. L. 1974. Relationship between deer and Floyd, T.J., L. D. Mech, and M. E. Nelson. 1979. An moose on a burn in northeastern Minnesota. 51 p. improved method of censusing deer in deciduous- M.S. Thesis, University of Idaho, Moscow, Idaho. coniferous forests. Journal of Wildlife Manage- ment 43(1):258-261. Jordan, P. A. 1967. Ecology of migratory deer in the Formozov, A. N. i946. Snow cover as an integral San Joaquin river drainage. 278 p. Ph.D. Disserta- factor of the environment and its importance in the tion. University of California, Berkeley, California. ecology Of mammals and birds. Materials for Jordan, P. A., D. B. Botkin, A. S. Dominski, H. S. Fauna and Flora of the USSR. New series Zoology Lowendorf, and G. E. Belovsky. 1974. Sodium as a 5, 152 p, (Translation from Russian) Boreal Insti- critical nutrient for the moose of Isle Royale. •tute, University of Alberta. Proceedings of the North American Moose Confer- Gilbert, P. F., O. C. Wallmo, and R. B. Gill. 1970. ence Workshop 9:13-42. Effect of snow depth on mule deer in Middle Park, Joyal, R., and B. Scherrer. 1978. Summer movements Colorado. Journal of Wildlife Management and feeding by moose in western Quebec. Canadian 34(!):15-23. Field:Naturalist 92(3):252-258.

. o 6 Julander, O_, and W. L. Robinette. 1950. Deer and McCaffery, K. R., and W. A. Creed. 1969. Signifi- cattle range relationships on Oak Creek range in cance of forest openings to deer in northern Wis- Utah. Journal of Forestry 48(6):410-415. consin. Wisconsin Department Natural Resources Karns, P. D. 1980. Winter--the grim reaper. Pro- Technical Bulletin 44, 104 p. ceedings of the Symposium on white-tailed deer McCaffery, K. R., and B. E. Kohn. 1971. Relation of management in the North Central U.S. [In press], forest cover types to deer populations. Wisconsin Kohn, B. E. 1970. Summer movements, range use, Department of Natural Resources Progress Report, and food habits of white-tailed deer in north cen- Project W-141-R-6, Study 201, 9 p. mimeo. tral Minnesota. 38 p.M.S. Thesis, University of McCaffery, K. R., J. Tranetzke, and J. Piechura, Jr. Minnesota, St. Paul, Minnesota. 1974. Summer foods of deer in northern Wisconsin. Kohn, B. E., and J. J. Mooty. 1971. Summer habitat of Journal of Wildlife Management 38(2):215-219. White-tailed deer in north central Minnesota. McEwen, L. C., and D. R. Dietz. 1965. Shade effects Journal of Wildlife Management 35(3):476-487. on chemical composition of herbage in the Black Krefting_ L. K., and H. L. Hanson. 1963. Use of Hills. Journal of Range Management 18(4):184- .phytocides to improve deer habitat in Minnesota. 190. Southern Weed Control Conference (Mobile, Ala- Mech ,L. D., and P, D. Karns. 1977. Role of the wolf in bama) 16:209-216 a deer decline in the Superior National Forest. U.S. Levin, D. A. 1976. The chemical defenses of plants to Department of Agriculture Forest Service, Re- pathogens and herbiVores. Annual review of ecol- search Paper NC-148, 23 p. U.S. Department of Agriculture Forest Service, North Central Forest ogy and systematics 7:121-159. Experiment Station, St. Paul, Minnesota. Linn, J. G., R.D. Goodrich, J. C. Meiske, and J. Staba. Miguelle, D. G., and P. A. Jordan. 1980. The impor- 1973. Aquatic plants from Minnesota: Part 4--nu- tance of diversity in the diet of moose. In Proceed- trient composition. Water Resources Research ings 15th North American Moose Conference • Center Bulletin 56, 22p. University of Minnesota, Workshop. [In press]. St. Paul, Minnesota. Moen, A. N. 1966. Factors affecting the energy ex- Longhurst, W. M., A. S. Leopold, and R. F. Dasmann. change and movements of white-tailed deer in 1952, A survey of California deer herds, their western Minnesota. 121p. Ph.D. Thesis, Univer- ranges and management problems. California De- sity of Minnesota, St. Paul, Minnesota. partment ofFish and Game. Game Bulletin 6,136 p. Moen, A. N. 1968. Surface temperatures and radiant L0nghurst, W. M., H. K. Oh, M. B. Jones, and R.E. heat loss from white-tailed deer. Journal of Wild- Kepner. 1968. A basis for the palatability of deer life Management 32(2):338-344. forage plants. North American Wildlife Proceed- ings_33:181.192. Moen, A. N. 1976. Energy conservation by white- tailed deer in the winter. Ecology 57:192-198. Lyon, L. J., H. S. Crawford, E. Czuhai, R. L. Fredrik- son, R. F. Harlow, L. J. Metz, and H. A. Pearson. Moen, A. N. 1978. Seasonal changes in heart rates, 1978, Effects of fire on fauna. U.S.Department of activity, metabolism, and forage intake of white- Agriculture ForestService, General Technical Re- tailed deer. Journal of Wildlife Management port WO-6. 41 p. 42:715-738. •MaUtz, W. W. 1978. Sledding on a bushy hillside: the Mooty, J. J. 1971. The changing habitat scene, p. 27- fat cycle in deer. Wildlife Society Bulletin 6(2):88- 33. In The white-tailed deer in Minnesota. M. N. 90. ' Nelson, ed. 88 p. Minnesota Department of Natural Resources. Maycock, P. F., and J. T. Curtis. 1960. The phytosoci- ology of boreal conifer-hardwoods forests of the Mooty, J. J. 1976. Year-round food habits of white- Great Lakes Region. Ecological Monograph 30:1- tailed deer in northern Minnesota. Minnesota 35. Wildlife Research Quarterly 36(1):11-36. McCaffery, K. R. 1976. Vegetation characteristics of Nagy, J. G., and W. L. Regelin. 1977. Influence of aspenstands. Wisconsin Department of Natural plant volatile oils on food selection by animals. Resources Final Report, Project W-141-R-11, XIIIth International Congress of Game Biolo- Study 205, 27 p. mimeo, gists:225-230.

