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White-Tailed in the Southern. Forest Habitat

ProceedZzg~of n Symposiam at Nacogdoches, Texas March 25-26, 1969

Southern Forest Experiment Station Forest Service, U. S. Department of Agriculture in coope~ationwith the Forest Committee of the southeaster^^ Section of the Society and the School of Forestry, Stephen F. Austin State University White-tailed deer were once nearly exterminated in the South. In the past several decades, however, improvements in game management and changes in land use patterns have enabled populations to recover. The deer today are more numerous than at any time since white men settled the country. Deer are highly valued by the public. They are heavily hunted by sportsmen, admired by nature enthusiasts, and desired by some land- owners as a possible source of additional income. Yet to many persons the deer are less than a blessing. They some- times severely damage farm crops and young trees. They are possible transmitters of disease. They often compete with other wild or for food. They create difficulties in resource allocation and management. These problems become more complicated as human popu- lations expand and as for land use intensifies. The potential values of deer, however, outweigh the undesirable characteristics, but enlightened management is necessary if these values are to be achieved fully. The symposium was organized to consolidate known information and to offer a means of expressing new ideas and philosophy pertinent to management of white-tailed deer in southern forests. Persons from Federal agencies, State conservation departments, universities, and private industry were asked to contribute their knowl- edge and viewpoints. Speakers emphasized the background, characteris- tics, and management of deer and their habitat. Of special importance were papers concerning the outlook for deer as influenced by sociological, economic, and political factors. It is believed that the information presented here represents the most complete compendium now available on the southern white-tailed deer.

Lowell K. Halls WILDLIFEHABITAT AND SILVICULTURELABORATORY SOUTHERNFOREST EXPERIMENT STATION NACOGDOCHES,TEXAS Contents Page

History of deer and their habitat in the South- John D. Newsom

Population dynamics of whlte-talled deer-

Walter V. Robertson . 5

Foods and feeding hablts of white-tailed deer- Daniel W. Lay 8

Physiology and nutrition of deer in southern upland forests- Henry L. Short 14

Seasonal changes in movements of white-tailed deer- Robert L. Downing, Burd S. McGinnes, Ralph L. Petcher, and Joshua L. Sandt 19

Deer In North Carolma and other southeastern States- Frank B. Barzck 25

Some considerations for diseases and parasites of white-tailed deer in the southeastern United States- Frank A. Hayes and Annie K. Prestwoocl 32

Critical factors In habitat appra~sal- William D. Zeedyk

Deer habitat quality of major forest types in the South- J. J. Stransky 42

Agricultural clearings as sources of supplemental food and habitat diversity for white-tailed deer- Joseph S. Larson 46

Philosophy of deerherd management- Jack A. Crockford 51

Hunting methods, limits, and regulations- Dean A. Murphy 54

Managed hunts by State agencies- A. Gordon Spratt 59

Effects of timber harvest and regeneration on deerfood and cover- V. E. Carter and S A. Dow 62 How size and distribution of cutting units affect food and cover of deer- Burd S. McGinnes ...... 66

Short and long rotations in relation to deer management in southern forests- Phil D. Goodrum ...... 71

Timber stand density influences food and cover- Robert M. Blair ...... 74

Effects of hardwood control on food and cover- E. B. Chamberlain, Jr......

The effects of prescribed burning on deerfood and cover-

Paul A. Shrauder and Howard A. Miller ...... _.. 81

The goals of the southern National forests in white-tailed deer management- Donald D. Strode and Wayne J. Cloward ...... 85

The goals of State conservation agencies in deerherd management- Frank P. Nelson ...... 88

The goals of private forest holdings in deer management- Raymond D. Moody ...... 90

Prevention and control of damage to trees- D. C. Denton, E. H. Hodil, and D. H. Arner ...... 93

Measuring habitat productivity- Richard F. Harlow and James D. WheEan ...... 98

Evaluating food use-new methods and techniques- H. S. Crawford ...... 109

Problems in censusing the while-tailed deer- James H. Jenkins and R. Larry Marchinton ...... 115

The use of models in resource management- Don W. Hayne ...... 119

Sociological and economic considerations in management of white- tailed deer- E. L. Cheatum, Lonnie L. Williamson, and A. Sydney Johnson .... 123

Regulatory legislation and public attitude on white-tailed deer man- agement- Leslie L. Glasgow ...... 127 History of Deer and Their Habitat in the South John D. Newsom ' Louisiana Cooperative Wildlife Research Unit Bureau of Sport Fisheries and Wildlife U. S. Department of the Interior Baton Rouge, Louisiana

The history of deer and their habitat in the South bama) was 250,000 pounds of hides The annual export is separated into four periods: Precolonial, Exploi- from Charles Town i South Carolina) between 1739 and tation, Recovery, and Today. 1762 was from 131,000 pounds to 355,000 pounds and In the Precolonial Period deer populations and from Georgia between 1765 and 1773 was more than their habitat remained relatively stable over long 200,000 pounds " George Barnard is reported to have periods of time even thoz~glrdeer were widely used shipped not less than 75,000 deerskins from near the for food, clothing, and tools by the Indians. present site of Waco, Texas, between 1844 and 1853, and at his death in 1883 was one of the wealthiest men Exploitation through settlement and clearing of in Texas, having gained h~swealth from trade in deer land, market hunting for meat and hides and hunt- and other sklns (Strecker 1927) ing without restrictions throughout the year reduced deer populations to an alltime low in the South Most of these records refer to the middle and late 18th aboz~t1920. century. I have not been able to establish the precise Recovery was initiated by the establishment of period during which a general decline in deer popula- National Forest Preserves in the early 1900's. It tions occurred throughout the South. It is assumed that progressed through the establishment and enforce- the initial decline in numbers occurred first in the east- ment of hunting regulations, reversion of farmland ern seaboard States and progressed westward with colon- to forests and Federal Aid to Wildlife Restoration ization and subsequent exploitation. This pattern of restocking programs. decline is probably reflected by the dates when game laws were first established by the various States: Vir- Currently, approximately 90 percent of the land ginia, 1699; Maryland, 1730; North Carolina, 1730; South area in the South is open to deer hunting, and the Carolina, 1755; Georgia, 1790; Mississippi, 1803; Ala- annual kill exceeds 300,000 deer. bama, 1822; Florida, 1828; Louisiana, 1857; Texas, 1860; Kentucky, 1861; Tennessee, 1870; and Arkansas, 1875. ACKNOWLEDGMENT The South was obviously a land of plenty, insofar as I am indebted to the following persons for providing decr were concerned, until the middle of the 19th cen- some of the information used in this paper: Charles tury. Kelley, Alabama; Gene Rush, Arkansas: Gordon Spratt, The chronology of evolution of deer and their habitat Florida; Hubert Handy, Georgia; Fred Hardy, Kentucky; in the South can best be described in stages. For my Bill Turcotte, Mississippi; Frank Barick, North Carolina; discussion, I will utilize the following: Stage I-The Frank Nelson, South Carolina; Roy Anderson, Tennes- Precolonial Period; Stage 11-Exploitatiol?; Stage 111- see; and Dick Cross, Virginia. Recovery; and Stage IV--Today. The white-tailed deer (Odocoileus ?~irginianus)is the big game animal in the Southern United States. . .. THE PRECOLONIAL PERIOD Writings of early travelers through the South are replete with accounts of the abundance of deer iDu- When we think of wildlife habitat in the South at the Pratz 1774; Bartram 1792). The dependence of early time of discovery and initial settlement by white men, settlers on the white-tailed deer for food and clothing we are prone to visualize a pristine wilderness. Obvi- is well known. During certain times the settlers depend- ously, most of the forested areas of the South were virgin ed heavily on the meat and skins of deer as itenis of trade. wilderness, characterized by mature forests with rc!a- Through some of these records we are able to obtain lively clean floors and little understory of value to deer. information on the relative abulidance of deer during I-Iou7evcr, even at that early date in our history the Indi- the period of coloinzation of the Soutll For instance ails were practicing habitat maiiageinent of a somewhat Young (1956) reports that 2,601,152 pound5 of deerskin crude form by maintailling rather large openings in the from about 600,000 deer were shipped from Savannah forest by the use oi' fire (Prunty 1965 j. Deer habitat (Georgia ) from 1755 to 1773, and in 1 year, 1753, 30,000 in this stage must have remained relatively stable over deerskins were exported from the colony 01 North Caro- long periods of time subject only to the effects of natural lina Harlow and Jones ( 1965 ) report as follouri "The ( lightning ) fires and those occasionally set by Indians, trade was llghter from St Augustine ( Florida ), u here and other natural phenomena. In the absence of authen- in 1771 only 4,000 pounds of hides were traded than tic records. I would guess that the three broad forest from Pensacola t Florida 1 where during the same year types would have rated as deer habilat in the following the conibiiied export from Pensacola and Mobilc (Ala- order : pine hardwoods, bottom-laud hardwoods and -- 'This papcxr is ,I contribution of the Louisiana Cooper,lti\e longleaf pine lands. None of these habitat types sliould Wlldlifc Rescdrch Unlt, Louisiana State Um\cisity, 1,oulriana have been as productive as they are today. Wild Lifc and Fisheries Cominlsslon, The Wildlife Mandge- ment Institute and U S Bureau of Sport Flshcries and Wild- The usc of decr by the Indians for food, clothing, and life Coopelating tools has been well authenticated. From this we can safely assume that deer were widespread and fairly abun- in 1916. National wildlife preserves were establisk dant throughout the South during this period. on a part of each of these areas. In the report of Leopc et al. (1947) only two areas in the South are listed EXPLOITATION having problems with overpopulations of deer-t Ozark National Forest (syLa&ore District) and 1 The American Indian was apparently a much wiser Pisgah. The establishment of the bulk of the southe conservationist, or game manager, than the white man National forests in the midthirties gave an additiot who succeeded him. In general, the Indian took from boost to the marginal deer populations. the land only that which was needed to sustain him. During the great depression of the late 1920's a There are exceptions, but the mere fact that the white 1930's, there is no evidence to indicate that substant man found such an abundance of natural resources when recovery was made by southern deerherds. Howev~ he arrived out this contention. some progress was made in restoration of habitat. . Exploitation of southern deer and their habitat must mentioned previously, the bulk of the National fore: have started soon after the first settler set foot upon of the South were established in the midthirties. In 19 southern soil. The settlers immedistely started to clear the Civilian Conservation Corps was established and the forests so they could grow their crops, and unlike 1942, 2.5 billion trees had been planted on denud the Indians, sought wildlife as a medium of trade in addi- forest land; the gradual exodus of the human populati~ tion to its use for food and clothing. The white-tailed from farms to industrial jobs in cities had begun a1 deer was one of the most sought after . farmland started the reversion back to forest land. E The unrestricted harvest of wildlife continued in the tween 1945 and 1953, 10 million acres of farmland South until the mid 19th century. At this time the the South reverted to forest (McGuire and Dickerm, enactment of game laws became a part of the deer man- 1958). agement picture in the South. But in spite of this, ex- The passage of the Federal Aid to Wildlife Restoratic ploitation of the species continued. It appears that rela- Act (more familiarly known as the Pittman-Robertsc tively unrestricted deer hunting continued in the South Act) in 1936 provided the key to the restoration of wil until about 1915-1920. In all cases restrictive laws were life throughout the United States. This act of Congre poorly enforced, or not at all. ushered in the era of in this cou According to the United States Census of Agriculture, try. Some Southern States initiated deer restockii there are approximately 332 million acres of land in the programs in the late 1930's utilizing Pittman-Robertsc 12 States covered in this discussion. By 1850 there were funds. 496,892 farms involving a total of 162,013,497 acres of It was during this period that the remaining de land in agricultural production. By 1920 there were populations in the South suffered several severe sr 2,481,101 farms operating 193,629,309 acres of land. The backs. This Mississippi River flood of 1927 virtual production of both softwood and hardwood lumber shows wiped out the deer in Louisiana's Delta hardwoc similar trends. Total lumber production in 1869 was swamps: the fever tick eradication program whi~ 1,624,843,000 board feet; in 1920, it was 15,086,372,000 started in Florida in 1937 and the overpopulation a1 board feet (Steer 1948). At this time the agrarian cul- subsequent die-off of deer on the Pisgah in North Car ture of the South was approaching its peak, and it was lina and the Sylamore in Arkansas set back the recove. characterized by high rural and low urban human popu- of deerherds in these States to a significant extent. lations with generally low financial status. With the advent of World War I1 forces were aga This unrelenting pressure from the white man had a exerted on deer and their habitat. To meet the deman, telling effect on the white-tailed deer in the South. In of war in 1942, lumber production in the South elimbc many areas deer were completely exterminated by the to the then alltime high of 18,380,706,000 board fec practice of hunting with all year long and killing exceeding the production of 1935 by almost 8 billic all deer regardless of sex or age. By about 1920, the board feet (Steer 1948). Throughout the South lar; white-tail reached the alltime low point in its population tracts of land, most of which were suitable deer habit: (Barick 1951 ). It had been reduced to scattered remnant were acquired by the U. S. Department of Defense f herds that were largely confined to the most remote military training purposes. Many of these areas we and inaccessible habitat types-hardwood bottom-land fenced and protected and ultimately harbored large de swamps and mountainous terrain. populations. A large segment of the hunting public w called into military service, but because of meat sho~ ages during the war years of 1941-1945 heavy deman RECOVERY were placed on game populations to supplement me In 1891 when the first forest reserves were established, rations. Approximately 54 million pounds of dressc the feeling of concern for forest resources in the country meat, principally deer, were taken from American fore; became evident. This feeling grew and resulted in the in 1942 (Young 1956). establishment of the U. S. Forest Service in 1905 and Although much of the foundation for recovery of de eventually to the establishment of the National Forest and their habitat in the South was laid between 19: System. These National forests have played a key role and 1940, it was not until after the end of World War in the recovery of southern deerherds. Some of the that real progress was made. With the return of mc earliest National forests established in the South were: from military service, a work force became availab: Ozark in north-central Arkansas in 1908; Ocala in north Federal Aid to Wlldlife Restoration funds had bu Florida in 1908 and the Pisgah in western North Carolina up during the war years and adequate financing w available to the States, and finally trained personnel Odocoileus virginianus virginianus, 0. v.macrourus, 0. v. became available to the States as a result of -the stepped- seminolus, 0. v. mcilhennyi, 0. v. osceola, and 0. v. up program of training in many State colleges and uni- clauium. During these restocking efforts, there was versities. The demand for outdoor recreational oppor- considerable mixing of the existing resident races of tunity was growing yearly. A general awareness of the deer plus the introduction of at least two new races: need for a broad-base wildlife conservation effort became 0. v. borealis and 0. 1,. texanus (table 1). The overall evident by the increase in the number of organized effects of this mixing is unknown, but at this time it is sportsmens groups and the reorganization of several obvious that reproductive capacities are adequate for State wildlife agencies. existing deer habitat. By 1949 every State in the South had an active pro- The history of recovery of southern deerherds can gram of deer restoration with the exception of South best be treated by the data presented in table I. The Carolina whose trapping and transplanting program was deer population increased from about 303,500 in the mid- initiated in 1951 (Barick 1951). Most of the forests forties to about 2,405,000 at the present time and the had been cut over for several years and young second kill increased from about 60,133 in 1950 to 274,184 in growth shrubby vegetation provided excellent browse 1967 in the I1 Southern States on which data are avail- conditions for deer. All States had initiated programs able. of acquiring land for the development of wildlife man- Concerted efforts toward reestablishment of the agement areas. Between 1938 and 1967, 22,686 white- South's deerherds were virtually completed by 1960. tailed deer were stocked in the 12 Southern States in Kentucky was the only State with an active deer re- areas where protection was provided and hunting was stocking program in 1967 when they restocked 458 ani- not allowed for a period of 5 years after stocking. mals. Deer hunting was once again a sport which could Originally there were six subspecies of the white- be enjoyed by the average person, and the number of tailed deer in 12 Southern States covered in this report: deer hunters was increasing annually.

Table 1.-History and status of deer restoration in the South Percent of Source of State open to State State open t deer hunting prior Deer kill to restocking

Alabama ' 1945 (1925) Alabama Wisconsin N. Carolina Arkansas 1945 (1940) Arkansas Wisconsin Florida 1948 (1938) Florida Wisconsin Texas Georgia 1950 (1928) Georgia Wisconsin Texas Kentucky 1946 Kentucky Wisconsin Louisiana 1948 (1920's) Louisiana Wisconsin Texas Mississippi 1944 (1937) N. Carolina Mexico North Carolina 1944 (1937) Wisconsin N. Carolina South Carolina 1951 S. Carolina

Tennessee 1947 Wisconsin Tennessee Virginia 1938 Wisconsin Pennsylvania Total ' Year when major effort was begun. - Year when initial restocking started. There were some problems developing, too Deer The unrestricted use of dogs for hunting deer and t huntlng in the South had historically been "bucks only " abundance of free-ranging dogs in many areas of t Early proponents of deer restora:ion did an excellent South continue t.o be prime limiting factors on deer pc job of selllng the gcilcral public on protecting doe deer ulations. This is an area that deserves major attenti I an1 sometimes prone to think that they did a much of responsible deer managers and game dt~parlmc better lob than was necessary admii~istrators. As deer populatioils continued to increase, reports of Deer habitat and dcer ~~opulationsIn the South timber reproductioil and agricultural crop damage by presently 111 good condition I helicve thdt continu1 deer increased. The constant threat of die-offs and public demand for good dcer hunting will make it I domestic livestock-deer disease relationships prompted pcrative that we continue to conduct our tlccr malr,l[ the States to organize the Southeastern Cooperative Deer inent activities to the fulle5t extent of our .tbilit~ Disease Study in 1947 to maintain survei!lance over Advances in deer management knou7-how art an ~b' disease conditions in southern deerhcrds. The time had lute necessity We must all work toward thdl end arrived when dcerherd reductions across the South were essential in order to maintain the progress that had been made. LITERATURE CITED In the late 1950's most of the States started working Barick, F. B. toward either sex deer seasons. Much progress has been 1951. Deer restoration in the southeastern Unit made, but all the problems have not been solved. Ken- States. Pap. presented at Southeast. Ass( tucky is the on!y Southcrn State that regularly permits Game and Fish Comm. Fifth Annu. Mt:c either-sex deer hunting by archers and gun hunters alike. ing. 18 pp. Several of the States (Arkansas, Kentucky, Louisiana, Bartram, W. and Tennessee ) allow lengthy either sex archery seasons. 1792. Travels tl~roughNorth and South Carolir And all Stat,es in the South now have the authority to Georgia, east and west Florida. 414 F establish either-sex deer seasons vested in the respective Facsimilie edition edited by M. Van Dorc State wildlife agencies and conduct restricted either-sex Dover Pub. deer hunts annually. But this aspect of deer management has not as yet been accepted to the degree that will allow Dupratz, L. most State wildlife agencies the latitude necessary to 1774. The history of Louisiana. 387 pp. (Engli properly manage southern deerhcrds. translation of earlier French volume^ Within the past 6 years another problem has developed. New Orleans: Pelican Press, Inc. The highly fertile river bottom lands, some of the most Harlow, R. F., and Jones, F. K. productive deer habitat, are being cleared for agricultural 1965. The white-tailed deer in Florida. F: crop production at an alarming rate. In Louisiana alone Game and Fresh Water Fish Comm. Tec 896,308 acres were cleared between 1962 and 1967. Some Bull. 9, 240 pp. of this land was supporting approximately 64 deer to the square mile. It appears that we must accept the fact Leopold, A., Sowls, L. K., and Spencer, D. L. that within the foreseeable future all of this highly fertile 1947. A survey of over-populated doer rang land will be placed in agricultural crop production, and in the United States. J. Wildlife Manak deerherds in these areas will be reduced to remnants of 11:162-177. their former ahundance. Deer habitat in the South will McGuire, J. R., and Dickerman, M. B. be restricted primarily to the upland pine and pine-hard- 1958. Forest land and timber. In Timber r woods areas, which cannot support nearly as many deer sources Tor America's future, pp. 113-14 as the bottom-land hardwoods are now supporting. USDA Forest Resource Rep. 14. Prunty, M. C. TODAY 1965. Some geographic views of the role of fi in settlement processes in the South. Four In the South today the white-tailed dcer is hunted Annu. Tall Timbers Fire Ecol. Conf. Prc on approximately 90 percent of the total land area; 1965: 161-167. Tallahassee, Fla. 30 years ago only 30 percent of the land area was open to deer hunting, and some of this land supported no deer. Steer, H. B. The 12 Southern States covered in this report currently 1948. Lumber procluction in the United Stat operate 317 management areas and refuges totaling 1799-1946. USDA Misc. Pub. 669, 233 F 10,013,604 acres of land where the general public may Strecker. J. K. hunt white-tailed deer on an annual basis. The limited 1927. The trade in deer skins in early Texas. deer stocking currently done in the South, with few Mammalogy 8 ( 2) : 106-110. exceptions, is not being done in the name of restoration. There are approximately 200,926,000 acres of forest land Young, S. P. in the South, nearly all of which are suitable deer habi- 1956. The deer, the Indians and the Americ. tat. The annual kill of deer in the South undoubtedly pioneers. In The deer of North Amerit exceeds 300,000 animals. pp. 1-27. Harrisburg, Penn.: Stackpole C Population Dynamics of White-Tailed Deer Walter V. Robertson Biology Department Stephen F. Austin State University Nacogdoches, Texas

Pop~~lationdynamics is discussed in regard to Table I.-Reproductive potenlinl of rr szngle pnzr of deer tlzrougli 10 gcn- changes in total number and population structure erntions and the nrea wi~ici~could be populated at one deei per 40 acres with reference to their importance to deerherd man- agement. Some speculation regarding man's possible influence on genetic changes within the population is included. Adult Deer populat~ons,llke populations of any other specles, Adult Year1111g undergo constant change and In thls respect are dynamlc Juven~le Thc terms stability and balance when used In reference Adult to populations and conditions in nature actually mean Yearling a dynamic stability or dynamic balance. Fluctuations Juvcniie Adult in total number and changes occurring in population Yearling structure which affect total number are of primary con- Juvenile cern in deerherd management. Adult Yearling The rate at which a species could reproduce under ideal Juvenlle conditions and in the absence of mortality is referred to Adult as the reproductive potential, and any species has the Yearling capacity to overpopulate its habitat in a relatively short Juvenile time. In sexually reproducing forms the rapidity at Adult Yearling which this could be accomplished depends upon the age Juvenilc of sexual maturity, the number of offspring which can Adult be produced in a given unit of time, and the reproductive Yearling longevity of the species. Considering the reproductive Juven~ie potential of a single pair of white-tailed deer and follow- Adult Yearling ing this through 10 generations gives some idea of the Juvenile theoretical capacity of this organism to reproduce. The Adult assumption is made that adult does would produce two Yearling young per year, yearling does would produce one young Juvenlle per year, juveniles (fawns) would not reproduce, and that no mortality would occur. A sex ratio of half males and half females is assumed. With the exception of the absence of mortality the above assumptions are con- servative since it has been indicated by Scveringhaus POTENTIAL and Cheatum (1956) that under favorable conditions in ---- REALISTIC New York some of the female fawns breed at 6 to 8 months of age and occasionally produce twins. Instances of does remaining productive for 10 years and longer are known. Successive increases shown in table 1 are similar to those developed by Lcopold (1948) and can be obtained by mathematical equations (Kelker 1947; Eber- hardt 1969). Allowing one deer per 40 acres, which would require good deer range, one section of land (640 acres) could be populated to near capacity in only 4 years and almost 12 sections in only 10 years. Plotting this population growth curve shows a typical geometric progression (fig. 1). It becomes obvious imme- diately that the reproductive potential is largely theoreti- cal and if attained at all in nature, would be maintained for only a short time. In a study of deer released in a 12345678510 1,200-acre enclosure in Michigan O'Roke and Hamer- TIME IN YEARS strofn (1948) reported a population growth from four Figure 1.-Potential population growth curve in white- does and two bucks to 160 animals in 6 years. This rate tailed deer based on table 1 (solid line) and of increase closely approaches the reproductive potential the probably growth curve under natural of the species. Other records of rapid population increase conditions (broken line) where the carrying in large game are numerous. capacity is approximtltely 100 animals. Comparing the curve for the potential increase with the first year represents 50 percent of the populatio a hypothetical increase similar to that expected under whereas loss of the same number of fawns during ti natural conditions shows that the population tends to 10th year would represent less than 2 percent of tl "level-off" as the carrying capacity of the habitat is population. In the first instance population growl reached (fig. 1). The difference between the theoretical would be greatly affected, but in the second the effc and the realized population growth represents mortality would be negligible. It is likely that populatio~is fa and decreased natality. to become established in many suitable areas becau: Several factors within the total environment tend to of disturbance or reduction during the initial phase I bring populations down or prevent them from attaining population growth. A second important period is ti unusually high levels. Prime among these are physical point at which the optimum carrying capacity is a] factors such as temperature, rainfall, and soil fertility proached. At this level range conditions would be favo which affect directly the available food supply. There able and natality would be high. Thus, the large incr' are also several biological factors which tend to stabilize ment added to the pop~~lationduring the summer woul the population. Such factors as predation, disease, para- increase the numbers above safe levels for the wintr sitism, movement out of the area, reduced fertility and range. A severe winter could result in high mortalit natality, and competition for space, food, and cover are or an unusually mild winter causing little niortalit closely interrelated and tend to intensify as the popula- would compound the problem the following year. tion increases. The influence of these factors would vary It becomes obvious that habitat evaluation is a bas in different localities or within the same locality at dif- consideration for the wildlife biologist. Equally in ferent times. Only through experience and familiarity portant is a knowledge of the deer population. includin with a given population can the wildlife biologist ap- its age composition and sex ratios. The imllortance ( proach accuracy in evaluating the effect of these vari- this knowledge emphasizes the necd for refinement c ables upon the population. methods and techniques for population analyses whic The term "leveling-off" does not imply that the num- reflect the greatest accuracy with the least rcquiremer ber and composition of the population would remain of time. Since certain of the data can be collected on1 fixed. The actual numbers would fluctuate around an by close examination of the animals, it emphasizes th average. Numbers would increase above the average value of information gained from inspection of harveste during the period of birth when favorable conditions animals over an extended period of time. Records c exist and decline below this average during more harsh this sort pertaining to deer populations in southern fore: periods normally corresponding to late winter or seasons areas are noticeably scarce. of drought. During a series of favorable years the popu- Table 1 can be used to show how the removal of di- lation is likely to increase well above the ~iormalcarrying ferent age and sex classes will affect the population. Fc capacity of the habitat with resultant overbrowsing. instance at year 9 there would theoretically be 26 malc The habitat is severely damaged and a large portion of in the adult and yearling class. Removal of half c the population dies from starvation or from interactions these would reduce the total population to 110. Assumin between malnutrition and other decimating factors. The each of the remaining bucks could service as many E die-off normally reduces the population back to much four does, plus the fact thst many of these killed woul lower levels than the original carrying capacity. Usually have bred before being harvested, the population th several years are required for the recovery of the habitat following year would be reduced by only 13, giving and the population. The removal or drastic reduction of total of 176. On the other hand, removal of 13 adu. normal mortality factors such as predation or parasitism, does in year 9 would result in a population of 150 th if not offset by increased removal by other means, may following year. The effect of female removal in othc produce similar results. age classes can be easily determined. The conclusio The pattern of population growth and cycles indicate can be drawn that removal of buck deer, within limil that a sizable portion of an established population is of course, will not appreciably reduce the populatior likely to be lost due to natural causes. But, if a portion Normally, dangers of overpopulation can be averte of the population is removed prior to the decline phase only by removal of does. of the cycle then the natural mortality is reduced in Table 1 also shows that the ratio of young deer (yea] the approximate proportions of the segment removed. lings and fawns) to adults remains high as the populatio In a sense then, that portion of the population normally increascs. But failure of an appreciable increment ( lost represents a "surplus." Efforts in wildlife manage- young to survive would result in more adults tha ment are directed toward harvesting this surplus and young. In actual management practice then, if a censL maintaining population levels at or near the carrying of a population reveals this high ratio of young to aduli capacity of the range. the assumption is made that the herd is increasing an There arc two portions of the population growth that that the population is within the carrying capacity t seem especially important for the welfare of the deer- the range. Ilowever, if the reverse proportions are ol herd. The first is associated with the establishment level served it indicates reduced natality or low fawn surviv: and the other is the point in the curve which approaches which may mean overpopulation although it could 1: the carrying capacity of the habitat. When the popula- due to other factors. Data from other sources would 1: tion is low the actual increase in numbers is low and necdcd before an accurate assessment of the populatio any factor which reduced natality or increases mortality in relation to its range could be made. will have the effect of holding the population at very It may seem premature to talk of overpopulation her, low levels. In the example given in tablc 1 fawn loss since many southern forest arcas secm to be undcrpopl lated But overpopulatlon can occur at low densities lation. If hunters were willing to make the same eco- If the range can support one deer per 50 acres and there nomic investment to kill a small deer as compared to are twice this many per acre, then the range is over- a large one, then a greater number of small deer might populated Maintenance of the population above the be desirable. However, present hunting philosophies proper level is likely to damage the habitat and further do not follow this line of reasoning. reduce the carrying capacity It has beer1 demonstrated Change in size is difficult to detect and evaluate since numerous times that under adequate protection deer- it relates so closely to nutrition factors and age. Differ- herds will increase beyond r~ingecapacity in a relatively ence in size as a genetic trait is readily observed in other short lime Adequate protection requlres cooperation mammals, especially in domesticated forms where arti- of the people who own and use the land Once this attl- ficial selection has been practiced. It seems feasible that tude is developed and deer populations are brought from man's influence on deer populations could modify natural scarcity to abundance there is often a reluctance by selective processes. Such genetic considerations in deer- landholders to follow the advicc of biologists and allow herd management may, in time, become quite important removal of antlerless deer Thus, in many areas the popu- as the demand for more intensive land utilization in- lation is held well above optimum levels creases. Since food is a major factor in limiting deer popula- tions, it is likely that in high populations some survival value would accrue for those animals which require less LITERATURE CITED food to sustain life. Since energy requirements are re- lated to body size it seems feasible that the genetically Eberhardt, L. L. smaller animals would have a selective advantage over 1969. Population analysis. In Wildlife manage- larger animals. This positive selection for smaller ani- ment techniques, pp. 457-495. Wash., D. C.: mals is further accentuated by hunting philosophies in Wildlife Soc. which a premium is placed upon the larger deer. Theo- Kelker, G. H. retically then, there are two selective forces which favor 1947. Computing the rate of increase for deer. survival of small animals. Such selection over a long J. Wildlife Manage. 11: 177-183. period of time could result in reduction of the size of the deer within the population. Surface area, and conse- Leopold, A. quently heat loss, is proportionally greater in smaller 1948. Game management. 481 pp. N. Y.: Charles animals and one would expect such tendency for decrease Scribner's Sons. in size to be offset by greater mortality of small animals O'Roke, E. C., and Hamerstrom, F. N. in areas where winters are severe. In the South this 1948. Productivity and yield of the George Re- offsetting selection would he of minor consequence. This serve deer herd. J. Wildlife Manage. 12: concept offers additional sound reasoning for keeping 78-86. populations at optimum levels and for a less-selective removal practice in established populations. Conversely, Sevcringhaus, C. W., and Cheatum, E. L. it indicates some value in making special effort to retain 1956. Life and times of the white-tailed deer. some larger animals for breeding purposes. Reduction In Deer of North America, pp. 57-186. in size would not necessarily be detrimental to the popu- Wash., D. C. : Wildlife Manage. Inst. Foods and Feeding Habits of White-Tailed Deer Daniel W. Lay ' Texas Parks and Wildlife Department Nacogdoches, Texas

Principal foods of white-tailed deer are browse, fruits, succulent herbage, mushrooms, and ngricul- tz~ralcrops. Deer diet is notable for great variety. Browse accounts for less than half. Preference var- ies by season and among species as does quality. The best insurance for proper deer nutrition on forest range is maximum habitat variety.

Deer diet is one of the oldest subjects for wildlife investigations and it remains an essential part of deer management in southern forests.