7 Nagy, J. G.I and R. P. Tengerdy. 1968. Antibacterial Stenlund, M. 1970. Deer wintering conditions and action of essential oils ofArtemisia as an ecological mortality report. Minnesota Quarterly 30(2):102- • factor, iI. Antibacterial action of the volatile oils of 103. Artemisia otridentata (big sagebrush) on bacteria Stone, R. N. 1966. A third look at Minnesota's timber. from the rumen of mule deer. Applied Microbiology U.S. Department of Agriculture Forest Service 16:441-444. Bulletin NE-1, 64 p. U.S. Department of Agricul- Nagy, J. G., H. W. Steinhoff, and G. M. Ward. 1964. ture Forest Service, Northeastern Forest Experi- Effects of essential oils of sagebrush on deer rumen ment Station, Broomall, Pennsylvania. microbial function. Journal of Wildlife Manage- Swift, E. 1946. A history of Wisconsin deer. Wisconsin ment 28:785-790. Conservation Department Publication 323, 96 p. Nelson, M. E., and L. D. Mech. 1980. Effects of winter Swift, E. 1948. Deer select most nutritious forages. severity on winter deer weights in Minnesota. Journal of Wildlife Management 12:109-110. Journal .of Wildlife Management [In press]. Taylor, W. P. 1956. The deer of North America. 668 p. Ohmann, L. F., and D. F. Grigal. 1979. Early revege- The Stackpole Company and The Wildlife Manage- tation and nutrient dynamics following the 1971 ment Institute, Washington, D. C. Little Sioux forest fire in northeastern Minnesota. Forest Science Monograph 21, Supplement t_oFor- Thompson, C. B., J. B. Holter, H. H. Hayes, H. Silver, and W. E. Urban, Jr. 1973. Nutrition of white- est Science 25(4), 80 p. tailed deer. I. Energy requirements of fawns. Jour- Ohmann, L. F., and R. R. Ream. 1971. Wilderness nal of Wildlife Management 37(3):301-311. ecology: virgin plant Communities of the Boundary WatersCanoe Area. U.S. Department of Agricul- Townsend, M. T., and M. W. Smith. 1933. The white- ture Forest Service Research Paper NC-63, 55 p. tailed deer of the Adirondacks. (1) Preliminary U.S. Department of Agriculture Forest Service, survey of the white-tailed deer of the Adirondacks, •North-Central Forest Experiment Station, St. (2) Ecology of the white-tailed deer in summer with special reference to Adirondacks. Roosevelt Wild- . Paul, Minnesota. life Bulletin 6(2):161-325. Ols0n_ H. F. 1938. Deer tagging and population studies in Minnesota. Transactions North Ameri- Ullrey, D. E., W. G. Youatt, H. E. Johnson, P. K. Ku, can Wildlife Conference 3:280-286. and L. D. Fay. 1964. Digestibility of cedar and aspen browse for the white-tailed deer. Journal of Orke, D. N. 1966. Patterns and intensity of white- Wildlife Management 28(4):791-797. tailed deer browsing on the woody plants of Itasca • State Park. 61 p. M. S. Major Report. University of Ullrey, D. E., W. G. Youatt, L. D. Fay, B. E. Brent, Minnesota, St. Paul, Minnesota. and K. E. Kemp. 1967. Digestibility of cedar and jack pine browse for the white-tailed deer. Journal Ozoga, J. J., and L. J. Verme. 1970. Winter feeding of Wildlife Management 31(3):448-454. patterns of penned white-tailed deer. Journal of Wildlife Management 34(2):431-439. Ullrey, D. E., W. G. Youatt, L. D. Fay, B. E. Brent, and K. E. Kemp. 1968. Digestibility of cedar and •- . Peek, J. M. 1975. A review of moose food habits balsam fir browse for the white-tailed deer. Jour- studies in North America. Naturaliste Canadien nal of Wildlife Management 32(1):162-171.

• 10i:193-215. U.S. Department jof Interior. 1977. 1975 national Pierce,D. E., Jr. 1975. Spring ecology of white-tailed survey of hunting, fishing, and wildlife-associated deer in north central Minnesota. 55 p. M. S. Thesis. recreation. 91 p. U.S. Department of the Interior. University of Minnesota, St. Paul, Minnesota. Urich, D. L. 1973. Nutrient levels in deer and moose Silver, H., N. F. Colovos, J. B. Holter, and H.H. forage in northern Minnesota as related to site Hayes. 1969. Fasting metabolism of white-tailed characteristics. 138 p.M.S. Thesis University of deer. Journal of Wildlife Management 33(3):490- Minnesota, St. Paul, Minnesota. 498. . Verme, L. J. 1962. Mortality of white-tailed deer Skinner, W. R., and E. S. Telfer. 1974. Spring, fawns in relation to nutrition. Proceedings Na- summer, and fall foods of deer in New Brunswick. tional Deer Disease Symposium 1"15-38. Journal of Wildlife Management 38(2):210-214. Verme, L. J. 1963. Effect of nutrition on growth of Stenlund, M. 1958. A history of Itasca County deer. white-tailed deer fawns. Transactions North Conservation Volunteer 21(125): 19-24. American Wildlife Conference 28:431-443.

. 8 Verme, L. J. 1965. Swamp conifer deer yards in Reichert. 1973. Accuracy of field estimates of deer northern Michigan. Journal of Forestry 63(7):523- food habits. Journal of Wildlife Management -529. 37(4):556-562. Verme, L. J. 1969. Reproductive patterns of white- Wambaugh, J. R. 1973. Food habits and habitat tailed deer related to nutritional plane. Journal of selection by white- tailed deer, and forage selection Wildlife Management 33:881-887. in important habitats in northeastern Minnesota. Verme, L. J., and D. E. Ullrey. 1972. Feeding and 71 p. M.S. Thesis. University of Minnesota, St. nutrition of deer. p. 275-291. In Digestive physiol- Paul, Minnesota. ogy and nutrition of ruminants, vol. 3: practical Watts, C. R. 1964. Forage preferences of captive deer. nutrition. D. C. Church, ed. publisher and Oregon while free- ranging in a mixed oak forest. 65 p. M.S. State University Bookstores, distributor Corvallis, Thesis. Pennsylvania State University, University Oregon. _ Park, Pennsylvania. Waddell, B. H., Jr. 1973. Fall ecology of white-tailed Weir, W. C., and D. T. Torell. 1959. Selective deer inlnorth central Minnesota.'46 p.M.S. Thesis. by sheep as shown by a comparison of the chemical University of Minnesota, St. Paul, Minnesota. composition of range and forage obtained Wallmo, D. C., and D. J. Neff. 1970. Direct observa- by hand clipping and that collected by esophageal tions of tamed deer to measure their consumption fistulated sheep. Journal of Animal Science of natural forage, p. 105-110. In Range and Wildlife 18:641-649. Habitat Evaluation--A Research Symposium. Wetzel, J. F. 1972. Winter food habits and habitat U.S.. Department of Agriculture Forest Service, preferences of deer in northeastern Minnesota. 106 Miscellaneous Publication 1147, 220 p. p.M.S. Thesis. University of Minnesota, St. Paul, WalImo, O. C.; R. B. Gill, L. H. Carpenter, and D.W. Minnesota.