FOODS

Browse

Leaves and twigs of woody plants constitute browse, which is generally considered a major component of deer diet. Field observations on a deer range reveal more sign of deer use of browse than anything else. Browse is the one component of the food supply which is avail- Wild persimmon. Fruits of many forest species able in some form year-around. But leafless twigs are are important to the white-tailed deer. rarely used in the South. Deer are not especially well equipped to digest the high fiber content of browse. The plants which produce fruits for deer usually pr Food habits studies often reveal only minor utilization vide browse also. of browse. Mushrooms and Fungi

Herbage Most mushrooms and other fungi are relished 1: deer. Bracket fungi have been found in stomachs in add Nonwoody plants include many deer foods. Most tion to the more common fleshy mushrooms. This is or broadleafed herbaceous species are likely to be utilized. food item that is difficult to appraise by field observ If deer have access to openings in early stages of plant tion, as no sign remains when one is consumed. Tk succession, they are sure to find many attractive herbs. supply also comes and goes in a matter of hours or day Grasses and sedges are taken in limited amounts. When early spring growth appears, some grasses (especi- Agricultural Crops ally those with winter rosettes) are taken. On ranges Many crops attract heavy deer feeding: peas, melon where other foods are scarce, grasses may be used year soybeans, corn-to name a few. Agricultural crops a1 around. Bracken and other species of fern are taken especially important after acorns are consumed. Damag by deer. may reach the point that certain crops cannot be prl Dried leaves, both woody and herbaceous species, are duced in the presence of deer populations. used. The extent of use and its significance are not clear. Fertilization makes crops especially attractive. Wintc greens such as oats, vetch, rye grass, and clovers a Fruits attract heavy deer pressure. Most fruits growing in eastern Texas are eaten by Probably heavy fertilization of evergreens such : deer. As shown in table 1 xed remains of 37 species or Japanese honeysuckle lLonicera jnponicu) would maE genera were found in 3,195 deer pellet samples. August them equally attractive. Pine (Pinus sp. I seedlings fro1 to January was the period of heaviest use although some fertilized nursery-beds may be browsed by deer whc seed remains were found every month. Acorns (Quer- p1;inted on a range, with little food present. Yet natur. cusj were recorded for every month but April. Three pines are not taken under moderate stocking. species were found every month of the year: yaupon (llex vomitoria), hawthorn (Crntc~egusiand partridge- Water berry (Mitchella repens). Some species are known to be used but the seed were not readily found in droppings, Deer in the South do not require or use surface watc due to tiny size or high digestibility. at all times. Apparently much of their requirement

Aid in Wildlife Rcstor;rtion PrOj- s~lppliedby the vegetation they eat. Most deer fooc ccts. are 50 to 90 percent water. Table 1.-Frequency of fruit remains in 3,195 east Texas deer pellet groups, 1956-68 - Name -- Arnpelopsis arborea 3 1213 10 Rerschemia scanderas 1 2 2 3 8 Callicarpa americana 911 37 93 122 106 71 33 473 Carya sp. 1 1 Celtis occidentalis 11 1 3 Cl~ionanth~~svirginicus 111612 7 6 1 44 Cornus floridu 11 5 1 4 27 21 69 Crutuegus sp. 41 21 6 8 19 10 9 39 40 42 10 14 259 Gleditsia sp. 14 8 17 3 33 Hypericum sp. 2 2 Ilex coriucea 1 1 I. deciclua 6 9 7 5 2 2 31 I. opaca 3 15 1 1 1 1 22 I. vo~nitoriu 58 39 13 5 16 14 8 10 5 10 42 36 256 Lonicera japorzica I 1 2 Mitchella repens 7 20 3 5 1 10 17 25 17 23 12 10 150 Myrica cerifera 18 5 1 9 33 Ngssa sylvalica 30 2 1 2 4 7 22 26 40 134 Quercus sp. 70 60 3 1 1 1 5 43 100 67 38 389 Prunus mexicanus 9 9 Prvnus sp. 1 1 Rhus sp. 22 10 2 3 5 15 17 18 17 109 Rubus sp. 24 43 11 4 1 83 Sambucus canadensis 2 7 3 12 Smilax glauca 1 1 S. laurifolia 1 1 2 S. rotundifolia 15 2 1 1 10 S. walteri 1 1 Smilax sp. 3332318 Styrax arnericana 12 3 Synzphoricarpos orbiczclatz~s 4 4 Symplocos tinctoria 1 12 5 2 20 Toxicodendron radicans 252 9 Vaccinium arboreuin I 1 Vaccinium sp. 1 2 1 4 Viburnunz rnolle 1 1 Vitis sp. 323 8 Totals 289 197 31 10 66 98 119 217 282 382 294 223 2,216 Number examined 354 493 218 198 213 262 247 291 220 281 223 195 3,195 Identifications per 100 examined

FACTORS AFFECTING DEER FEEDING HABITS territory. Deer die offs have occurred in pastures located within a mile of understoclted range with surplus food. Animal Needs of the deer vary with season and sex, and Diet is limited to what is available within the travel selectivity doubtless represents search for nutrients as range of deer. An individual deer may be observed required by lactatiori, growth, reproduction, and antler regularly in a key spot of habitat--such as a honey- development. Selectivity of the anirnal may result in suckle thicket--that may not exceed 1 acre. a diet with 66 pcrcelit more protein than hand-collected material of the same species, according to Wilson (1969). Enclosed deer may spend a lifetirlle on tracts of less than 100 acres, but there is some evidellce t2i:rt confined The most critical diet requirement, considering thc deer are not as well nourished by a given habitat as numerous authorities, is for 13 or 14 percent of protein unconfined deer. A possible reason for this would be and 0.40 percent phosphorus (Murpl~yand Coates 1966). the better use of uncommon food items a deer may find Also, the fiber level should be modest to assure adequate in a larger territory. digestibility. Deer move to seasonal supplics of acorns and other Daily feeding routines are not especially notable. When fruits. Yet they will not lnove out of their established undisturbed deer generally have two feeding periods- early and late daytime. With disturbance they feed at taken at a 55 percent rate, the medium choice specif night or when they can. Corn, for instance, when placed will be taken at a 30 percent rate, and the low choic in front of a hunting blind is generally taken when the species at a 5 percent rate-in Texas experience. Thi blind is empty. means that not more than a third of the current growl The reach of deer limits feeding to the zone up lo 4.5 on all browse plants should be considered available t feet high. When necessary, feeding may be accompanied deer. A range needs to have available, according t by standing on the hind legs and reaching higher. This these estimates, 273 pounds (dry basis) per acre to carr occurs in periods of scarcity and favors the stronger a deer to 10 acres. and taller individuals. In north Georgia, Ripley and McClure 11963) founm an average of 36 pounds per acre of browse forage wit Environmental-Season only 16 pounds of that desirable species. They estimate' it would take 52 acres of National forest or 33 acres o Seasonal changes in the deerfood supply are a major private land to carry one white-tail. This was basel aspect. The composition of foods changes with the stage on 2.5 pounds consumption for 100 days at a level of 41 of growth. Many food items ripen or develop at a specific percent utilization. Two questions arise: is this a satis season. Others are taken only at certain seasons due to factory deer density and what do deer eat the other 261 changes in available foods. Evergreens have a special days each year? value in winter. On some ranges winter browse is almost nonexistent The major support for deer on a range may be one Dunkeson i 1955) reported this for the Ozarks where onl: or two good evergreens such as yaupon (1. vomitoria) the unpalatable pine and cedar [Juniper) are generall: or Japanese honeysuckle. Seasonal availability deter- available. mines use of such desirable foods as fruits. Some fruits In Ozark forests studied by Segelquist and Greet dry in place and are found by deer long after the period r 1968), mast was five to 10 times more abundant thar of fresh fruit availability. Others may be scratched out winter browse-98 pounds of acorns to 9.2 to 19.9 pound: of drifts or used when they germinate-as acorns. of browse. Environmental-Weather A study of browse utilization by penned deer in a 51 acre enclosure at Kirbyville revealed less than half o The mild southern climate exerts little direct influence the diet was supplied by browse (table 2 ). on deer feeding. Rarely does sriow or ice cover the food supply and then for short periods. Bunching in "deer- Table 2.-Estimated contribution of browse to deer die yards" does not occur. Summer heat seems to cause more in a 58-acre enclosure at Kirbyville (air-dr~ stress than winter cold. "Die-offs" occur more often in weights) late summer. This is the period of declining forage quality and increasing strain of lactation and growth on does and fawns (Short et al. 1969; Goodrum and Reid 1962 ).

Environmental-Topography The southern terrain does not limit deer access and little consideration for topography seems to be required. Deer do use certain trails, runs, and feeding spots more than others for obscure reasons.

Food Availability Food supply is the major determinant of deer range carrying capacity. Much effort has been devoted to Fruits may be more abundant than is generally recog. measuring the browse supply as an index to how many nized. Yields of some cultivated crops range into the deer can be supported. Often the results indicate more tons per acre. deer could be grown if limiting factors, such as excessive The current study of deer in two enclosures at the hunting, were not operating. Stephen F. Austin Experimental Forest, Nacogdoches Methods of appraising the food supply sometimes need has revealed something of the productivity of fruit ir refinement. Too often species or plant parts are included southern pine-hardwood forest habitat. About 34 fruit. which are not nutritionally adequate. Also, summer ing, woody species are present. The average fruiting surveys need careful interpretation as to what would be population is 138 plants per acre in the pure pine stand available at other seasons. and 165 in the pine-hardwood stand. Pines are no1 It is known that a 100 pound deer needs about 2.5 counted. The most common species is American beauty. pounds of dry forage per day (Davenport 1939, Nichol berry (Callicarpa c~rnericar~a)and estimates of its fruit 1938; Smith 1950). To maintain a herd of one deer to yield have ranged to about 60 pounds per acre. The 10 acres, each acre would contribute 91 pounds of dry yield per plant has been about 0.25 pounds for bcauty- weight of acceptable food ( 2.5 x 365.'10 ). berry and 0.95 to 3.3 for dogwood (Cornus florida). Browse utilization should not be more than about half Yields of 20 species are shown in table 3, based on counts for the welfare of the plant. If the choice species arc of fruit on 658 plants. Table 3.-Sl~mmt~ry of whole-tree fruit counts in eastern Tesns in as nutritional quality, seasonal conditions, preference pol~ndsper tree ' of deer, and short-term foods like mushrooms-all com- plicate the process. Species Food Quality

American bcaulyherry Food quality is more limited than quantity. Phos- i(:ullzccirpa amc?tcanai '! 42 0.47 0.24 1963 100 .19 phorus and nitrogen or protein are generally inadequate 1964 100 .30 except in spring. 'I'able 4 gives estimates of seasonal 1966 58 2.8 composition of browse by three categories of palatabiIity White fringetree ( Lay 1957). (Chzontinthi~s~*irginzc?~s) 1958 41 - 1.75 1.1 1963-67 47 1.66 .81 Flowering dogwood Table 4.-Seasonal variation in protein and phosphoric (Cornus (lo~idtij 1958 26 4.0 3.3 acid in browse forage ' 1966 35 3.72 .95 1967 38 3.17 2 35 ATumber Season Protein Phosphorus Blueberry hawthorn species (Crataegz~sbrachljacantha) 1938 10 3.0 2 4 Parsley hawthorn Spring 10 14.5 0.18 (C.7narshall1) Summer 22 9.5 .08 Common persimmon Fall 10 8.2 .08 iDiosp?/rosi>i?'g!nianai 1959 1 4.1 13.4 Winter 25 7.2 .08 ' Two-inch tips, corrected to ovendry matter base (Lay 1957 ) . Possumhaw illex deciduai 1958 8 1.8 2.2 Gallberry Crude protein dropped from about 15 percent in spring (I corzaeeaJ to 7 in winter, phosphorus dropped from 0.18 to 0.08. These may be compared to the 13 percent protein and the 0.40 phosphorus needed. Seasonal differences are Blackgum great and most species are best at only one time during /iV{gssa sylvalzcai the year. Red bay persea Special recognition should be given evergreens. These (Persea horboniai 1959 6 2.56 .93 may approach adequate levels for protein even in winter. Black cllrrry Smilaz smalli has 15 percent protein and 0.12 phosphorus /l'rz~?zusserotir~ui 1959 1 3 05 1.73 (Lay 1957). Flaturoods plum Some good densities of deer do occur. Several methods (Prrrnz~sumbellutc~! 1958 8 3.1 1.18 are used to offset the scarcity of N and P. Deer feed Chines? tallowtree with care, often taking only a tip here and a leaf there. iSapzzirr~sahiferun~i 1960 3 4 88 5 75 They make heavy use of such high quality items as Common swcelleai mushrooms which in winter in east Texas may average iS;,niplocos tlnctorzai 1958 38 2.4 .64 23 percent protein and 0.55 percent phosphorus (Miller and Halls 1969). Succulent greenstuff, however, is the common source of P and N. Kentiicliy viburnum Fruits, including acorns, may be high in fats and (ViO~rl.n~rmmollri 1959 51 .87 .08 digestible carbohydrates. Acorns may have 30 percent fat and 50 to 80 percent nitrogen-free extract (Fraps 1919; Caillouet 1960; Goodrum 1959). Fat content of some other fruits are: whole dogwood 17 percent, -

.- wood flesh without seed 34 percent, whole arrowwood 658 - fruit (Viburnzin~dentaturn,' 26 percent, yaupon 13 per- ' All trees wcre randomly selecteti among those having sorne fruit and cent, and American beautyberry 11.5 percent. Parsley we1-e located In a fully stocked pilie foresl, crcept the single-tree haw (Crataeyus viridis) and dogwood each contain more samples. than 1.5 percent calcium (I-lastings 1966). Diameter was measured tl inches high. The fiber shown for fruit is largely in the seed which is passed without digestion. The digestible part is low in fiber and this enhances its contribution to the diet. The great contribution of fruit to the diet is energy. Browse availability is important because it lends Any hunter knows deer fatten soon after acorns start itself to measurements, is present ycar-around, and falling, if tlkey have not already fattened on late summer serves as an indicator of the degree of pressure. Caution crops of blueberry hawthorn (C. braclzyacantha) and/or should be used in converting pounds of forage present American beautyberry. These enable the building of into deer carrying capacity. Unmeasured variables such reserves for the rut and for winter. Food Palatability a time and place where they can grow and fatten on ; Some ranges, such as one with a long history of high abundant supply of foods with adequate protein, pho phorus, and energy deer and/or livestock stocking, may have few first-choice foods present. Second and third-choice foods may be The use of many kinds of fruits, forbs, musliroon taken with seeming relish; but this does not denotc and other succulents in addition to many species I preference. Observations are more significant on lightly browse demonstrate the importance of variety ( Li stocked ranges with no livestock present and with a 1967 ). This is necessary because most foods have certaj previous history of little or no stocking. times when they are most nearly adequate. The role of the digestive fauna must be considered. The best insurance for proper deer nutr~t~onon forc Deer need time to adapt to new foods-even to corn. range is maximum habltat variety They must have or acquire the proper kind of digestive fauna to break down a given food with a net gain in LITERATURE CITED energy. Some workers (Longhurst et al. 1968) have shown Caillouet, C. W. relationships between palatability and evolutionary de- 1960. Nutrient content, of some s:-eds anci frui velopment of the plant species. Some plants contail1 utilized as food by wildlife in 1,ouisi;in protective substances which cause deer to avoid eating M. S. Thesis. La. State Univ. Baton Roug them. 72 PP. Deer may be forced to eat pine but this does not Davenport, L. A. denote palatability or adequacy. It. should be considered 1939. Results of deerfeeding experiments at Ct an alarm that deer are ill fed and in a precarious position. sino, Michigan. Fourth N. Amer. Wildlii There is a sequence of utilization, even among the Conf. Trans. 1939: 268-274. third-choice species. Pine browsing, for instance, follows heavy use of wax myrtle (Myrica cerifera). Such species Dunkeson, R. L. have special value as indicators of pressure on the food 1955. Deer range appraisal for the Missou. supply. Ozarks. J. Wildlife Manage. 19: 358-36' To add to the complex of palatability, individual ani- Fraps, G. S. mals differ significantly. 1919. Feeding values of certain feeding stuff Composition tests may not show why deer prefer one Tex. Agr. Exp. Sta. Bull. 245, 29 pp. species over another. I suspect choice usually is guided by nutritional requirements, and that deer recognize Goodrum, P. D. food quality better than we. The avoidance of high fiber 1959. Acorns in the diet of wildlife. Southeas content may be as necessary as the attainment of re- Assoc. Game and Fish Comm. 13th Annt quired N and P. Conf. Proc. 1959: 54-61. Goodrum, P. D., and Reid, V. H. Food Competition 1962. Rrowsit~g habits of white-tailed deer i Deer die with stomachs full of the wrong things. The the western Gulf Region. First Nat. White Tailed Deer Dis. Symp. Proc. 1962: 9-11 deer manager can anticipate starvation by observiilg the competition for food. The herd should be reduced Hastings, E. F. before such die offs occur. Often this happens in un- 1966. Yield and chemical analyses of fruit prc spectacular loss of fawns and does in late summer. duced by selected deer-browse plants i Close observation and imagination is required to grasp a loblolly-shortleaf pine-hardwood fores the true degree of food competition among deer. On Ph.D. Thesis. La. State Univ. Baton Rougc ranges where the fawn survival is less than 50 percent, 230 pp. serious competitio~lis indicated. Carcasses of dead deer Lay, D. W. are not often found because they decompose or are con- 1957. Some nutrition problems of deer in th sumed rapidly. Also, visibility in dense cover may be southern pine type. Southeast. Assol only a few feet. Game and Fish Comm. 10th Annu. Con Livestock eat many foods taken by deer. Cattle reduce Proc. 1956: 53-58. the carrying capacity for deer it1 southern forests by eating deerfoods and by suppressing their reproduction. Lay, D. W. X'lany browse species, especially evergreens, are superior 1967. The importance of variety to southern deci to forest grasses and they attract cattle as much as deer. Southeast. Assoc. C;anie and Fish Comn Some browse species are taken by cattle but not by 18th Annu. Conf. Proc. 1964: 57-62. deer. Flowering magnolia (Mugnolicr grniztliflo~rr) anti M. American holly (1. opczcn) are examples. Longhurst, W. M., Oh, 11. K., Jones, B., and Kep11e1 It. E. SUMMARY-THE IMPORTANCE OF VARIETY 1968 A basis for thc pal~tabilityof deer forag plants Thirty-Th~rrlN Amer M'lldl~fc an Dfer in i;outhcin forest5 no] m,rlly '11 c se,irt lliilg fox Ndtur Resour~eiConf Tran5 1968 181 food5 whicl~wlll ~mprovcthe~r dict Rlydo lhcy iind 189 Miller, H. A,,and Halls, L. K. Segelquist, C. A,, and Green, W. E. 1969. Fleshy fungi commonly eaten by wildlife. 1968. Deer food yields in four Ozark forest types. USDA Forest Serv. Res. Pap. SO-49 tin J. Wildlife Manage. 32: 330-337. press). South. Forest Exp. Sta., New Or- leans, La. Short, H L, Newsom, J D, McCoy, G L, and Fowler, J F Murphy, D. A,, Coates. ,J. A. 1969. Effects of nutrition and elimatt: oil southern 1966. Effects of dietary protein on deer. Thirty- deer. Thirty-Fourth N. Amer. Wildlife and First N. Amer. MTildlife and Natur. Re- Natur. Resonrces Conf. Trans. 1969 : 137- sources Conf. Trans. 1966: 129-138. 145. Nichol, A. A. 1938. Experimental feeding of deer. Ariz. Agr. Smith, A. D. Exp. Sta. Tech. Bull. 75, 39 pp. 1950 Feeding deer on browae 5pecles during win- ter J Range Manage 3 130-132 Kipley, T. H., and McClure, J. P. 1963. Deer browse resources of North Georgia. Wilson, A. D. USDA Forest Serv. Resource Rull. SE-2, 1969. A review of browse in thc nutrition of 20 pp. Soutl~east.Forest Exp. Sta., Ashe- grazing anim;rls. J. Range Manage. 22: ville, N. C. 23-28. Physiology and Nutrition of Deer in Southern Upland Forests