. 9 APPENDIX °

Table 1.---Plants eaten by white-tailed deer in northern Minnesota in January

(In estimated percents of the diet) z -.

Feedinqsites Rumens North-central Northeast (northeast) 1971-19742 1971s 19724 1971s Beakedhazel ' 37 30 35 16 Mountainmaple 10 29 25 3 Dogwood 1 15 10 6 Juneberry 12 6 14 2 Arboreallichens ' 14 Balsamfir <1 < 1 13 Green.alder 8 1 1 8 Blueberry 8 Redmaple 4 6 Quakingaspen 2 4 1 Paperbirch 4 1 1 Grasses <1 4 ,' Goldthread 4 Cherry 2 1 3 3 Labradortea 3 2 1 •Whitecedar < 1 3 < 1 White pine 2 1 Blackspruce 2 Wintergreen 2 Willow 1 1 1 <1 Speckledalder < 1 < 1 1 •Bush-hOneysuckle < 1 1 Arrowwood 1 <1 Redoak <1 <1 Honeysuckle <1 <1 Jackpine <1 Unidentifiedleaves 6 Unidentifiedtwigs 15 Numberexamined 9 26 16 8 Numberof bites 2,064 24,797 7,027 _Not.includedintables1-3and12are22taxawhoseusebydeerdid.notexceed1.5percentofthedietinanystudyinanymonthduringDecember-March.These taxain¢!udeclubmoss,brackenfern,mosses,large-leafaster,blackmedic,strawberry,marshmarigold,goldenrod,thistles,trailingarbutus,redraspberry, gooseberry;rose,:redelderberry,leatherleaf,sweetgale,swambirch,p sumac,mountain-ash,blackash,balsampoplar,andredpine. 2Mooty(unpublished)..DataforDecember-Marchwereobtainedbyexamining43feedingsitesinItascaCounty,north-centralMinnesota,during1971-197For4. asummaryofthesedataby2-monthintervals,seeMooty(1976).Deerdensityduringthestudywas4to8deerpersquarekilometeranddeclining,accordingto annualspringpelletcountsconductedbytheMinnesotaDepartmentofNaturalResources. 3FromWetzel(1972).DataforDecember-Marchwereobtainedbyexamining82feedingsitesand32wolf-killedrumensinSt.LouisandLakeCountiesin •northeasternMinnesotaduring1968-197Deer1. densitywas4to6deerpersquarekilometeranddeclining(P.D.Karns,1979,personalcommunication;Mechand Karns1977Rogers; etal.1980). 4FromWambaugh(i973).DataforJanuary-Marchwereobtainedbyexamining50feedingsitesin1972inthesameareathathadbeenstudiedbyWetzel (above).

' i0 Table 2.--Plants eaten by white-tailed deer in northern Minnesota in February

" (In estimated percents of the diet) 1

Feedingsites Rumens North-central Northeast (northeast) " 19722 1971a 19724 19713 White.cedar 8 52 Beaked.,hazel 35 33 48 4 Mountainmaple 29 37 Labradortea " 14 1 Whitepine 3 12 Blueberry 10 Jackpine 9 8 Juneberry 8 2 4 1 Dogwood 8 1 2 Willow ' 6 1 <1 Speckledalder 5 2 <1 5 Balsamfir • 1 5 Redmaple 5 2 Arboreallichens 4 Honeysuckle 3 <1 Quakingaspen 2 2 Arrowwood 2 <1 , Greenalder. 2 <1 <1 Hawthorn 2 Americanhazel 2 Cherry 1 1 1 <1 Paperbirch <1 1 <1 1 Wintergreen 1 Sweetfern 1 Redoak <1 < 1 Bush-honeysuckle < 1 Blackspruce <1 Grassessedges, <1

• Unidentifiedleaves 1 Unidentifiedtwigs 4 Numberexamined 7 37 20 15 , Numberof bites 1,458 35,200 12,616 _Notincludedin,tabl1-3esand12are22taxawhoseusebydeerdidnotexcee1d.5perceontfthedieint anystudyinanymonthduringDecember-MaThrch.ese taxaincludclubmosse bracken, fernmosses, large, -leasteraf black, medicstrawb, errymarsh, marigoldgolden, rodthistles, trailing, arbutus,redraspberry, , gooseberryrose, red_ elderberryleatherleal sweetf, gale,swambirch,p sumac,mountain-ash,blackash,balsampoplar,andredpine. 2Mooty(_npublished).DataforDecember-Marchwereobtainedbyexamining43feedingsitesinItascaCounty,north-centralMinnesota,during1971-197For4. asummaryofthesedataby2-monthintervals,seeMooty(1976).Deerdensityduringthestudywas4to8deerpersquarekilometeranddeclining,accordingto annual.springpelletcountsconductbyedtheMinnesotaDepartmentofNaturalResources. , 3FromWetzel(i972).DataforDecember-Marchwereobtainedbyexamining82feedingsitesand32wolf-killedrumensinSt.LouisandLakeCountiesin northeasternMinnesotaduring1968-1971.Deerdensitywas4to6deerpersquarekilometeranddeclining(P.D.Karns,1979,personalcommunication;Mechand Karns1977R; ogerseta/.1980). ' 4FromWambaugh(1973).DataforJanuary-Marchwerobtaine ebydexamining50feedingsitesinthesamare eathathadbeestudin ebydWetzel(above).

' 11 Table 3._Plants eaten by white-tailed deer in northern Minnesota in March

o

" (In estimated percents of the diet) z _

Feedinqsites Rumens North-central Northeast (northeast) . 19722 1971a 19724 1971a 1937s Beaked.hazel 52 44 51 4 Balsamfir 1 <1 5 43 Whitecedar. _ 26 35 3 Jack.pine 5 17 Mountain-maple , 12 16 Pine 14 Juneberry. 8 , 2 11 Dogwood 3 1 4 9 <1 Paperbirch 5 1 <1 1 5 Greenalder 4 ' 5 2 <1 Arrowwood <1 <1 5 Arboreallichens 5 Red,-maple <1 2 4 <1 Willow 1 1 <1 1 4 •Cherry 1 1 < 1 3 < 1 Speckledalder 1 3 <1 Blackspruce 1 3 ' Redoak <1 < 1 3 < 1 Grasses,sedges <1 3 Labradortea <1 3 Honeysuckle 3 Quakingaspen 2 1 2 White pine 2 1 Blueberry 2 Large-toothedaspen 1 Bush-honeysuckle < 1 Wintergreen <1 Bunchberry <1 Unidentifiedleaves 1

• . .Unidentifiedtwigs 8 Othermaterial 7 ' NUmberexamined 15 15 14 4 51

.Numberof bites 3,700 19,666 6,228 _Notincludedintables1-3and12are22taxawhoseusebydeerdidnotexceed1.5percentofthedietinanystudyinanymonthduringDecember-MarThesch e taxaincludeClubmoss,brackenfern,mosses,large-leafaster,blackmedicstra, wberry,marshmarigold,goldenrod,thistles,trailingarbutus,redraspberry, gooseberry,rose,redelderberry,leatherleaf,sweetgale,swampbirch,sumacmountain-ash,, blackash,balsampoplar,andredpine. 2Moot(unpublished).y DataforDecember-Mwerarcehobtainedbyexamining43feedingsitesinItascCounty,a north-centralMinnesota,during1971-1974.For asummaoryfthesedataby2-monthintervals,seeMooty(1976)Deer. densityduringthestudwy as4to8deerpersquarkilometere anddeclining,accordingto annualspringpelletcountsconductedbytheMinnesotaDepartmentofNaturalResources. 3FromWetzel(!972).DataforDecember-Mwearcrehobtainedbyexaminin82gfeedingsitesand32wolf-killedrumensinSt.LouisandLakeCountiesin northeasternMinnesotaduring1968-1971.Deerdensitywas4to6deerpersquarekilometeranddeclinin(Pg.D.Karns,1979,personalcommunication;Mechand Karn!977;s Rogersetal.1980). .4FromWambaugh(1973).DataforJanuary-Marwcerehobtainedbyexamining50feedinsitesg inthesameareathathadbeenstudiedbyWetzel(above). 5FromAldouands Smith(1938).DatawereobtainedinnortheasternMinnesotabyexaminin51grumensfromdeerthathaddiedofstarvation,predation, sickness,ekposurelorhavingbeenstrucbyk vehicleslateinthewinterof1936-1937.Deerdensitywas4to8persquarekilometaecr cordintogdeerdrives ConductedbytheCivilianConservationCorpsintheSuperiorNationalForestin1936.

.12 Table 4.---.Plants eaten by white-tailed deer in north- Table 5.--Plants eaten by white-tailed deer in north-

. , ceritral Minnesota in April of 1970 and 1971 central Minnesota in May of 1970 and 1971 (In esti_mated percents of diet) 1 (In estimated percents of diet) 1

Feedingsites Rumens Feedingsites Rumens White-cedar 48 Grasses 45 7 Beakedhazel 21 2 Sedges 17 •Greenalderandspeckledalder 2 18 Willow 3 6 Bush-honeysuckle 15 Goldenrod(dead) 5 .Grasses 14 Marshmarigold 5 1 Redosier .dogwood 10 Redosier dogwood 4 2 Latelow blueberry 9 Woodanemone 4 Juneberry 6 Brackenfern 4 -Goldenro.d(dead) 5 Trailingarbutus 3 Sedges. 5 Falselily-of-the-valley 3 Blackash 3 Rose 3 Honeysuckle 3 Yellow bellwort 3 Marshmarigold 3 Latelow blueberry 3 Lichens 2 Large-leafaster- 2 UnidentifiedWoodybrowse 30 Clinton'slily 2 Numberof sites or rumens Unidentifiedleaves 64 examined 11 5 Unidentifiedforbs 13 Numberof bites 1,833 Numberof sites or rumens 1FromPterce(1975)Da. tawereobtainedbyexamining39feedingsites examined 16 3 . and3 rumensinItascacountynort, centrah Minnesol duringta, April-June Numberof bitestallied 4,595 t970-1971Five. rumensfromdeerthatstarveindAprilinItascaCountywere 1FromPierce(1975).Foradditionaldetailsofthestudy,seefootnoteto examined.Deerdensitywas4 to8 persquarekilometeranddeclining, table4. Itemsusedbydeerinamountslessthan2percentinMayinclude accordingto annuaslpringpellecount tconducs tbyedtheMinnesota hazel,quakinaspg enand, pyrola. DepartmenofNt aturaResoul rces.Itemsusedinamounltesss thantwo percentinAprilincludetwigsd ofpaperbirch,quakingaspen,andcherryand sprigsofjackpineandwhitepine.Bush-honeysuckle,grass,sedge,and marshmarigoldwereatene mainlyinlateAprilaftesomr snowe hadmelted.

°

13 Tabl e 6.--Plants eaten by white-tailed deer in the northern Great Lakes Region in June . (In estimated percents of the diet) _

- Feedingsites Observationof tame deer (Minnesota) Rumens (Michigan) Forest2 Burn3 Minnesota4 Wisconsins Aspen6 Clearcut6 Aspen 1 16 6 29 21 37 Redmapleand mountainmaple 8 22 14 5 1 9 Brackenfern 13 < 19 7 Dogwood 1 18 Beakedhazel 9 4 14 16 Bush-honeysuckle 1 . 16 5 Strawberry 2 5 15 4 9 Wildsarsaparilla 14 < Lilyfamily 2 12 3 <