Henry L. Short ' Southern Forest Experiment Station Forest Service, U. S. Department of Agriculture Nacogdoches, Texas

Upland forests in the South generally have in- Rates of food consumption of yearling females ha fertile soils and produce roughages that are season- a similar but less variable pattern (Short et al. 1969 ally deficient in net energy, protein, and phosphorus Does in late gestation consumed more food than nor for deer. To increase the number and size of deer, pregnant does. During and immediately after lactatioi palatable and nutritious foodstuffs must be produced does with fawns might ingest one-third more food tha at seasons when naturally occurring foods are nu- those without fawns. tritionally inadequate. Table 2 shows the approximate nutritive content c White-tailed deer (Odocoileus virginianus L.) on the several east Texas foodstuffs. Energy contents of fooc extensive upland habitats of the Southern United States stuffs (kcallg.) were calculated from proximal analyse are often small and occur at relatively low densities. and Tyler's (1964) conversion factors for protein, fa This situation is caused largely by low soil fertility, and carbohydrates. Caloric values for different types c which seriously limits the production of nutritious for- foodstuffs are similar to those cited hy Golley (1961 age. This paper reviews findings about seasonal nutri- Acorns have relatively high gross energy (5.4 kcal/g. tive requirements of deer and the seasonal nutritive loblolly pine browse has intermediate values (4.4 to 4. quality of forest foodstuffs, which are sometimes defi- kcallg.), and mixed browse, mixed grasses, and mixe cient in net energy, protein, and phosphorus. forbs have low energy levels (3.9 to 4.3 kcal/g.). Acorn have a higher gross energy than do most fleshy fruit ENERGY and legume seeds, and all have more ether extract (an Energy intake of deer varies with age, sex, and re- therefore more gross energy) than do grass seeds (Kin productive condition. In a recent study (Short et al. and McClure 1944). Fleshy fruits and berries are c 1969), ad libitum consumption of a nutritious ration by moderate energy value (Wainio 1941). yearling bucks was high during spring and early summer Gross-energy intake of yearling bucks on an adequat (table 1) . Intake diminished in midsummer, apparently ration was 5,000 to 6,000 kcal/day from April to Octobe because of climatic stress. Food intake increased in late and 3,500 to 4,000 kcal/day during late autumn an summer or early autumn, when heat and humidity mod- winter (table 1). Consumption of only 1 to 2 kg. (over erated. It decreased during rut and remained low dry weight) of nutritionally adequate foodstuffs per da throughout- the winter, when nutrient deficiencies in would satisfy these energy requirements. forage normally exist. - The digestibility of a foodstuff determines the amoun 'The author is on the staff of the Wildliie Habitat and Silvicul- of the gross energy that is available to animals. Dry ture Laboratory, which is maintained by the Southern Forest Experiment Station in cooperation with Stephen F. Austin matter digestibilities of the foods in table 2 were estj State University, Nacogdochcs, Texas. mated from their fiber components. Though these value

Table 1.-Weight gains and average rates of nutrient ingestion for deer on rations of constant and varying nutrient value'

Month

Mo. Kg. Kcal. Grams Grc~ms Kcal. Grams Grams

Feb 8 32 9 10 0.8 Mar 9 34 8 12 1.1 AP~ 10 39 4 26 2.4 May 11 452 26 2.4 June 12 50 7 23 2.2 July 13 56 0 25 2.3 Aug 14 59 0 22 2.2 Sept 15 62 3 16 1.9 Oct 16 642 16 1.8 Nov 17 63 1 11 1.2 Dec 18 58 9 11 1.2 -- Total 11 months 6.0 kg. - -- - .6 kg. 'Data from Short et a1 Table 2.-Nutrient content and predicted dry-matter digestibility of several foods sampled from southern upland forests at different seasons of the year Estimated Cell- Predicted Crude Forage gross Phosphorus wall ADF Lignin dry-matter protein energy I content digestibility ' Kcal./g. Percent dry weight Pct. ADF Percent Woody twigs with leaves, when present (terminal 2 inches) May 4.3 0.3 16 32 27 July 4.3 .1 10 34 27 Sept. 4.3 .1 10 37 31 Nov. 4.1 .1 6 32 29 Feb. 4.2 .1 6 57 46 Current twigs of woody plants (terminal 4 inch average summer-winter) 4-6 70 53 Green leaves of browse plants 8-14 32 24 Dead fallen leaves 4.1 .4 5 49 36 Mixed forbs (terminal 2 inches) May 4.2 .3 15 34 29 July 4.3 .2 11 37 32 Sept. 4.2 .2 9 41 35 Nov. 4.2 .1 8 43 37 Mixed grasses (terminal 2 inches) May 4.2 .2 13 67 38 July 4.1 .1 9 67 40 Sept. 4.2 .2 12 68 39 Nov. 4.0 .1 7 71 44 Feb. (includes fertilized winter grasses) 3.9 .2 15 57 39 Loblolly pine (terminal 2-inch twigs with needles) May 4.4 .2 9 45 36 July 4.4 .1 7 45 39 Sept. 4.4 .1 7 47 40 Nov. 4.5 .1 8 45 39 Feb. 4.6 .I 8 41 37 Acorns (whole but without cups) 5.4 .I 6 28 22 ' From equations for estimating digestibility of dry matter (Van Soest 1965). may not accurately depict the usefulness of these forages Dead grasses have low potential digestibility, but fertil- for deer I Short, H. L., unpublished data), they indicate ized cool-season grasses are as digestible in winter as the relative usefulness of different foods and the effect mixed warm-season grasses are in summer. Predicted of seasonal changes on digestibility. digestibility of the grass samples in table 2 is less than Woody twigs have a relatively high lignin content. actual digestibilities listed for some orchard grass, brome Composite samples of the terminal 2 inches of twigs with grass, and timothy samples cited by Van Soest (1963). attached leaves may be 57 to 61 percent digestible. Di- Pine browse samples contain intermediate CWC levels, gestibility diminishes to about 41 percent after leaf drop. relatively high lignin levels, and more ADF than other Green leaves of woody plants are 64 to 67 percent di- browses. Pine samples are about 50 percent digestible gestible, while dead leaves, after nutrients have been throughout the year. leached, are only 39 percent digestible because of a high Digestibility percentages may be almost 20 points lignin content. lower in autumn and winter than in spring. The net The terminal 2 inches of mixed forbs have a higher energy values of most plant tissues, therefore, are far cell-wall content (CWC) and acid-detergent-fiber IADF) lower in autumn and winter, even though similar caloric content but a smaller lignin content than browse. Thus, contents are present throughout the year. After the forbs are predicted to be slightly more digestible than growing season has terminated, high levels of fiber com- browse. The terminal 2 inches of mixed grasses char- ponents limit the value of foods such as woody twigs and acteristically have high CWC and Iow Iignin content. dried leaves. Energy deficiencies exist for deer when- ever the net energy derived from foods fails to fulfill particularly efficient in utilizing the available soil nib maintenance or production requirements. Because of gen. their relatively high metabolic requirements, lactating Stand conditions affect the total nitrogen availab does and weaned fawns are particularly vulnerable to to deer. In a model of nn old-field community that SUI low-energy diets. ceeded to pines i Switzer et (~1. 19681, the total nitrogc in the standing herbaceous cover that was physicall PROTEIN available or contained in plants palatable to deer progre, Protein is essent~alfor body maintenance and pro- sively diminished. When the pines were 5 years of agi duction Yearling bucks fed a control ration of about 15 the total nitrogen in the ecosyst.em was about the san: ptrcent proteln (N x 6 25) consumed 8 9 kg N from as it had been 5 years before, but only about 60 pcrcei February through December (table 1) They consumed was in vegetation potentially useful to dccr. At age ' about 30 g N per day from Apr~lthrough October and the pine canopy had closed and, even tilough total nitrc increased body we~ghtat least 5 percent per month from gen in the ecosystem had increased, little or none w; February througll September available to deer. 1,ittle forage nitrogen will be avai Bucks fed diets formulated to resemble natural for- able to deer until the canopy opens and forage agal ages which are seasonally ava~lablein upland forests develops near the ground i Swit~eret al. 1968) consumed only 6 kg N from February through December When deer-forage species do ~nvadethe plantatlor During spring (dietary protein about 17 percent) and their contribution to the nitrogen requirements of dee early summer (dietary protein about 12 percent), they may be small The total annual production of browi consumed somewhat less nitrogen per day than d~d and herbage in a Lou~sianaplantation of 105 to 13 control deer Body weight increased at least 5 percent square feet of bad area was about 90 to 135 kg acr per month only from April through August Gall1 essen- (Rla~r1967) Many browses and hcrbages contallled n tially ceased in late summer (d~etaryprotein less than 9 more than I to 15 percent N 19 percent or less proteir percent) During autumn and winter (dietary prote~n after growth ceased (table 2) Thus, in this plantatlo about 8 percent) these bucks lost more weight than d~d only up to 2 kg acre of nltrogen were potentially ava~l control-fed deer. Deer fed the control ration consumed able to deer Stands of shortleaf and loblolly p~nesant 48 percent more protein and gained nearly 30 percent mixed hardwoods in east Texas produced approximatel: more in body weight than did deer on the experimental 195, 330 to 385, and 725 kg.Jacre at basal arcas of 9E diets (table 1). 76, and 26 square feet (Schuster and I-Ialls 19631. Th Deer fed low-protein diets (7 percent) throughout the most open stands probably provided no more than 11 kg year were physically stunted, and does fed diets of 7 of forage nitrogen per acre. to 11 percent protein produced fewer fawns than those Plant tissue has a h~ghnitrogen content only whei on an adequate diet i Murphy and Coates 1966 ). growth is rap~d At this t~me,nitrogen may make u] Protein deficiencies in vegetation restrict the growth more than 3 percent of the dry m,ltter in new growth and development of deer Deer kllled durirlg autumn After tissues mature, nitrogen content is d~lutedby th on upland forests In Louisiana weighed only 68 percent rapid accumulation of carbohydrates Some translocatioi as much as eaptlve deer fed an adequate rat~on(Short and leaching occur from leaf tissues as early ,is Jul: et a1 1969) Protein deficiencies also restrlct populat~ons (Kramer and Kozlowsk~1960) Thus, the nitrogen con in uplands. Inadequate protein during lactation prob- tent of the terminal portion of many forest forages i ably reduces fawn survival and hinders the recovery of often lcss than 15 percent in autumn and winter (tablt lactating does prior to breeding. Low protein levels also 2) Furthermore, much of the nitrogen in pcmrly digcstcc probably adversely affect the growth and survival of fibrous t~ssuesin stems, twlgs, and dead leaves is no newly weaned young. metabolically available to deer. The predominant red and yellow podzolie soils of up- Foods which conta~nadequate levels of crude proten land habitats tend to have high aluminum and iron oxide at seasons other than sprlng are legume, pine, and gras contents, a low reserve of organic matter and many soil seeds (36, 26, and 12 percent, respectively) (King an( nutrients, and a low exchange capacity. These highly McClure 19441, mushrooms (up to 30 percent) [M~lle leached soils are moderately to strongly acid and support and I-lalls 19691, and leaves of some evergreen broad forest vegetation high in carbon and low in nitrogen. leaved species These highly nutr~tiousfoods are oftel Such environments support relatively low densities of scarce in the forest (Albrecht 1957). Soil fertility varies by site, and leaves from plants growing on nitrogen-rich soils PHOSPHORUS contain more nitrogen than those on deficient soils. Plant species themselves vary in nitrogen requirements. Phosphorus is needed for metabolism in Red, white, and chestnut oak and red maple grow on and is 3 major constituent of bones, antlers, and sof soils low in nitrogen, while such species as wl~iteash, tissues. Phosphorus deficiencies in fcmale herbivore yellow-poplar, and basswood need soils with a high affect Estrus and therefore influence productivity (May nitrogen content (Kramer and Kozlowski 19601. nard and Loosli 1962 ). Does require relatively high level Plant parts and plant species vary as sourccs of nitro- of phosphorus during gestation for normal developmen gen. 1,eaves are better sources than twigs (Cressel 19621, of the fetus, and they secrete extensive quantities in milk and hardwood leaves are better tkan coniferous needles Phosphorus levels necessary for deer l~avebeen esti iKramer and Kozlowski 1960). Some plants, such as mated at 0.25 to 0.30 percent for maintenance and 0.51 legumes, can fix atmospheric nitrogen, and others are percent for optimum growth (Magruder el al. 19671 Apparently adequate growth and development of deer nutritional quality during seasons when many unman- occurred with d~ctithat prov~dedc~ther 0 25 or 0 44 aged deer foods are deficient in essential nutrients. percent phosphorus (Short et ul 19691 Yearhng bucks fed the control ration (044 percent P) 1nge5ted over LITERATURE CITED 5 g P day dur~ngthe rap~dgrowth period of sprlng and summer, and about 3 5 to 4 0 g day dur~ngthe Albrecht, W. A. other months of the year (table 1) Young bucks on 1957. Soil fertility and biotic geography. Geogr. the experimental diets of varying composition (0.25 to Rev. 47: 87-105. 0.18 percent P) ingested slightly more than 2.2 g. P 'day Blair, R. M. during the rapid growth period of spring and early 1967. Deer forage in a loblolly pine plantation. J. summer. Phosphorus intake was less than 1.2 g. day Wildlife Manage. 31: 432-437. durnig autumn and winter, when phosphorus levels In the diet were only 0 16 to 0 11 percent Preliminary Blair, R. M., and Halls, Id. K. evldence suggests that the p1'1sma phoiphorus levels of 1968. Growth and forage quality of four southern experlmental deer dropped in autumn and wlnter and browse species. Southeast. Assoc. Game and that the levels were lolver than those measured in control Fish Comm. 21st Annu. Conf. Proc. 1967: 57-62. deer i Matthews 1958 I. These data suggest inadequate bone nutrition i Maynard and Loosli 19621, which could Causey, M. K. result in diminished slccletal size. Control bucks in- 1964. Nutritional analysis and seasonal variation geited 167 percent more phosphorui dur~ngan 11-month of some herbaceous deer browse plants in the perlod, were larger (l,~ble1 I, arrd had more antler po~nts pine-hardwood areas of Winn and Union (Short ct a1 1969 I than did those on experlmental ra- Parishes, 1,ouisiana. M. S. Thesis. La. State tions Production capabilltics of wild deer ihould be Univ. Baton Rouge. 47 pp. very l~mitedon foodstuffs with less than 0 16 percent Gessel, S. P. phosphorus. 1962. Progress and problems in mineral nutrition The same soil characteristics which frequently limit of forest trees. In Tree growth, pp. 221-235. protein in forages grown in upland forests cause phos- N. Y. : The Ronald Press Co. phorus deficiencies in many plant tissues at seasons other than spring and early summer. Phosphorus is closely associated with plant metabolism and accumulates in 1961. Energy values of ecological materials. Ecol- actively growing tissues such as buds and newly de- ogy 42: 581-584. veloping leaves. Phosphorus content declines in older, Halls, L. K., Knox, F. E., and Lazar, V. A. senescent leaves because phosphorus is translocated back 1957. Common browse plants of the Georgia Coastal into twigs. Phosphorus deficiencies after spring have Plain: Their chemical composition and con- been reported in common browse tissues by Blair and tribution to cattle diet. USDA Forest Serv. Nalls (1968) and Halls et nl. i 1957 I, and in range forage Southeast. Forest Exp. Sta., Sta. Pap. 75, 18 by many authors. Only occasior~alherbaceous species PP. contain adequate phosphorus during autumn and winter (Causey 1964). The phosphorus content of several mush- King, T. R., and McClure, 13. E. rooms was 0.20 to 0.58 percent, legume and pilie seeds 1944. Chemical composition of some American wild nearly 0.60 percent, and grass seeds and several fruits feedstuffs. J. Agr. Res. 69: 33-46. about 0.25 percent, while acorns were clearly deficient Kramer, P. J.,and Kozlowski, T. T. with only 0.10 percent (King and McClure 1944). 1960. Physiology of trees. 642 pp. N. Y.: McGraw- Hill Book Co., Inc. FOREST MANAGEMENT AND DEER NUTRITION Magruder, N. I)., French, C. E., McEwen, L. C., and Thc population and physical size of dcer on upland Swift, R. W. habitats in the South are limited by nialnutrition. Even when they are abundant, forages grown on poor soils 1957 Nutrit~onalrccluirenients of wh~tc-ta~leddeer ollen have high carbon and fiber contents and seasonally for growth dnd antler deveIopmetit II-Ex- inadequate levels of net energy, protein, and essential perimental rehults ol the th~rdytur Penn. minerals. Some studies indicate that deer partly com- Agr Exp Sta Bull 628, 21 pp pensate for poor roughage quality by utilizing a large Matthcws. P. J. variety of supplemental foods, including fleshy fruits, 1968. Blood physiology of white-tailed deer on seeds, nuts, fleshy fungi, and succulents. varying planes of nutrition. M. S. Thesis. La. In many instances, manipulation ot' the cariopy 011 State Univ. Baton Rouge. 188 pp. upland sites may simply increase the supply of rough- Maynard, L. A,, and IJoosli, J. K. ages which are seasonally deficient in inlportant nutri- 1962. Animal nutrition. 533 pp. N. Y.: McGraw-Hill ents. I-labitat managers shocld consider alternatives to Book Co., Inc. provide nutritious foods if large numbers of c~ualitydcer are to be produced on upland habitc~ts. Supple~nents Miller, 1%A, and Ifalls, L K might be provided directly to forest deer. A better :a[)- 1969 Fleshy lungi conin~only catc.n by wilcll~fc. proach might be to fertilize areas within the forest USDA Fore5t Scrv Res Pap SO-49 (I11 press). and seed plant species selected lor palatability anci high South Forest Exp Sta , New Orleans, La Murphy, D. A,, and Coates, J. A. Tyler, C. 1966. Effects of dietary protein on deer. Thirty- 1964. Animal nutritio~?.253 pp. London: Chapm: First N. Amer. Wildlife and Natur. Resources and Hall. Conf. Trans. 1966: 129-138. Van Soest, P. J. Schuster, J. L., and Halls, L. K. 1963. Use of detergents in the analysis of fibro~ 1963. Timber overstory determines deer forage in feeds. 11-A rapid method for the tieterm shortleaf-loblolly pine-hardwood forests. Soc. nation of fiber and lignin. J. Assoc. Offi Amer. Forest. Proc. 1962: 165-167. Agr. Chem. 46: 829-835. Short, H. L., Newsom, J. D., McCoy, G. L., and Fowler, Van Soest, P. J. J. F. 1965. Comparison of two different equations fc 1969. Effects of nutrition and climate on southern the prediction of digestibility from cell cox deer. Thirty-Fourth N. Amer. Wildlife and tents, cell wall constituents, and the ligni Natur. Resources Conf. Trans. 1969: 137-145. content of acid-detergent fiber. J. Dairy Sc 48: 815. Switzer, G. L., Nelson, L. E., and Smith, W. H. 1968. The mineral cycle in forest stands. In Forest Wainio, W. W. fertilization-theory and practice, pp. 1-9. 1941. The chemical composition of forest fruits an Muscle Shoals, Ala.: Tenn. Val. Authority, nuts from Pennsylvania. J. Agr. Res. 62 Nat. Fert. Develop. Center. 627-635. Seasonal Changes in Movements of White-Tailed Beer