Watermilfoil 11 ! Raspberry 8 5 Willow 8 1 4 < Hawkweed 8 2 < Honey_,uckle 8 Grassesandsedges 7 3 6 6 2 2 Chokecherry 1 7 6 Aster,mainly large-leafedaster 6 1 6 2 'Paperbirch 1 6 2 < Sweetpea 5 5 Yellowbellwort 4 Goldenrod 3 2 3 < Bunchberry 3 < < Juneberry 2 2 MiScellaneousspecies 20 Unidentifiedleaves 9 Numberof sitesor rumensexamined 12 6 3 24 Numberof bites 2,763 492 1Plantswhoseusebydeerdidnotreach2percentofthedietinanystudyinJuneweremushrooms,pyrola,bluebell,spreadingdogbane,violet,goosefoot, • . bedstraw,commondandelion,yarrowmpearlyeverlasting,indianhemp,hedgebindweed,scarletcolumbine,fireweed,thistle,latelowblueberry,thimbleberry, currant,rose:arrowwood,blackash,buroak,andwhitespruce. . 2FromPierce(1975).Foradditionaldetailsofthestudy,seefootnotetotable4. 3FromIrwin(1974).Datawereobtainedbyexamining52feedingsitesintheLittleSiouxBurnareainSt.LouisCounty,northeasternMinnesotaduringApril- December1973.Deerdensitywas4 to6 persquarekilometer(P.D.Karns,1979,personalcommunication). 4FromKohn(1970).Datawereobtainedbyexamining31feedingsitesand9rumensinItascaCounty,north-centralMinnesota,duringJune-August1968-1969. DeerpopulationdataareasforPierce(1975)(above).Kohndidnotmentionbyname11browseand20forbtaxaeachofwhichaccountedfornomorethan2 percentofthedietinanymonthofstUdy, SFromMcCafferyetal.(1974).DatawereobtainedinnorthernWisconsinduring1969-197by0examining76rumensfromroad-killeddeer(15inApril-May,42 inJune-September,19inOctober-November).Seventygenera,excludinggeneraofgrasses,sedges,andmushrooms,werefoundinthe76rumens,but17items accountedfor80percent.otheaggregatef volumesfoiA" pril-November.These17itemswereaspenleaves(16percent),graminoids(1percent),barrenstrawberry (7percent),aster(5percent),bush-honeysuckle(6percent),strawberry(4percent),cherry(4percent),oakacorns(6percent),wintergreen(3percent)(4 percent),mushrooms(2percent),mapleleaves(2percent),solomon'sseal(2percent),falselily-of-the-valley(2percent),andbunchberry(2percent). Approximatelymonthlybreak-downsareavailableforJuneonly.Deerdensitywas6 to8persquarekilometeranddeclining(Wm.F.Creed,1978,personal communication). 6FromBauer(1977).Datawereobtainedbyobservationof3tame,harnesseddeerinalO-acrematureaspenstandandinanadjacent1O-acre1-year-oldaspen clearcutdur!ngJune-August1976inDickinsonCountyinMichigan'sUpperPeninsula.Deerdensitywas11persquarekilometeranddeclining(L.F.Verme,1979, personalcommunication). Table 7.--Plants eaten by white-tailed deer in the northern Great Lakes Region in July o (In estimated percent of the diet) _

Feedingsites Rumens Observationsof tamedeer (Minnesota) (Minnesota) (Michigan) Forest7 Burns Forest4 Aspen6 Clearcut6 Aspen 7 3 22 26 20 Beaked-hazel 26 9 20 Strawberry 3 11 10 17 •Redandmountainmaple 10 5 16 8 ,13 Goldenrod _ 1 12 4 2 3 Chokeandpin cherry <1 12 4 3 2 Brackenfern . 3 3 12 3 Asterslmainly large-leafedaster 9 10 3 5 Raspberry 1 10 6 Roughcinquefoil , 10 Willow 9 4 2 ; Juneberry 3 8 ;2 I Spottedjewelweed 3 8 BUsh-honeysuckle < 1 2 7 Grasses.andsedges 1 6 2 2 Violet <1 5 Paperbirch 3 4 1 <1 • Commongeranium 4 ' Watermilfoil 4 Fireweed 3 Clover 2 2 Hedgebindweed 2 2 Falselily-of-the-valley 1 2 <1 Hawkweed 2 <1 : Spreadingdogbane 1 2 Bicknell'sgeranium 2 Mushrooms <1 2 Arrowwood 2 Unidentifiedleaves 7 Miscellaneousspecies 17 , ' Numberof feedingsites or rumensexamined 21 8 3 Numberof bitestallied 5,098 778 _Plantswhoseusebydeerdidnotreach2percentofthedietinAugustincludedHawkweed,commondandelion,roughcinquefoil,pyrola,pearlyeverlasting, rue, strawberry, hedge t ' bsarsalacksnakeroot,parilla,honeysuckle;meadowgreenindianalder,hemp,andwhitebluebell,spruce.sweeffern,barren arrowwood,thistle,bedstraw,mint,wildlettuce, bindweedwild • 3FromIrwin(1974).Datawereobtainedbyexamining52feedingsitesintheLittleSiouxBurnareainSt.LouisCounty,northeasternMinnesotaduringApril- December1973.Deerdensitywas4 to6 persquarekilometer.(P.D.Karns,1979,personalcommunication). 4FromKohn(1970).Datawereobtainedbyexamining31feedingsitesand9rumensinItascaCounty,north-centralMinnesota,duringJune-August1968-1969. DeerpopulationdataareasforPierce(1975)(above).Kohndidnotmentionbyname11browseand20forbtaxaeachofwhichaccountedfornotmorethan2 percentofthedietinanymonthofstudy. 6FromBaue(19r 77)Data. wereobtainedbyobservationof3tame,harnesseddeerina1O-acrematureaspenstandandinanadjacent1O-acre1-year-oldaspen clearcutduringJune-August1976inDickinsonCountyinMichigan'sUpperPeninsula.Deerdensitywas11persquarekilometeranddeclining(L.F.Verme,1979 personalcommunication). 7Mooty(unpublished).DatawereobtainedbyexaminingfeedingsitesinItascaCounty,north-centralMinnesotaduring1968-1971.Forasummaryofthesedata by2-mOnthintervals,seeMooty(1976).SomeofthedatausedbyMootywerecontributedbyKohn(1970)andWaddell(1973).Deerdensitywas4to8persquare kilnmeteranddeclining,accordingtoannualspringpelletcountsconductedbytheMinnesotaDepartmentofNaturalResources.

I 15 Table 8.rePlants eaten by white-tailed deer in the northern Great Lakes Region in August - (In estimated percent of the diet)_ ° , Feedingsites Observationsoftamedeer ' - (Minnesota) Rumens (Michigan) Forest7 Burns (Minnesota)4 Aspen6 Clearcut6 Aspen 8 12 23 9 Beakedand americanhazel 15 <2 15 23 BuSh-honeysuckle 7 20 9 " Strawberry 1 <1 9 17 Mushrooms 17 Chokeandpincherry 3 16 6 4 <1 Asters,mainly large-leafedaster . 15 <2 6 13 2 Redandmountainmaple 5 5 9 2 15 Raspberry.and B!ackberry <1 13 4 Clover 4 11 Paperbirch 6 ' 8 <1 1 Fireweed 8 Willow 6 5 3 7 2 Spreadingdogbane 6 1 Spottedjewelweed 6 Buckbean 5 •Wildsweetpea 3 4 Grassesandsedges 1 3 1 2 Juneberry 1 <2 4 ' Lillies,mainly •Clinton'slily 3 2 <1 AmericanVetch 1 3 BristlySarasparilla <1 3 Fringedbindweed 3 Goldenrod 2 <1 2 2 Rose 1 2 BraCkenfern 1 2 <1 Dogwood 2 1 Thimbleberry 2 Violet 2 Blueberry 2 Basswood 2 Unidentifiedleaves 14 Miscellaneousplants 21 Numbeofr feedingsites or rumensexamined 26 8 3 Numberof bites 7,964 570 , _Plantwshosuesebydeerdidnorteac2hpercentofthedietinanystudyinAugustincludedHawkweed,commdandelion oroun,gcinquefoil,h pyrola,pearly everlasting,blacsknakeromeot,adowrueindi, ahnemp,bluebeswll, eeffebarrrn,esntrawberry,arrowwood,thistle,bedstramint,w, wildlettuchedge, binde weed wildsarsaparillhona,eysucgkle,reenalder,andwhitespruce. 3FromIrwin(1974).Datwea reobtainbyedexaminin52fgeedinsgitesintheLittleSiouxBurnareainSt.LouiCsounty,northeasternMinnesotaduringApril- December1973:Deerdensitywas4to6persquakilomre eter.(P.D.Karns1979,, personcaommunil cation). '=FromKohn(1970)..Dawertaoebtainedbyexaminin31fgeedinsgiteands 9rumeninsItascaCounty,north-centrMinnesotal duringa, June-Augu1968-196st 9. DeerpopulationdatareasforPierce(1975)(abovKohne). didnotmentionbyname11browseand20forbtaxaeachofwhicahccountedfornotmorethan2 percentofthedietinanymonthofstudy. 8FrOmBauer(1977).Datwaerobte ainbyedobservaofti3otname,harnesdeersedina10-acmreaturasepestnandandinanadjacent10-ac1-year-re aspenold CiearcudutringJune-Augu1976sint DickinsoCountyn inMichigan'Uppers PeninsulaDeer.denswityas11persquarekilometeranddeclining(L.F.Verme,1979 personcaomml unication). 7Mooty.(unpublished)Datwea r.oebtainedbyexamininfeegdinsiteg insItascCoa unty,north-centrMinnal esoduringta 1968-1971.Forasummaryofthesedata by2-monthintervalses,eMooty(1976).SomeofthedatuasedbyMootywerecontributedbyKohn(1970)andWaddell(1973).Deerdensitywas4to8persquare kilometeranddeclining,accordingannuto sapl rinpgelletcountsconducbytedtheMinnesotDepartmenta ofNaturalResources. .16 Table 9.--Plants eaten by white-tailed deer in north- Table lO.--Plants eaten by white-tailed deer in north- " ern Minnesota in September ern Minnesota in October . . (In estimated percentages of the diet) _ (In estimated percentages of the diet) _ Forested Burned Forested Burned feedingsitese feedingsites3 feedingsitese feedingsitesa Redmaple 49 Bush-honeysuckle 24 36 Grassesandsedges 29 5 Asters,mainly Bush-honeysuckle 13 I largelea, fedaster 33 3 Beakedandamericanhazel 1 12 Pincherry 20 Clover 7 11 Fringedbindweed 11 Aster,mainlylarge- Clover 10 9 leafedaster 11 Mountainmaple ,9 4 Bunchberry 8 2 WilloW 6 2 Goldenrod 8 Bristlysarsaparilla 6 Paperbirch 1 7 _Clint0n's lily 3 <2 Wintergreen 5 2 Goldenrod 3 , Quakingaspen 5 2 Brackenfern 3 Jackpine 4 Redmaple 2 <2 Juneberry 4 Rose " 2 <2 Raspberryand blackberry 2 2 Miscellaneousplants 9 Strawberry 2 2 Numberof feedingsites 12 5 Commonthistle 2 Numberof bitestallied 2,932 490 Hedgebindweed 2 Miscellaneousplants 8 1piantswhoseusebydeerdidnotreach2percentineitherstudyinAugust . includedra.spberry,blackberry,grass,sedge,bunchberry,strawberry, Numberof feeding thistle,beakedhazel,Americanhazel,quakingaspen,andarrowwood, sitesexamined 13 6 3FromIrwin(1974).Datawereobtainedbyexamining52feedingsitesin Numberof bitestallied 5,483 480 theLittleSiouxBurnareainSt.LouisCounty,northeasternMinnesotaduring April-December1973.Deerdensitywas4to6 persquarekilometer(P.D. _Plantswhoseusebydeerdidnotreach2percentofthedieint eitherstudy KarnS,1979personalcommunication), inOctoberincludedrose,willow,alder,andfireweed. eFromWaddell(1973).Datawereobtainedbyexamining41feedingsites 3FromIrwin(1974).Datawereobtainedbyexamining52feedingsitesin duringAugust-October1970-197in1ItascaCounty,north-centralMinnesota. theLittleSiouxBurnareainSt.LouisCounty,northeasternMinnesotaduring •Waddelldidnotmentionbyname56planttaxa,eachofwhichaccountedfor April-December1973.Deerdensitywas4to4 persquarekilometer(P.D. notmorethan2 percentofthedietinanyofthe3 monthsofstudy.Deer Karns,1979,personalcommunication). densitywasfourtoeightsquarekilometeraccordingtocensusdatacollected 8FromWaddell(1973).Datawereobtainedbyexamining41feedingsites bytheMinnesotaDepartmentofNaturalResources. duringAugust-October1970-1971inItascaCounty,north-centralMinnesota. Waddelldidnotmentionbyname56planttaxa,eachofwhichaccountedfor notmorethan2 percentofthedietinanyofthe3 monthsofstudy.Deer . densitywas4-8persquarekilometeraccordingtocensusdatacollectedby , • theMinnesotaDepartmentofNaturalResources.

17 Table ol l.---Plants eaten by white-tailed deer in the northern Great Lakes Region in November .

(I_nestimated percentages of the diet as determined from rumen analyses).

NortheastMinnesota NorthernWisconsin 1967-19691 19362 1943a Large-leafaster 25 Easternhemlock 20 Quakingaspenandbalsampoplar 4 17 11 White-cedar 9 15 Balsamfir 13 12 Grasses , 11 3 <2 Willow 1 11 Birch .10 <2 Bdnchberry 8 2 <2 Redosierdogwood , 8 Pine(mainlyjackpine) 1 7 7 GreenandSpeckledalder 3 <1 5 Labradortea 5 Apple 5 Boglaurel 4 Currantor gooseberry 4 Wintergreen 4 • BeakedandAmericanhazel 1 <1 3 Twinflower 3 <1 Rose 3 Maple 2 2 Mushrooms,lichens,andmoss 2 <2 Commonthistle 2 Woodfern 2 Wild sweetpea 2 Compositefamily 2 Unidentifiedtwigs 12 12 1 Numberof rumensexamined 14 21 387 1FromWetzel.(1972).DataforNovemberwereobtainedbyexamining14rumensfromdeerkilledbyhuntersinSt.LouisandLakeCountiesinnortheastern Minnesotadurin1967-1g 969De.edensityr was4to6persquarekilometer(P.D.Karns,1978,personalcommunication),andthepopulationwasdeclining(Mech • " andKam1s977;Rogersetal.1980).Wetzeomil ttedanddidnotname25itemsthaht efoundonlyintraceamountdurings the3yearsheanalyzerumed ns. 2FromAldousandSmith(1938).Datfora Novembweerer obtainedbyexamining21rumensfromdeerkilledbyhunteinrsnortheasternMinnesotain1936Items. ' whicAh ldouands smithfoundtocomprisleessthan1percentofthedietincludeblad cspruk ce,basswood,clubmoss,raspberry,juniper,goldenrod,clover, grape,grapefern,hardhack,ironwood,Ioosestrife,sheepsorrel,elm,cherry,andoak. 3FromDahlbergandGuettinger(1956).DataforNovemberwereobtainedbyexaminin387g rumensfromdeerkilledbyhuntersinnorthernWisconsinin1943. , Fiftyplantspeciesin,additiontothoselistedintable11,werelistedbyDahlbergandGuettingerascontributingtraceamountsto thediet.