Robert L. Downing Bureau of Sport Fisheries and Wildlife Blaclcsburg, Virginia

Burd S. McGinnes Virginia Cooperative Wildlife Research Uriit ' Bureau of Sport Fisheries and Wildlife Blacksburg, Virginia

Ralph L. Petcher Former VPI Graduate Student, now teaching at St Paul's College Zaria, Nigeria, West Africa

Joshua L. Sandt Former VPI Graduate Student, now with Maryland Department of Game and Inland F~sh Flintstone, Maryland

Locutions of 36 white-tailed deer tagged as fawns of the deer in the South do not move very far during in a 2,322-(tcre enclosnre at Radford Army Ammzi~li- their lifetimes, since they may be responsible for the tion Plant were recorded c~lnlostevery month for spread of the species into unoccupied habitat. Sanderson the first 2$5 years of their lives. Striking changes (1966, p. 231), discussing another problem associated with in movement occurred during hnnts when more than movement studies, very aptly stated, "Emphasis two-thirds moved outside their normal ranges. Al- will have to be shifted from the movements themselves inost 40 percent of the :yer~rlir~ybucks ,rno71e(ioutside to the reasons for the movements." their norrnal ranges during the rut, but none as far We have begun a study to detcrm~nethe reasons for as :ji-mile from the center of activity. At 255 years long, irregular deer movements and will attempt to of age, however, 60 percent of the bucks moved out- interpret our flndlngs for the ~nformat~onof others Thls side their norrnal ranges, 30 percent moving more will be followed by a discuss~onof the slgniflcance of than ."'r-mile. Does did not move outside their ranges these movements as they relate to s~milarpubl~shed and durin,g the rut. Another period of increased moue- unpublished reports Our work was done in a 2,322-acre ment was the summer, especially June. The one area enclosed by an 8-foot high chain-link fence at the observed permanent clzange in range took place clur- Radford Army Ammun~t~onPlant, Dublin, Virginia. ing this time. Many deer cxxtended their ranges in The enclosure contanied 200 to 450 deer durlng the study summer but continued to use most of their original period from 1965 through 1968 During these 4 years, ranges. These results are compared with other 177 fawns were captured and marked for long-range movements reported in the literature. visual identification. Primarily, we will discuss the ir- Most studies have shown the white-tailed deer (Odo- regular movements observed for 36 of the 60 fawns coileus virginianus) to be a relatively sedentary animal marked in 1966, with occasional reference to interesting during most of its life. However, under some circum- movements of other individuals in the area. Our discus- stances it will extend or shift its home range, disperse, sion emphasizes irregular movements because they seem or even migrate. Long movements are documented in more likely to result in dispersal or changes in range. the Northern States due to work by Shiras (19351, Bart- Irregular movements are those generally %-mile or more lett 119381,Dahlberg and Guettinger (19561, and Carlsen from the center of home range activity. We are deeply and Farmes 11957). Only a few movement records are indebted to personnel of the Radford Army Ammunition available from the milder climates (Progulske and Ras- Plant for their cooperation and assistance in this study. kett 1958; Thomas et al. 1964; Michael 1965; 1,cuth 1966). Special tllanks are due Idt. Col. John W. Sevareid and In addition, almost every State has instances of deer Lt. Col. Dewey G. Weeks. having spread from small nucleus herds to occupy ad- jacent and sometin~esdistant ranges. We should keep in METHODS mind that we are dealing with a potentially mobile The fawns captured in 1966 were born between May species that has shown tho ability to seelc solitude, cover, 26 and June 13 and were captured at 1 to 12 days of and better food in rather distant places, and to escape age. Each fawn was uniquely marked with 2%-inch from floods, fire, deep snow accumulations, predators, long streamers of lli-incli wide plastic tape affixed to the and hunters if necessary. It seems that the niajority lower edge of each ear by means of aluminum "Perfect"' ' Cooperatively siipportc>dby the. Buroau oC Sport Fishcries anct ear tags (Salt Lake Stamp Co., 380 W. Second South, Wildlife, Virginia Polytcclinic Institute, Virginia Conimission Salt Lakc City, Utah 1. Observalions were made almost of Came and Inland lpislicrics, and Wildlife Mnnagcrnent Institute. every month to identify individuals and to study their Rcferencc to tradr nnn1cxs dnos not inrply cndorsemcnt of movements and behavior. The habitat, which is mostly conlmereinl products by thf E'edcral Governrncnt. rolling, open grassland, has scattered clumps of hard- woods and cedar, and several young pine plantations. to stay in the large openings, and even here they wer Because nearly 90 percent ol' the area is visible from a sometimes stalked by impatient archers. 'Iar:issmcr network of paved roads, 1,447 observations of the 36 of the deer was probably as intense as with any othc deer under study were made during tlie 32-month period. type of hunting except dog hunting, and a great de: The least number of sighting for any of the decr markea of movernent was notcd for all sex and age classcs (fig in 1966 and discussed here was 19, the most was 108, 1 and 21. Even so, no st deer were seen near their center and the mcan was 43. of activity on hunt days and no change in home rang To define some point from which to measure plotted resulted. movements for each deer, we established a center of activity based on all observation points recorded during Movements to Seek Food and Cover its first year of life. This center point was tlie intcr- Trvo sources of unusual food were provided during th section of a north-south and an east-west line, each study. A 22-acre stand of hardwoods clearcut in earl line equally dividing the observation points. With only summer of 1968 seemed attractive to decr during car1 two exceptions, this point proved to be within or very fall and winter; 27 being seen there at one time. IIom near each deer's annual home range as ascertained in ever, no significant long-range movement of tagged dee subsequent years. to reach it was noted. A 20-acre planting of new gra: in early fall, 1968, proved extremely attractive to deei RESULTS 133 were seen feeding there at one time. Nevcrtheles: no marked deer were observed to move a great distanc Hunting Season Movements to reach this planting. All observations during November 1966 were on hunt We have too few winter observations to draw cor days. Up to 80 archery hunters per day were allowed clusions about long-range movements during this se: in the area, and virtually every clump of trees was son. Several deer made relatively long movement occupied by one or more hunters. Deer soon learned during the winter, but no change in range resultec

MON NQ DEER-I38 7 7 10128 6 11 126 2 1311 6 0 9 8 0 4101 0 6 94 74 7 1035 OBSERVED

Flgure 1 -Percent of female deer rt?o?>znq '2 to 1 mzle and over 1 nzzle frorn cc.ntcr of actz?)tt?], Radford Army Ammunztzon Plant, Vzryzrzza 100

90 S I. 80 ? B 70 3 Q g 60 p: 50 3 40 2 8 30 h 2 8 20 5 10

0 MONTH-J JASONDJFMAMJ JASONDJFMAMJJASONDJ NO DEER-22 12 10 16202317 16 19 19 9 4 222315 012 13 0 09 0 0 6 13 11 7 8 910 23 OBSERVED Figure 2.-Percent of rnnle deer moving '5 to 1 nzile and over 1 mile from center of activity. Radford AT~IL~Rinin~71itiorz Plant, Virginia.

Ffowever, no long movement to reach food or cover was turned to his original range for the remainder of the expected, because most home ranges appear to offer year. plenty of each.