'18 Table 12._Plants eaten by white-tailed deer in northern Minnesota in December

(In estimated percentages of the diet as determined through examination of feeding sites) _

Forestedfeedingsites Burnedfeedingsites Northeast2 Northcentral3 Northeast4 Dogwood 44 16 Redmaple 10 <1 40 Sweetfern 23 BeakedandAmericanhazel 13 22 Jackpine _ 2 19 Mountainmaple 15 <1 2 Blueberry ' 15 Wintergreen 12 . Labradotrea <1 9 ' Juneberry 7 6 2 Greenalder , 6 Bunchberry 4 Bush-honeysuckle 3 2 Willow <1 3 Quakingaspen 2 <1 <2 Sedge 2 Miscellaneous,plants 5 Numberof feedingsitesexamined 4 12 10 Numberof -bitestallied 2,112 1,861 782 I _Plantswhoseusebydeedidr notreach2percentinanyofthestudiesinDecemberincludedthistles,trailingarbutus,clubmoss,strawberry,honeysuckle,red t raspberry,balsampoplar,paperbirch,cherry,staghornsumac,rose,redoak,blackash,andwhitepine. 2FromWetzel(1972).Foradditionaldetailsofthatstudy,seefootnote3 oftable1. :3Mooty(unpublished).Foradditionaldetailsofthatstudy,seefootnote7, table7. 'i 4FromIrwin(1974).Foradditionaldetailsofthatstudy,seefootnote3, table6.

, COMMON AND SCIENTIFIC NAMES OF PLANTS , EATEN BY DEER IN THE i| UPPER GREAT LAKES REGION

I , Commonname Scientific name Seasoneaten_ ' TREES: Americanbeech Fagusgrandifolia Americanelm Ulmusamericana Americanlarchor tamarack Larixlaricinia , Apple Pyrus spp. F Ash Fraxinusspp. Aspen Populusspp. Su, F,W Balsamfir Abiesbalsamea F,W Balsampoplar Popu/usba/samifera F Basswood Tiliaamericana W

19 Birch Betulaspp. F,W Bitternuthickory Caryacordiformis Blackash Fraxinusnigra W Blackcherry Prunusserotina W Blackspruce Piceamariana Bluebeech Carpinuscaroliniana Box elder Acer negundo

.. Bur oak Quercusmacrocarpa W Butternut Juglanscinema Easternhemlock Tsugacanadensis F Elm Ulmusspp. Greenash Fraxinuspennsylvanica W Hornbeamor Ironwood Ostryavirginiana Jack pine , Pinusbanksiana F,W Large-toothedaspen Populusgrandidentata Su, W . Maple Acer spp. Su, W ' Mountain-ash Sorbusamericana W Oak , Quercusspp. Su, W Paperbirch Betulapapyrifera Su, F,W . Pine Pinusspp. F,W . Quakingaspen Populustremuloides Su, F ' Redmaple Acer rubrum Su, F,W Redoak Quercusrubra Redpine Pinusresinosa Silver maple Acer saccharinum , Sugarmaple Acer saccarum Swampbirch Betulapumila White-cedar Thujaoccidentalis Sp, F,W White pine Pinusstrobus W White spruce Piceaglauca Wild crab Pyrusangustifolia Yellowbirch Betulalutea

SHRUBSANDVINES;

Alder Alnus spp. Sp, F,W Alderleafbuckthorn Rhamnusalnifolia , Alternate-leaveddogwood Comusaltemifolia W ' , • • Americanhazel Corylusamericana Su, F, " Arrowwood Viburnumspp. , Beakedhazel Coryluscomuta Sp, Su, F,W Bearberry Arctostaphylosuva-ursi Blackberryand raspberry Rubusspp. Su , Black-haw Viburnumlentago . Blueberry Vacciniumspp. Sp, W Bogbirch Betulapumila Bog laurel Kalmiapolifolia .. Bog rosemary Andromedaglaucophylla Buffaloberry Sheperdiaargentea . Bush-honeysuckle OiervillaIonicera Sp, Su, F •- Canadianfly honeysuckle Loniceracanadensis • Cherry Prunusspp. Su Chokeberry Pyrusmelanocarpa W Chokecherry Prunus virginiana Su Commonelder Sambucuscanadensis .

20 Creepingsnowberry Gaultheriahispidu/a " Currant Ribesspp. - Swampcurrant Ribes/acustre Dogwood Comusspp. Sp,Su, W Downyarrow-wood Viburnumrafinesquianum W Easternblackberry Rubusallegheniensis Elderberry Sambucusspp. Gooseberry Ribesspp. Graydogwood Comusracemosa W Greenbriar Smilaxspp. Greenalder Alnuscrispa Sp, W .Hairyhoneysuckle Lonicerahirsuta Hardback Spiraeatomentosa Hawthorn , Crataegusspp. Hazel Corylusspp. Sp, Su,F,W Highbushblueberry . Vacciniumcorymbosum , _ Highbushcranberry Viburnumtri/obum W Honeysuckle , Loniceraspp. Su Juneberry Amelanchiesrpp. Sp, Su, W Juniper Juniperusspp. ' Labradotear Ledumgroenlandicum F,W Latelowblueberry Vacciniumangustifolium Sp Leatherleaf Chamaedaphnecalyculata Leatherwoodor moosewood Dircapalustris Lilac Syringavulgaris Meadowrose Rosablanda ' Meadowsweet Spiraealatifolia,S. alba Mountain-holly Nemopanthusmucronatus Mountainmaple Acerspicatum Su, W NewJerseytea Ceanothuasmericanus Ninebark Physocarpusopulifolius Northernhollyor winterberry Ilexverticillata Northernyew Taxuscanadensis Pincherry Prunuspennsylvanica Su Pipsissewa Chimaphilaumbellata PoisonIvy Rhusradicans Prairiewillow Salixhumilis , Pricklyash Xanthoxylumamericanum . , Raspberry Rubusstrigosus Su Red-berrieedlder Sambucuspubens , Redosierdogwood Comusstolonifera Sp,Su,F,W Red-panicledogwood Comusracemosa Rose Rosaspp. , Round-leaveddogwood Comusrugosa W ' Smallcranberry Vacciniumoxycoccus Smoothclimbinghoneysuckle Loniceradioica Smoothsumac Rhusglabra Snowberry Symphoricarposa/bus Speckledalder Alnusrugosa Sp, W Sumac Rhusspp. W Swamphoneysuckle Loniceraoblongifolia Sweetfern Comptoniaperegrina W , Sweetgale Myricagale Thimbleberry Rubusparviflorus

° 21 .Thornapple Crataegusspp. " lrailing arbutus Epigearepens Twinflower Linnaeaborealis -Velvetleafblueberry Vacciniummyrtilloides Wild -grape Vitisriparia Wild plum Prunusamericanus Wild raisin Viburnumcassinoides Willow Salix spp. Sp, Su, F,W ' " Wintergreen Gaultheriaprocumbens F,W " Witch hazel Hamamelisvirginiana -Woodbineor virginia creeper Parthenocissusinserta HERBACEOUSPLANTS: ' Medicagosativa Americanvetch Viciaamericana . Arrowhead Sagittarialatifolia Aster Aster spp. Su Barrenstrawberry , Waldsteiniafragaroides Su Bedstraw Galiumspp. . Blackmedic Medicagolupulina .- Blacksnakeroot Sariculamarilandica Bladdercampion Silenecucubalus Bladderwort Utriculariaspp. Bluebell Mertensiapaniculata Bristlysarsaparilla Araliahispida Su , . Buckbean Menyanthestrifoliata Su Buckwheat Fagopyrumspp. Bunchberry Comuscanadensis F Bur-reed Sparganiumspp. Clinton'slily Clintoniaborealis Clover Trifoliumand Melilotus Su, F • Commondandelion Taraxacumofficinale Commongeranium Geraniumbicknellii Commontwisted-stalk Streptopusroseus Compositefamily Compositae Coontail Ceratophyllumdemersum Corn Zeamays .. ' Cowparsnip Heracleumlanatum • .- ' Dock Rumexspp. - EarlySweetpea Lathyrusochroleucus Su ' Falselily-of-the-valley Maianthemumcanadense FalseSolomon'sseal Smilacinatrifolia Filamentousalgae Spirogyraspp. , Fireweed Epilobiumangustifolium Su Flat-topwhite aster Aster umbellatus Forkingcatchfly Silenedichotoma • Fringedbindweedor false buckwheat Polygonumcilinode Su .Garden.pea , Pisumsativum Goldenrod Solidagospp. Sp, Su, F • 6oldthread Coptisgroenlandica : 6oosefoot Chenopodiumspp. Graminoids Gramineaeand Cyperaceae Grasses Gramineae Sp, Su, F Harebell Campanularotundifolia !

I.

I

. Hawkweed Hieraciumspp.

. , - Hedgebindweed Convolvulussepium Hempor marijuana Cannabissativa Ho_seweed Erigeroncanadensis IndianHemp Apocynumcannabinum Kidneybean Phaseolusspp. Lady'sthumb Polygonumpersicaria . Large-leafaster Aster macrophyllus Su, F Lily family. Liliaceae Sp, Su Loosestrife Lysimachiaspp. , Marshcinquefoil Potentillapalustris Marshmarigold Calthapalustris Sp Milkweed Asclepiasspp. I _ _ Night-floweringcatchfly Silenenoctiflora NorthernSt. John's-wort Hypericumboreale . _. Paletouch-me-not Impatienspallida ' Pearlyeverlasting Anaphalismargaritaceae Pigweed . Amaranthusspp. , Pitcherplant Sarraceniapurpurea , • Purplewatershield Braseniaschreberi .- Pyrola Pyrolaspp. Ragweed Ambrosiaspp. Ribbonleafpondweed Potamogetonepihydrus Roughcinquefoil Potenti//anorvegica Su Scarletcolumbine Aquilegiacanadensis ,- Sedges Cyperaceae, Sp, Su, F includingCarexspp. , Sheepsorrel Rumexacetosella , Showysmartweed Polygonumamphibium Smoothaster Aster laevis Solomon'sseal Polygonatumspp. • Glycinemax • Spikerush Eleocharisspp. Spottedjewelweed Impatienscapensis Su Spreadingdogbane Apocynumandrosaemifolium Su Strawberry Fragariaspp. Su Sunflower Helianthusspp. , . ' Sweetclover Melilotus spp. ,. ' Sweetpea Lathyrusspp. Su Sweetwater-parsnip Siumsauve ' Thistle Cirsiumspp. . Three-waysedge Oulichiumspp. Violet Violaspp. Su ;_ ' " . Waterhorsetail Equisetumfluviatile ' Watermilfoil Myriophyllumspp. Su Wheat Triticumaestivum , " Whiteclover Trifoliumrepens Wild carrotor QueenAnne'slace Oaucuscarota Wild lettuce Lactucacanadensis • Wild rice , Zizaniaaquatica • Wild sarsaparilla Aralianudicaulis Su Wild sweetpea Lasthyrusvenosus Su Woodanemone Anemonequinquefolia Yarrow Achilleamillifolium Yellowbellwort Uvulariagrandiflora

. , " Yellowtrout lily Erthroniumamericanum FUNGI: Bracketfungi Daedaliaspp. Lenzitesspp. Polyporusspp. .. Schizophyllumspp. ' Mushrooms- Su

LICHENS: Oldman'sbeard Usneaspp. Arboreallichens2 . Main/yUsneaspp.and Evemiaspp. W

FERNSAND FERNALLIES: Brackenfern ' Pteridiumaquilinum Su , Clubmoss Lycopodiumspp. • Grapefern Botrychiumspp. Shieldfern Dryopterisspp. Woodfern Dryopterisspp. Mosses Unidentifiedmosses

t Pigeonwheatmoss Polytrichumspp.

1Seasonofuseisdesignatedonlyforthosespeciesfoundtocomprise5 percentormoreofthedietinatleastone studyinthe givenseason.Sp = April-May,Su = June-September,F = October-November,W = December- March. 2Alectoriaspp.wasreportedaseateninonestudybutlaterwasfoundtobeamisidentificationofUsneaspp.and Everniaspp.lichens.

o 24 II_U.S. GOVERNMENPRT INTING OFFICE: 1981-767-167/3 REG.#6