Breeding Season Movements Summer Movements It is noteworthy that none of the eight does observed Only one ot the 13 does ventured rnore than br mile in Novembcr-their normal breeding month- was seen from her center of ,~ct~vitydurlng the first year (note more than $5 mile from the centcr of activity in 1967, April 1967 In fig 1) However, a marked illcrease In and only one of 10 inore than !2 mile in 1968 (fig. 11. movemellt was notcd during June 1967 when 39 percent Many bucks were seen outside their home range during of the 1-year-old does moved '2 to 1 mile and another the rut, but usually they wcre not actually chasing does, 23 percent moved more than 1 mile This increased and several were alone or with other bucks. A different act~vitycontinued through July and apparently iubslded behavior due to age seemed apparent during the actual in August Movement? beyond '1 mllc wcre noted again rut, as a higher percentage of the bucks moved longer dunng t11(, summer of 1968 when the does, now 2 years distances during November I!i(i8 than during November of age, agan trequently moved Inore than '2 mile from 1967 (fig. 21. Too few 315 year old bucks were observed their center of activity Seven does t,lgged in 1965 ex- to indicate whether this greater movemcnt with in- hlbited a slmilar pattern of increased movement during creased age applies to older age classes. the sumnler at ages 1 and 2, but failed to show any No permanent change in range resultetl from rutting increase in movement at age 3 movements; however, one 3'5 year old buck was ob- Long summcrtime movei~ientsof bucks at 1 and 2 served three timcls during a 2-week period more than years old were similar to those of does, but they began 1 mile from his norrnal r;~nge. This buck was seen in these movelnents earlier. There was also a tendency the same area d~~ringthe rut as a 2'5 year old but re- for summer movements of bucks to increase with age cnote the hlgher percentage and longer duratlon of buck has taken up residence in an area of higher (leer pol movements at age 2, flg 2). lation 3 miles from his original range. Most of the moy One scemlngly permanent chnngc in home range that inent of Illarchinton's deer was on its o\vn arici n:it wh could be regarded as a disperial took plC%ceOne male being chased, but was i~nc1ouhtcdly tri~geretl by t marker1 in 1966 moved from his home range of the firit initial chase. year to a new one more than ,I mile distant sometime Our Radford deer readily re'icted to hunteri hy ]no\ i between April 5 and June 2, 1967, and hai never been considcr'ible d~st~inccswithin the eii~loiurc Vio~ct1-i seen since within the original home range Two other two-tllirds of the deer we studied nlovc>tl outilrie thl males shifted their home ranges in June 1968, but itill normal range5 during hunts, rn~inyof thrm goil-ig ai f use a small part of thew origlnal ranges If the deer had ,is the enclosure fence would allow not been enclosed within a fence, perhaps more of them The fact that movements take place tiuring huilti would have moved greater distances and found more makes us wonder how well hunter returns of tagg favorable habitat. This supposition is based as much animals represent normal dislxx-sals and migrations, a on behavior of the animals as on distance movcd. In how much of this movement is of a temporary natuj many of the cases in which yearlings were observed taking place only during hunt days. We have not prov outside their normal range during June and July, they that permanent changes in range occur as a result were alone and obviously "on the move." This behavior hunting, but we do know that hunter returns of tagg was evidenced by rapid walking or running, which deer do not necessarily indicate the normal range of t rcsultcd in the~rbeing seen sevrral times in the same animals. hour in distant port~onsof the enclosure Such brhavior was commonly observed during the early June fawning Migrations to Reach Better Food or Cover time and may have been triggered by their mothers' antagonism toward them during this period Studies of deer migrations in Northern States ha shown that deer move many miles to escape deep snc or to obtain better food and cover. Severinghaus a] Cheatum ( 1956, p. 158) said! " . . . it appears that migr Escape from Hunting tion movements are minor or nonexistent in deer rang where seasonal contrasts in weather are not great. Deer do not seem reluctant to make whatever move- Whe ments are necessary to escape a markedly unpleasant contrasts are marked, in mountainous regions. and acrc the Northern United States and Canada, seasonal migr situation. Schoonmaker ( 1938, p. 504) said, " . . . when tions of whitetails are common and sometimes pr persistently hunted the animal is apt to go beyond the nounced." limits of the home range. Also, when wounded and tracked by hunters and when trailed by dogs the deer Severinghaus and Cheatum (1956) cite studies wlti, may leave its territory, but barrlng death or acc~dent indicate that in severe winter weather, cover may ~t usually returns " Progulske and Baskett t 1958) re- morr important than food. They ;ilso said ( p. 158 I, "T' ported a deer being chased 3' 3 miles in ]/, hour by dogs spring movement back to summer range appears to Tester and Heezen (1965)also reported a deer that movcd a release from a restricted food supply during which t' 1 mile outside its home range during a dr~vecensus animals move out to the newly available spring foragc During the same drive, however, another telemetered (1938, p. 29), in discussing the mild winter deer stayed within its home range 1937-1938 in the high mountains of North Carolina, sai Urbston (personal communication), working ;it the "As the weather remained unusually mild and the groui Savannah River Project, Atomic Energy Commission in free of snow, a large number of animals remained South Carolina, reported that a buck tagged as a fawn t'riese areas all winter, not moving into lower coui~t May 15, 1967, was killed by hunters 14.5 miles distant until the last of March or early April, when early grel in November 1968. Dog hunts were conducted near the growth was available as food. Thereafter, as tht! grow capture site in 1967 and again in 19611, only 3 weeks advanced into higher altitudes, the deer again follow^ before the date of the kill and may have contributed to it upward." The senior author noted a similar altitudin this movement. However, hunting did not fully account moven~entin the North Carolina mount~iinsduring 1% for the movement, since pressure was higher where the An adult doe was captured and equipped with a rad buck was killed than in his original home range. transmitter in January 1968 near a food plot where s' Marchinton (personal communication), working in the had previously been captured more than a year befor same locality as Urbston, chased a telemetered buck with During a week of radio tracking, she remained with dogs several miles outside his normal range on several 1% mile of this location relevation 2,200 to 2.1100 fee occasions and found that the deer returned home within The transmitter ceased functioning after about 10 da: a few hours. However, on one occasion, the chase was but the doe, her tagged fawn, a tagged yearling buc continued with fresh dog packs, and the buck moved and several untagged deer were seen repeatedly for t over 7 miles from his normal range. Instead of returning next 2 months in or near the same food plot. About t immediately, he worked his way back very slowly, first of April she and the other deer were see11 movil taking 2 weeks to return. Then he stayed only 1 or 2 clownstream from this plot. A few days later the enti days before resuming his movements, becoming for a group was seen over 6 miles downstream in a broad £1 while a wanderer. This wandering behavior coincided valley at least 1,000 feet lower in elc.vation than h with the rut and may have been due to breeding activity. original location. The date of this movement coincidm Nevertheless, the buck has not yet returned home, but with spring green-up at lower elevation. Several wee later this doe wtis sccii ;igain in the rnoiiiit:i~ris where ~novc~irioiitstlrit~i~ig suiiiriirr, t~s1?c~ciallytiuri~ig the pc;ik she was originally tuggetl. of tlic, f;i\r-t~irigsc3;isoii in ,Jii!re. 'I'r;icIc coul-it d;it;~1jrc:- Progulske arid R:iskc)tt (15158, :inti 1liciri~ic:i I l!1(35) re- sc.ntc~l hy I>ti\vliirig (.I ril. i I!)(i(i~ indicntc, 11i;it ignifi- ported short migr:itioils not cc~ii~ic~c~tc~tlwith riioiiiit;iirious c,;iirtly more niovc~lilcrit cvas rccortiod in .June tl1:111 in terrain. Ellisor 119691 iiotict~ii t1i;it m;irlic.rl t1oc.r nio\.c\tl Aiigiist. bill. ,J~iric1r;ic.k coii~itswclrc 1c.s~tii;i~i Octobcr, across an intc~rmitteiitstre~ilii rl~iri~ig wiritcr. to reach ;I I rigII~I i sol ri .\l:irchintcin (per- patch of oats. IIc also iiotc~lthat ai 1ic;ivy str~ckirigof solial ci~~iiti~ii~iici~ti~~~i~~CIXII.~~tliiit ;I buck that was riitlio- cattle caused 13 m~irkc3ti dew to niove at? ;ivc,r;ige of track(~tifor 7 mc,i-itlis ;it Eglii? AF13 in 1i710rid~itook ;I 1- more than 1 rnile to a nioric~r:iti~lystoclic~tl r;riige. but it or 2-tiay exciirsioii 1noi-c. thiin 2 111i1es frc~i~ihis normal is not known if this was e;ii~secl hy tiistilrh;iiice or by riiilgc dnritig siimmc~r. a change in habitat. Ilahll~crg atltl Gucltiiiger il956, In tliscussirig rnove~~ic~ntsof niulc dclor 10. I~ctnion?tsi, IS. 59) said. "It is ;in ~:it;ihIi~h(~tlS;!et in U'isconsin that I,oopolri c.1 111. ( 1951, p. 48 i said. "During the summer the nothing will nlove ciccr like ;I loggi~igoperation. Wherc rloes rear their f;i\vns: the ycurlings b:li~ig temptir~lrily thesc opcrotio~isarc? begun hcfore heavy snowfall anci rlispossesscd, tcnd to dis~~crscand wandcr . . . . " Miller conclucted in suitable cover, they iiievit;ihly attract deer ilOfiGi, in a study of black-tailecl rlecr; ;I subspecies of that apparently 11,ive 1)revioilily wilitercti eliewhere rnule (leer, tiotetl more t.xteri~i!~e movements during Artificial feeding wlleli bcgiln t>'irly in the wintcr before May anti early June, which he called ;I prc~fawliingsl~ift. deep snowi, icemi to h,ive a iimil,ir effect " If the norm'il p'ittcrn 01 Junc movemc>nt li evcrywhcre At Radford, we obscr\~ecioiily one buck more than 1 as extensive as noteti :it Rnciford, why has this not been mile from h~stiorn~al r,irigci tluring the wintcr. ,ind he iiotc1d marc. frc~queiitlyfrom the recent rash of telemetry subsequently r tturrled home No portion of the cncloi~ire studies'? We can offer only one explanation--few of affords better food or caber than any otllcr io 11ttlc thesc movements result in a pcrmancnt change of range, winter movement was cxupecteci The clcarcutt~rig and and since deer usually move extcrisively for only a short new grass plot were attractive to deer, but no marked perlod of time, the movement may go ~mdetected deer moved to them during sumrlier and fall Choice foods were exhausted beiole the ~trldcrper~odb of winter CONCLUSIONS AND RECOMMENDATIONS Our actual count of 133 deer feeding on a small grass In spite of numerou? studies showing that deer move- plot indicates that dcer will move to a choice fooci supply ments are confined within a small area, there 1s ample Movcment of this type may be more common than 1s proof that under iome conditions perrn'lnrnt changes in generally rccognizrd range take place. Our study has shown that irregular movements may be expected during three periods of Breeding Season Movements the ycar: (1)Hunting seemed to cause extensive move- Schoonmakcr I 1938 p 503 1 iaid "A buck in quest of ments of all age classes. 121 Mature bucks moved con- a mate may travrl iar ciuriiig the rutt~ngseason siderable distances during the rut. These rnovcments Marchlnton (1968)notcd two unmai ked bucks during the were not known to be permanent. (3) Summer move- rut in an area where tl~cyhad never bcen seen bcfore ments involved yearling and 2-year-old deer and were I-Ie also noteci that one mdrkcti an~malmoved a mlle out- most prevalent during fawning season. Our limited data s~dehis range during the wme pcrlod Mareh~nton(per- and the limited size ancl diversity of the study area failed sonal cornmunic~itionialro reported a buck ne,lr Athens, to show any winter movcment that could be considered Georgia, th,rt moved 2 to 3 mllei outside his normal ;i migration. range during the rut Elliior (1960) reported a buck Future movement studies should be conducted during that moved 2 6 milcs between obiervatlons during the hunts, the rut, and during the peak of fawning. Research- breeding season in brusll habitat in south Texas. How- ers should look for factors that trigger long rtiovements. ever, no permanent changes in rangc as a result of rutting An attempt should be made to correlate habitat and dis- activity have bcen reportcti. Farther north, it may be turbance conditions of old and new locations with the difficult to separate these activities from normal migra- adoption or rejection of the new environment. We know tions. Most Statcs set hunting seasons to coincide with of many locations in the southeast where decr arc spread- the rut; therefore, movements to escape hunters may ing into adjoining good cieer range rather rapidly, while confound rutting movements and migrations. We sus- other areas that look as good remain unoccupied. pect that when a great deal of movement takes place Is there a disturbance factor such as free-ranging dogs during the rut, a few ~ndlv~tiu~ilsare apt not to return that rlecds control during thesc periods of greatest rnovc- home. We are therelore suggesting that the rutting ment? Is hunting pressure too high on the "outsidc" period may be another tirne of disprrsal. Our dtlla of herd n~iclcusareas because of overly restrictive rcgu- suggest that bucks are niorc: apt to move during rutting lations inside? In other words, could hunting or other than does. tiisturbances be utilized to drive deer into unoccupied habitat rather than out of it? Could particularly choice Summer Movements food patches or browse cuttings be used to draw deer Scveringh;rus and C!leatuni I 39563, p. 119) said, "Tlic into unoccupied territory? Through the use of Isroper summer season for decr is ir quiet, lazy time. The bucks stresses and enticements, might it be possible to create are being careful of the riltrasensitive growing antlers, widcsprcad annual migrations where none had occurred and the does are tentling thcir yoiii~g." We have not bcfore'? We hope that these questions will stimulate found this to be true. More than two-thirds of the 1- your thinking and possibly some aclditioiial research on and 2-year-old decr in our stiiciy made relatively long this subject. LITERATURE CITED Miller, F. L. 1966. Distribution patterns of black-tailccl deer loci Bartlett, I. H. coileus hemionus columbianz~s)in relation 1938. White-tails---presenting Michigan's deer prob- environment. M. S. Thesis. Oreg. State Uni lem. Mich. Dep. Conserv. 64 pp. Corvallis. Carlsen, J. C., and Farmes, R. F. Progulske, D. and Baskett, T. S. 1957. Movements of white-tailed deer tagged in Min- R., nesota. J. Wildlife Manage. 21: 397-401. 1958. Mobility of Missouri deer and their harassme by dogs. J. Wildlife Manage. 22: 184-192. Dahlberg, B. I,., and Guettinger, R. C. 1956. The white-tailed deer in Wisconsin. Wis. Dep. Ruff, F. J. Conserv. Tech. Wildlife Bull. 14, 282 pp. 1933. The white-tailed deer on the Pisgah Nation Game Preserve, North Carolina. USDA Fore Downing, R. L., Moore, W. H., and Kight, J. Serv. South. Reg. 249 pp. 1966. Comparison of deer census techniques applied to a known population in a Georgia enclosure. Sanderson, G. C. Southeast. Assoc. Game and Fish Comm. 19th 1966. The study of mammal movements-a re vie^ Annu. Conf. Proc. 1965: 26-30. J. Wildlife Manage. 30: 215-235. Ellisor, J. E. Schoonmaker, W. J. 1969 Mobility of white-tailed deer in Texas J. Wild- 1938. Notes on the white-tail deer in New York stat life Manage. 33: 220-222. J. Mammalogy 19: 503-505.

Leopold, A. S., Riney, T., McCain, R., and Tevis, L., Jr. Severinghaus, C. W., and Cheatum, E. L. 1951. The Jawbone deer herd. Calif. Dep. Natur. 1956. Life and times of the white-tailed deer. In T1 Resources Div. Fish and Game Bull. 4, 139 pp. deer of North America, pp. 57-186. Harrisbur Leuth, F. X. Penn. : Stackpole Co. Interpretation of some Alabama deer trapping 1966. Shiras, G., 111. data. Southeast. Assoe. Game and Fish Comm. 1935. Hunting wildlife with camera and flashligh 19th Annu. Conf. Proc. 1965: 110-117. Vol. I. 450 pp. Wash., D. C.: Nat. Geogr. Soc Marchinton, R. L. Tester, J. R., and Heezen, K. L. 1968. Telemetrie study of white-tailed deer move- ment- and in the Southeast. 1965. Deer response to a drive census determined k Ph.D. Thesis. Auburn Univ. Auburn, Ala. 138 radio tracking. BioScience 15: 100-104. PP. Thomas, J. W., Teer, J. G., and Walker, E. A. Michael, E. D. 1964. Mobility and home range of white-tailed dec 1965. Movements of white-tailed deer on the Welder on the Edwards Plateau in Texas. J. Wildlil Wildlife Refuge. J. Wildlife Manage. 29: 44-52. Manage. 38: 463-472. Deer Predation in North Carolina and Other Southeastern States

Frank B. Barick Division of Game North Carolina Wildlife Resources Commission Raleigh, North Carolina

A questionnaire survey of wildlife management relevant information in place of the completed questron- areas in the southeastern United States indicates naires. that predation by free-running dogs and Thus, the study is a compendium of replies to a ques- accounts for 6 and 2 percents, respectively, of the tionnaire completed by resident managers of wildlife annual drain of deer thereon. This compares with areas, and an evaluation of these replies. 63 percent through legc~lgun harvest and 20 percent It is obvious, of course, that our evaluation of data through illegal hunting. from other States is more tenuous than that of North Carolina data. Thus, our report is based primarily on STUDY OBJECTIVES North Carolina data, with somewhat gross checks for The primary objectives of this study were to identify corroboration and comparison in other States. the principal predators of deer in the southeastern United States and to determine their impact on deerherds and STUDY UNITS deer management. In doing this we have attempted to Manned wildlife management areas in North Carolina also identify other forms of deer mortality and assign range from 6,000 acres to over 86,000 acres in size. The numercial values to each, in relation to total population larger ones have several managers assigned and the and annual drain. average assignment per manager is about 15,000 acres. However, some are assigned as few as 6,000 acrcs while STUDY PROCEDURE others have over 28,000 acres. Managed hunts are conducted on all but one of the In conducting a study such as this, two general ap- North Carolina areas included in this study. The deer proaches might be considered. One would be to study harvested are brought through check stations. Resident intensively a single or small number of areas over a long area managers spend a substantial portion of their time period of years. Another method would be to study less through the year patrolling for illegal hunting and they intensively a larger number of areas over a shorter are authorized by law to kill dogs running deer as well period of time. Since we were assigned this subject only as to control other predators. a few months ago, we chose the latter course. A limited amount of food planting and browse cutting The individual study units were manned wildlife is done but in most cases this provides only a very minor management areas rather than counties or States since portion of the total food supply. In most cases basic it was felt that personnel on such areas could provide productivity of the land is lower than that of surrounding the most nearly accurate information due to their close privately-owned land. Deer populations are considered personal observation of limited land areas. While pre- to be at or above an estimated capacity of one per 50 dation is usually "controlled" on such areas, the mag- acres since annual harvest averages less than 200 acres nitude of loss in spite of control, as well as extent of per kill. control exerted, provide valuable insight into the mag- Wildlife management areas in States other than North nitude of the predation problem in localities not subject Carolina included in this study are similar in some re- to intensive protection. spects and different in others. Some areas are much To secure information on these areas, a questionnaire larger and some are much smaller. Some do not have was devised for completion by resident wildlife area resident managers and some have professionally trained managers throughout the southeast. The questionnaire biologists as managers. Most provide public hunting was filled out by all area marlagers in North Carolina but some are recently established areas that have not that have any responsibility for deer management-28 yet been hunted and at least one is a refuge area closed in all. Each man was sent a copy of the questionnaire to all hunting. and instructed to study it but not fill it out. They were filled out in the course of a series of meetings attended QUALITY OF DATA by the author so as to insure clear understanding of the meaning and intent of each question. While this does not pretend to be a "scientific" study Five copies of the questionnaire were sent to the in which basic data are compiled by trained biologists, director of each southeastern State with the request that it does, in our opinion, contain the best information avail- they be completed by managers of five representative able within the specified limitations of time. The North areas. Completed questionnaires were returned by 10 Carolina data may be characterized as having four de- States (Arkansas, Florida, Georgia, Kentucky, Louisiana, grees of reliability: Mississippi, Missouri, Oklahoma, South Carolina, Ten- 1. Numbers of deer killed and checked out on man- nessee). Some States enclosed supplementary informa- aged hunts may be considered to be as near accu- tion with their replies and one State (Virginia) sent rate as field data can be. 2. Mortalities classed as "known" or "observed" are, is considered to be much more serious on some area in about one-third of the North Carolina data, based than others. on records kept by the area manager. In other The questionnaire specified that all data be in refer cases it was purely memory. or a combination of ence to one single calendar year-January 1 t.11roug some records, and memory. December 31, 1968-so that averages derived could b 3. "Estimates" of mortality were based on the assump- considered as trnnual values. Also, since much of th tion that it would he impossible to observe all study is conceri~ed with western North Carolina area mortalities and therefore "estimated" mortalities where dog predation is a significant problem, and sinc are greater than "known" mortalities. It was also these areas average about 15,000 acres ill size, avcrag assumed that valucs for "cstimated" mortality figures in regard to them may be corlsidered as annua more closely approximated actual mortality than values for 15,000-acre units. did values for "known" mortality. Data in regard to individual area size and numbers o 4. "Estimates of potential n~ortality" might also be dogs controlled were omitted from tabulations to assur, called "educated guesses" and would, of course, anonymity and to preclude controversy irrelevant t( have the lowest level of reliability. the purpose of this study. Examination of replies showed iome "estimates" as Rcsponde~ltswere asked to indicate whether their re being extremely low and others as extremely high plies in regard to dog predation were based on records fIowevcr, there was througho~~tJ. fairly close grouping memory, or both so as to provide insight in regard tc about the means There might have been some reluctance reliability. Of 23 western North Carolina area managers to divulge information on extent of dog coiltrol in spite nine indicated that their replies were based on records of legal authority for same but we believe this was, to seven on memory, and seven on some of each. In corn a substantial extent, overcome by assuring anonymity paring the number of dogs reported controlled by thosc in each group, both the range in values and averagc ANALYSIS OF DATA values were quite similar. The observed number o deer killed by dogs, however, varied substantially. Thost North Carolina data indicate that predation of deer basing replies on records averaged 0.7 observed dee: reaches significant levels only in the western mountain killed per year per 15,000 acres while those basing replie: section of the State. It also appears that predation is on memory averaged 1.6 and those basing replies on botl least significant in the Eastern Coastal Plain section records and memory averaged 2.9. In spite of thi: which is characterized by vast wetland areas. Accord- divergence, the average of estimated deer kill per are: ingly, the data are grouped so as to derive average values was almost identical-11, 9, and 10, respectively. Whilc for 23 mountain areas, three Coastal Plain areas and these observations may not be a reflectioii of reliability two mid-State Piedmont areas. While no attempt was they do indicate a co~isiderableconsistency and the prob made to similarly regionalize the data from other south- ability that this element of consistency characterize: eastern States, it was obvious that the predation problem all estimates.

Protective coloration and meager body scent protect this fawn from predators.

26 In the case of both dog predation and predation, Table I.-Deer lossc~sin l!i(jh' 071 ~nannr?dwilrllzfe manngc?nzcnt oreirs questions were asked in regard to control so as to verify zn laesfer7r. cr~?ifrnl.and ctisfern pa?-ts of North Ccirol~rici the presence of these predators on the areas. Similarly, a question called for the number of poachers appre- hended to determine the level of concern with illegal hunting and illegal kills.

In computation of total drain and total population, SU>lBI:R values for estimated losses rather than known losses Dogs were used. This approach was used because known losses Observed liill 1.fi 2 2 are obviously minimal and it is obvious that many un- Sex of deer kilietl 4-.T-.-l observed losses occur. Hence we assumcd that estimated Estimated kills 10 7 ij I'otential kills 3 7 10 7 losses would more nearly approximate the actual. The Bobcats term "total drain" is defined as all removals of deer Observed from the population, whether by death or live transfers. Estlmnted Total deer populations were computed in the same Illegal kills on hunts manner on each area, in accordance with an obviously Known or observed 3.4 5 6 arbitrary procedure. It was assumed that the populations Estimated 14.0 10 13 were stable on all areas, i. e., that reproduction added Other illegal kills Known 25 percent to the spring population and that annual drain 2.0 2 2 Estimated C 0 4 2 1 removed this number by the following spring. Potential illi~galkills Since these computations were based on the assump- On hunts 8'7 0 I10 183 tion of a 25 percent increase through reproduction, the Other 174 0 220 333 "total population" was considered to be five times the Legal kills lost annual reproduction which was considered to be the Known 3 2 1 5 same as the annual drain. Hence "total population" was Estimated 9.1 5 11 computed as five times the annual drain. Other mortalitie5 Observed 4 5 While these assumptions are rather gross, they are Estimated 12 0 14 3 1 nevertheless reasonable and well within the "ball park." Legal kills and collections Since there was no evidence of starvation or disease Archery hunt 2.6 30 4 during the survey year, no allowance was made for it. Buck hunt 33 0 48 6 1 Also there was no attempt to include consideration of Either-sex hunt 39 0 47 53 Other collections .9 0 0 prenatal mortality or immediately postnatal mortality. . - Total 77.0 125 118 Total drain PREDATION BY DOGS Known Estimated There are some skeptics who question the ability of Computed deer population dogs to kill deer. Some claim that they are capable of based on known drain killing only fawns or pregnant does or deer that have Fall population 457 695 777 been wounded by hunters or incapacitated by disease Acres per deer 58 60 60 or parasites. While there may be reason, in some cases, Computed population based to question reports even of "known" kills by dogs, we on estimated drain Fall populnt~on 682 805 1,095 have too many reliable reports of observed kills of Acres per deer 30 38 47 healthy deer to discount dogs as predators. In addition, Annual drain ctrmposition many deaths by car, train, fence, drowning and cold (based on estimated drain) water shock may be attributed to chasing by dogs. Checked out legal kills 5 2 Most dog kills occur in the mountain region of North 1,ost legal kills 6.6 Illegal kills on hunts 9.6 Carolina whcre swamps and lakes are not available as Other ilicgal kills 6.7 means of escape (table 1). However, each year we receive Killed by dogs 9.5 reliable reports of a few deer being run down and killed Killed by other prcbclators 5.1 R?iscellaneous kills 9.7 by dogs on eastern wetland wildlife management area -~~ hunts where the use of dogs is allowed. (Use of dogs 'Figures rn colti~~in;ire niiiiibcrs adult blrcks. adult docs, and fawns. in hunting deer is not allowed in the central and western lncludps fences, cars. diseasc~,cliffs, tr;iins, sttidy coilcctions, drow~iings and cold wnlcr shock and predation by and fox. parts of North Carolina.) None of the several hundred dogs controlled on western contrary, most "strays" were described as appearing to North Carolina areas were accompanied by their owners be recently separated from human ownership. but 62 percent of them showed signs of being owned and Dog ownership was even more evident on eastern areas even the "strays" showed signs of domesticity. In where most dog control activ~lyoccurred durlng the questioning area managers about this point we attempted hunting season. (Use of dogs is allowed in deer hunting to determine whether any of the "strays" could be con- in this section.) Only about 5 percent of the dogs were sidered as true wild dogs that had been born and reared strays and 89 percent of the dogs were picked up during in the wild and had developed into a special breed com- the hunting season. Thus, dog predation is not a year- pletely independent of civilization. None of the area round threat in eastern swamplands In strong contrast, managers could certify to this development. On the however, dog activity, as evidenced by season of control, was practically uniform throughout the year on western quartered on the area rather than in town several mile areas (table 2). away. And 11 also demonstrates the importance of havin, legal authority to exercise co~itrol. Table 2.-Pel-cei~t of stray clogs c.ontl-olled, by One furtl-~eraspect of dog l>redation should bc con scn.son oj yerrr, oir Nortlz Carolincl sidcrcd, namely that of ownership, for herein lies thl wildlife rnanrcgen~eilt areas u~lzel-e crux of the problem. If dogs were wild animals rathe dogs are or clrc not ullou;ed in deer than personal progerty it would be a relatively eas: I~?rizti?ig matter to bring them under control. But the fact tha Dogs dogs are personal ljroperty precludes some very cffectivl )og" Season control procedures. And the fact that they are mas owed allowed produced by people on a "sustained yield" basis, an( not only allowed but actually encouraged to roam un - - - Pel ct.nt - - - controlled, inagnifics tremendously the problem of dee Spring 5 25 restoration. Summer 6 17 However, these circumstances also identify the solutior Fall 53 32 to the problem, i. e., cooperation of dog owners by keep ing their dogs confined. Thus, the best tools for contro Winter 36 26 of this problem include magazine and newspaper articles 100 100 radio arid personal commurrication. Also llelpful is cour prosecution of people who allow their dogs to run dee: where prohibited by law. Since actual control of dog: On eastern areas most dogs controlled were deerhounds by wildlife protectors is limited to designated wildlift (85 percent) w~ththe remainder about equally divided management areas, establishment of deerherds by over between beagles, blrd dog5 and "mixed" breed5 On flow into the surrounding areas is virtually impossible western areas 51 percent were inlxed breeds and 28 if the local people are not sufficiently concerned tc percent were hounds Most of the remainder were control their dogs. beagles, shepherds, and collies Eighty-three percent While most of this section on Predation by Dogs i: were in the act of ehas~ngdeer when they were con- based or1 data from western North Carolina wildlifc trolled and 9 percent had actually cornered their quarry management areas, data from other States indicate tha and were in the act of kill~ngor feeding on them In the problem is not peculiar to North Carolina. Replie: most of these cases the deer were saved and only 37 to questionnaires indicate that this is also a seriou, "known" deer kills were listed. They included I0 fawns, problem in parts of Georgia, Virginia, Tennessee, Okla 17 does, and 10 bucks. 'The number of "known" dog kills homa, South Carolina, and Mississippi. per area varied from zero to nine and averaged 1.6 on western areas. PREDATION BY BOBCATS On intensively protected wildlife management areas in western North Carolina, the average annual loss of Bobcats are the third most important predator-afte deer to dogs per 15,000 acrcs is 1.6 known, 10 estimated, man and dogs--on deer in the southeast (table 3).Thirtee~ and 37 potential (without dog control). The reliability "known" kills were reported for 1968 by North Caro of the figure for potential kill is of course a matter of lina wildlife area managers and 151 "estimated" kills. Re opinion. I will only observe that, while predation is ports from 10 of the 11 States responding to our question pretty much an around-the-cloclt process, the area man- naire indicated an "estimated" 201 deer kills on 51 area: ager's various duties-not to mention his need for a aggregating 2,906,985 acres. This indicates that thi night's ileep--keep him from hearing more than a frac- bobcat has been able to s~~rvivein good numbers ir t1on of the dog races remote localities throughout the southeast. It woult To furthe1 explore the impact of thii level of predation, appear that most "wild areas" of 10,000 acres or morc let us cons~dertwo hypothetical examples Consider fir5t have at least a few bobcats. a 15,000-acre area with a deer population of one per While some may question the ability of a 25-poun( 50 acre5 or a total populat~onof 300 deer and at a 25 bobcat to kill a deer several times its own weight, ther~ percent reproduction rate an annu,rl increment of 75 deer has been a sufficient amount of testimony to this effec per year In this case J. dog predation i'ite of 37 deer from our own personnel to convince us that this doe per year would remove 50 percent of the annual incre- take place. Furthermore, review of the survey figure ment. Next considrr alrot11c:r 15,000-acre area, less re- indicates that this has the potential of being a morl motely located, closer to hurnan Ilabitation, more heavily serious problem than commonly suspected. Whilt infcsted with free-running dogs, that has been stocked "known" bobcat kills of deer averaged only 0.4 pe with 50 deer. It is immediately obvious that uilless dog 15,000-acre area in western North Carolina, and thi control is initiated prior to stocking, the deerherd will Sigurc: is only one-fourth the value derived for "known have l~ttlc,if any, chance to exist, much less inultlpiy dog kills, many of the dog kills were the result of race and expand intercepted by the area manager. In contrast, kill It is also obvious that undcr such circumstances, con- are silent and usually in more remote localities thal trol of free-running dogs is the most important single dog kills, and furthermore eats usually cover their kill f~inctioirof the wildlifr area manager. These considera- thus malting t,hem even less likely to be discovered b tioirs also undcrllne the importance of his bc~nghead- man. Thus, it is not at all inconceivable that cat kill Table 3 -Annual drazn of deer on wzldlzfc management ureas zn southcc~sternUnztcd btutes (us cslzmcrted by arrn man- agers) -- 1 1 / ~LterklllsI Predator kills I- - Mlicellancoui lcills

State areas -- Arkansas Florida Georgia Kentucky Louisiana Mississippi Missouri North Carolina Oklahoma South Carolina Tennessee Totals 79 3,304,315 9,368 2532,829 885 352 32 3 35 9 724 57 33 23 15 11 10 5 2 32 -- - - Percentages 63 2 20 6 2 0% 5 1% Grand total of deer drarn, all causes-14,678

could approximate and even exceed dog kills on areas only one of the five Florida area reports showed the where the cat population is not kept under control. species present, with an estimated population of only five The potential kill by bobcats may be further explored animals. by considering that on those areas where cat control While hears are much more widespread, only a few has been exercised the annual take of bobcats has ranged North Carolina area managers listed them as potential from one to 15, for an average of 4.5 per 15,000-acre predators. Only one "known" kill was rcported and area. On a few areas trapping has netted an average another report indicated fawn hooves in the spoor of annual catch of six to 10 per year. Thus, if annual a bear. The relatively low population of bears and the reproduction of cats averages about 25 percent, the fact that their season of heavy feeding is in late summer normal November population could average about 40 leads us to discount the species as a serious predator of cats. The average of western North Carolina arca man- deer. In our opinion, bear feeding on deer is probably agers' estimates of cat populations is a very reasonable limited to carrion consumption. 43 per 15,000-acre area. Foxes were listed as suspect by scveral North Carolina We are thus led to the conclusion that an uncontrolled area managers but none reported any "known" kills. bobcat population cotrld increase to the point that it con- Similarly, were listed as predators by area man- sumes about half the annual reproduction of a 300- agers in several other States but none rcported any deer animal herd with a 25 percent reproductive rate on a kills by them. 15,000-acre area. This level of drain, in addition to that Six of eight Arkansas wildlife arca managers reported clue to stray dogs, would be capable of wiping out an substantial populations of but only two of ihe established herd and certainly capable of preventing six indicated predation on deer. In one case an estitnated the establishment of a new herd. population of 170 wolves was shown as killing an esti- In contrast to stray dogs, which are domesticated mated 20 deer on a 250,000-acre area. On another slightly animals rather than a part of the native fauna, bobcats smaller area the population was estimated at 185 should be considered to have a rightful place ctn wildlife and their predation on deer at 12 There were no cases management areas. In addition to the interest they of "known" kllls of dcer by wolves rcported engender, they also serve a useful purpose in removing weak and diseased animals, thereby preventing disease MISCELLANEOUS MORTALITY outbreaks. Reasonable control could probably be exerted by classing them as game animals and restricting hunting Deer mortality by running into motorized vehicles to declared open seasons. was reflected at significant levels in most States. It is felt that this type of rnortality is probably more obvrous than that due to other causes and it is llkcly that PREDATION BY OTHER PREDATORS these mortalities are much hrghcr on less remote dcer Predation on deer by panthers in the southeast appears range than that of wildlife management areas covered to be limited to the State of Florida (table 3). However, by this study. Of 912 dcer mortalities due to miscel- laneous cases in 11 southeastc?rn States ( table 3 I. 73 'I'hus thc total tlr;\in clue to illegal hunting and ul percent were attributed to cars, 12 percent to fenlei, retrieved kills was estirnateti to average 31.1 deer pt 5 percent to trdins, 2 percent cdch to frilllllg off cliff5 15.000 acrcs per year. This amounted to 40 percent t and tick bite, 1 percent each to drowning and tiiwaic, the checkcd out legal kills which averageci 77 per 15.OC 3 percent to unknou~ncause\, and a trace each to told water sh~ckand study collections 7';iblc 4. --Anrtz~al drain of deer on individur~lu;ildlij The overall average of thew deaths on all ioutliea5tern n~anc~gcinci~tctrecis in sov.tl~cnslern Unite areas was 4 6 per 15 070 acrcs pcr year In a herd of Stales ihclsed on estirnrtfes of urea ~~~unngc:r! 300 animals rrproclucing at the rate of 25 percent per year this would amount to a little uvcr 6 perccxnt of the Pmdat~onby 1 xisc Iiuntei kills State -- - annual increment Wh~lethis average value i.i not eille- I Dogs / Other mortality Illcgal / Legal cially significant, individual values on some arcas were very substantial. For example, 12 known car mortalities on a 7,000-acre area in R4issouri made up 37 percent Arkansas 5 55 0 6 34 of the annual drain. In Arkansas, miscellrineous mortali- 0 25 2 18 55 ties, mostly due to cars, accounted for 16 percent of the 0 10 3 24 62 annual drain on eight reporting areas. On a 42,000-acre 19 0 23 58 0 area in Tennessee, miscellaneous mortalities accounted 0 0 55 45 0 for 26 percent of the annual drain. They accounted for 0 0 10 31 59 27 percent of the annual drain on a 100,000-acre area in 0 0 2 52 46 Mississippi, and 30 percent on a 14,000-acre area in 2 4 33 7 54 Oklahoma. Virginia data showed a statewide loss to Florida Trace 0 17 4 7 9 vehicles of 1.502 deer in 1967. 1 2 10 25 62 6 2 6 15 71 ILLEGAL KILLS AND UNRETRIEVED LEGAL KILLS 3 5 2 29 62 Georgia 2 2 1 12 83 Deer hunting on North Carolina wildlife management 10 5 0 17 68 areas covered by this report is by daily permit and 3 0 3 20 74 hunters are required to subrrrit their bag for examination 30 1 0 12 57 when they leave the area. Hence a complete record is 14 0 12 12 62 secured of all legal kills. The areas have well developed 11 0 0 5 84 road and foot trail systems in which the overall objec- 2 0 2 10 86 tive of having no part of any area more than bh-mile from developed access has been brought to near accom- Kentucky 3 0 20 13 63 plishment. Thus, it may be presumed that there are 2 0 6 48 44 relatively few unretrieved kills. 0 0 0 25 75 Illegal hunting is held to a minimum by intensive 10 0 14 14 62 patrolling, at the average rate of one wildlife protector 3 0 15 25 56 per 15,000 acres. Boundaries are painted and posted Louisiana 0 0 1 2 96 with appropriate signs and large entrance signs are 5 9 4 8 74 placed at major points of entry. Hunting and use re- 0 0 1 1 98 strictions are substantially more severe than those apply- 0 0 0 4 96 ing to adjoining "nonrefuge" public lands. Enforcement of hunting regulations on "nonrefuge" lands by "county" Mississippi Trace Trace 7 25 69 protectors averages about 200,000 acrcs per protector. 26 0 27 15 32 Thus, intensity of protection on the management areas 23 0 23 6 48 is about 13 times as great as throughout the State gen- Missouri 0 0 0 5 95 erally. 3 3 5 2 88 In spite of this intensive development and protection, 3 0 37 29 31 there is a substantial loss to illegal hunting throughout Oklahoma 24 9 I I 37 18 the year, and to illegal kills during the hunting season 10 0 30 44 60 (table 1 ). In addition a substantial number of legally 2 3 11 4 80 killed deer are not retrieved. On 23 manned areas in the western part of North Caritlina averaging 15,000 South Carolina 25 5 0 54 17 acres each, an average of 4.3 people were arrested per 3 1 7 4 84 year for hunting deer illegally. The "known" loss to 5 5 6 9 75 illegal hunting other than during the hunting season 4 2 5 7 81 averaged 2 0 and the "estimated" average loss was 6 0 Tennessee 0 0 1 36 63 deer per year The "known" loss due to illegal deer 15 0 1 15 70 k~llsduring the hunting season averaged 3 4 per area 9 6 17 10 58 and the "estimated" number was 14 The "known" loss 3 0 26 10 61 due to unretrieved legal kills averaged 3.2 deer per area I 0 4 10 85 and the "estimated" loss averaged 9.1. acres. This is also three times the loss to stray dogs listed by area managers was the bobcat but some also and five times the loss to bobcats. Thus, more deer are suspected thc black bear and foxes. wasted by sloppy and illegal hunting than through both Dogs and bobcats were listed as t.he prin1;lry preci;itk)rs dog and bobcat predation comhined. Man is not only the in the othcr States r~sponciing to o:ir questionnaire. most effective predator but also the most wasteful. Wolves were cited in a few western States of the south- east region and coyotes were suspected in scveral. It LEGAL HARVEST is interesting to note that only one State ~Floridailisted Legal harvest of deer on southeastern wildlife man- panthers. agement areas open to hunting accounts for only a little Other causes of mortality were cars, fences. trains, over half of the annual drain ( table 3). On some areas drowning, cliffs, ticks, cold water shock, anti disease. it is estimated to be as low as 14 percent of the annual No States listed starvation. drain while on others it is estimated to be 98 percent. From table 3 we find that hunting by man accounts On about three-fourtl~sof the areas it ranged between for 85 percent of the annual drain but only 63 percent 40 and 80 percent of the annual drain. is legal harvest. The other 22 percent is taken illegally We feel that one of the primary objectives of manage- or lost in the woods. Predators account for an average ment is to insure that as much as possible of the annual of 8.5 percent of the annual drain with free-running drain be in the form of legal harvest. In examining the dogs getting 6 percent and bobcats only 2 percent. Mis- data from areas showing low percentage of legal harvest cellanecus mortalities account for 6.3 percent with cars the major competing factors were cited as illegal hunting killing 5 percent and the rtmainder succumbitig to a and stray dogs in all cases and, in addition, highway variety of causes. kills in some cases, and other predators in others. The percentage composition of annual drain indicates In view of the fact that only a little over half the that the pattern in regard to predation is rather varied annual drain is consumed by legal hunting on these on individual wildlife management areas in various intensively protected areas, it is interesting to speculate States (table 4 1. Estimates of loss to dogs range from what proportion is taken by legal hunting in lcss pro- 0 to 30 percent of the annual drain. Estimated values tected localities. In regions where hunting regulations for miscellaneous mortality ranged from 0 to 55 percent are not sufficiently stringent and where the protection and their individual values averaged about the same as staff is inadequate, legal kills may constitute less than those of losses to dogs. Estimated losses to wild preda- 10 percent of the annual drain. tors also ranged from 0 to 55 percent of the annual drain In our opinion an "ideal" ratio of legal kills to annual on individual areas but they averaged only about one- drain would be about 85 percent, with about 10 percent third of the magnitude of losses to dogs or miscellaneous going to unavoidable kills such as vehicle collisions and kills. another 5 percent going to wild predators such as bobcats. SUMMARY COMPONENTS OF ANNUAL DRAIN A questionnaire survey of 79 wildlife management Man, of course, is the primary predator and the extent areas in 11 southeastern States indicates that predation to which we can restrict annual drain to licensed hunters on deer is largely limited to free-running domesticated marks the degree of success of our management. It is dogs and to bobcats. Our analysis of data received indi- somewhat startling to note, therefore, that in spite of cates that the dogs do three times as much damage as intensive protection, legal hunter harvest constitutes only the bobcats and that their combined impact accounts a little more than half the annual drain on North Carolina for only about 8 percent of the annual drain. However, management areas. the data also indicate that when no control is exerted The primary predator, other than man, in North Caro- these two predators could effectively preve~itherd estab- lina, is the uncontrolled dog. The only other predator lishment or increase. Some Considerations for Diseases and Parasites of White-Tailed Deer in the Southeastern United States Frank A. Hayes and Annie K. Prestwood ' School of Veterinary Medicine University of Georgia Athens, Georgia

Accounts of cutastroplzic wlzite-tailed deer nzortal- reportedly were the first to observe unusual number ity in the southeastern United Slates PI-ovided the of bloated deer carcasses along streams. Untold nun~ber impetus jor establishing u regionczl u)ilcllifc. diseuse of deer were lost throughout the mountainous sector diagnostic and research service. This joint-State of the southeast, and mortality on some managemer orgu~zizutiorzis described with ernplzccsis on a phil- areas was estimated to be in excess of 90 percent. Cor osophy thut pertains io diclgnosing disertses of wild cern and consternation naturally were precipitated, bu animals. 'l'welo~ fundc~mental cattscs of xildlife before positive action could be taken, the mysteriou morbidity or mortality ure presented. Deer popi~la- killer vanished as suddenly as it had appeared. In it tiorzs of the so~~thea.stcan conlilzue to thrive even wake. heavy losses had been inflicted and many hope though they ure c~ffected,by nzcLny diseases. The were dampened. real threat to deer is unticipati:d to be from foreign During the early 1950's optimism was restored, an, shores as an injectio~~sentity thut currently does regional deer restoration programs flourished. It wa not exist in tlzis country. The Southeastern Asso- evident that white-tails would return to their one ciation of Game und Fish Commissioners fully prom~nentposition In the soc~aland economic structur recognized this potentic~lity,and preventive meus- of the southeastern Un:ted States, and the 1949 kille ures were enacted by game and fish agencies was soon forgotten The pcr~od of reassurance wa throughout the region. Merger of wildlife and do- short-lived. mestic unimal interests are disczcssed. Similar efforts In 1954, at the same time of year, KILLER X struc: huve not been inaugumted on a nationwide busis, again; then in even greater int.ensity only 12 month but inference was rrzade that progrcss is u~zderway later, in the late summer and fall of 1955. Heavy dee for accomplishing this finc~lobjective. mortality occurred from the Appalachians.. into th Scattered records from preconservation years in the Ozarks, but once again the strange marauder defiel southeastern United States suggest that white-tailed diagnosis, leaving only skin with bones and much cor deer (Odocoile~~svirginiunusi expericiiced numerous jecture. "die-offs" from unknown causes. Following the Civil Sportsmen and game officials alike had skirmishes War era, however, wantoll slaugllter of these animals ellough with the sitlister intruder, which three occ,- far exceeded deaths from natural phenomena, and at sions had gained entrance without provocation and lcf the turn of the century o~ilyisolated remnants of this without notice, A serious threat therefore was recog one-time great resource remained. nized, and a regional approach was conceived as th It was not until after World War I that serious con- most expedient way to cope wit11 future misfortunes o sideration was given to restoration of white-tailed deer this type. in this region; and not until after World War I1 that As a result of untiring efforts and encouragement b, earnest efforts were inaugurated to accomplish this goal. many individuals, the Southeastern Association of Gam Immediate results were limited but spectacular, with alId Fish Commissiollers approved a joint-state orgal: many inctividuals and agencies rightfully proud of the izatioll to be supported on a pro rats share basis. Thu progress made. on July 1, 1957, these collective endeavors were realize' Perhaps the first authentic diseclse threat to this big and designated as the Southeastern Cooperative Dee game animal restoration program occurred in the late Disease Study, headquartered at the Univcrsity of Geor summer and early fall of 1949. At that time, Sishermen gia's School of Veterinary Medicine. --- ' From the Southeaster~~Coopcrativc Wildlile Disease Study, Tfcpart~i~ctitof Pathology and P;irasitology, School of Vetclrinar Medicine, University of Gcorgia, Alhcns. This 1s Lhc first rc,gional diagnostic and research servicc c?stablishod in the Unitel States for the specific purpose of investigating disc'ases of game animals. Participating Slates include: Alabama, Arkansa! Florida, Georgia, Kentucky. Louisiana, Alaryland. Mississippi, Nortli Carolina, South Carolina, Tennessee, Virginia, and We. Virginia Study sponsored and coordinated 1indc.r ;i~~spiccsof the Southrnstern Association oi Game and E'ish Con~missioner: the Federal Aid in W~ldlifeRestoration Act (50 Stat. 917); ancl through Contract No. 14-16.0008-777, Bureau of Sport Fisllcric and Wildlifv. U. S. D(~pa~-tmcmtof the Interior. Thc authors' indeblcdnc.ss cxtcl~dsso widely thut standard acknowlcdgerncnts could not approach an ndeqiintc coverag of all persons and agcncics who hrr\~cmadc this report possible. 'I'his acadcrnic courtesy had to be furgonc, whcrc~by th autf~orsrc:spectfi~lly commit themselves to the rilagnallimity and understanding of inany wildlife biologists, conscrvatio officers, and game officials througliout tllc sout11c;istern United States. I~lvaluablcinformation also ii;is been drawn from Proceedings of the First White-T;?ilcd Deer Disease Sylnposium hell in 1962 at tlle Univcrsity of Georgia. Further indcbtedliess is expressed to Mr. Leonard E. Foot?, Soulhcasterrm Representative of tlle Wildlife Managemt7r Irlstitrltc, Dr. C. W. Watson, Federal AI~Supervisor (Ret.), Region 4 of the Bureau ol' Sport Fisheries aud Wildlife, and innu mcr~iblcother individuals and agcncic?~:ictivcly involvcd in the e;irly concepticin of this regional study. Speciz appreciation is c:xtended to the Congrrss of tlii. Ullited Statcs lor making possible the basic 1.csc~arclhfrom which much data an many concepts liavc been procured for inclusion in this report. Shortly after inception of this pilot program, a most 12 to three or four most likely factors. Although this disheartening experience for those involved was realiza- is rather elementary, we often have found it advantage- tion of the relative dearth of information on diseases of ous in minimizing lost motion upon arrival at the des- white-tailed deer. In comparison to the wealth of data tination. A resume' of this approach therefore is prc- pertaining to human health and livestock diseases, the sented for consideration. few individuals of that day working with wildlife dis- Anomulzes-Sign~flcant mortal~tydue to congcn~tal eases were essentially "functioning in a vacuum." Ac- or genetic abnormalities is not expected and usually celerating pressures also were being exerted for procur- confined to an occasional animal. Under normal cir- ing of vital information on disease interrelationships cumstances a subject of this kind soon succumbs to the which may exist between wild animals and man or do- rigors of environment. The history of the area fre- mestic livestock. quently will reveal similarly affected subjects. Condi- The Southeastern Association of Game and Fish Com- tions of paramount concern include: cleft palate; cauli- missioners readily appreciated the increasing demands flower antler; wooly coat; piebald; opaque cornea; lack upon the newly created Southeastern Cooperative Deer of rods and cones; lack of iris and lens; congenital cata- Disease Study. The title subsequently was changed to ract; lack of eyeballs; cndershot mandible; curled ten- the Southeastern Cooperative Wildlife Disease Study, dons; hydrocephalus; two-headed fetus; antlerless males; thus encompassing all forms of wildlife, and the annual antlered females and hypogonadism. Anomalies are rare pro rata budget was increased proportionately. Through in wild animal populations and exert little impact on the interest and efforts of the Southeastern Association deerherds in the southeast. Anomalies usually are of of Game and Fish Commissioners, in 1963 the Congress academic interest only. of the United States enacted an annual appropriation for Stress.-Varying degrees of mortality may be asso- support of basic research delineated by this regional ciated depending upon the nature of stress or stressors. organization, with funds to be administered and research History and study of the area involved frequently will coordinated by the Bureau of Sport Fisheries and Wild- reveal source(s) of stress which may include: adverse life, U.S. Department of the Interior. weather conditions such as prolonged drought, deep Significant means thereby were provided for closing snow, high water, etc.; excessive dust, usually localized; the dangerous gap in information relative to the expand- poor nutrition; chronic toxicity; chronic infectious dis- ing association of wildlife with domestic animals and ease; chronic ecto- or endoparasitism; and head in- man. Concomitant with the collective efforts of 13 jury resulting in a brain abscess. Stress is considered southeastern State game and fish agencies, many other to be a specific response to nonspecific stimuli, but individuals and institutions throughout the region be- population pressures alone are not a mortality factor came interested and actively engaged in similar research for deer in this region. The stress syndrome usually re- programs directed toward fathoming the multitude of flects other conditions and is of only academic interest mysteries shrouding wildlife diseases. relative to white-tailed deer mortality. Although within the last decade a tremendous amount Trauma.-This frequent cause of mortality often is of progress has been made in this field, the surface of manifested by discovery of occasional carcasses over adequate knowledge has as yet only been scratched. It a large area. Sometimes trauma cases may be concen- must be recognized, however, that inclusion of even trated, giving the initial impression of an infectious the limited data now available far exceeds the scope disease entity or toxic condition. A careful investigation of a manuscript of this type. Therefore only the high- in the immediate vicinity frequently suggests a source lights of 10 years' experience for projection of a practical such as: collision with an automobile; collision with a approach toward investigating, controlling, and pre- wire fence or similar object; gunshot wound; previous venting deer diseases will be presented. fighting during rut; and occasional attacks by dogs. The Upon receipt of a State call to investigate deer mor- most common traumas of wild deer result from automo- tality, we first consider population density of the ani- biles and light caliber gunshot. Only the latter thus far mals affected. We then must be keenly concerned with has been incriminated as a source of major deer mor- the association of diseased animals and other fauna in tality and usually this is restricted in location. the area. The location within the region is equally im- Suffocation.-The degree of mortality usually reflects portant, with consideration given for soil types, drainage the type of asphyxiation. I-Iistory of the area and careful systems, recent weather conditions, season of the year, investigation frequently will reveal the nature of in- and past experience in similar areas. With these factors volvement, which is usually: drowning; collapsed trachea in mind, augmented by case histories obtained via tele- resulting from trauma; or verminous pneumonia. Of phone or other communications, we then engage in an these, drowning and collapsed trachea are rare among initial process of elimination based upon what we cur- white-tailed deer, but under certain circumstances ver- rently consider 12 fundamental causes of morbidity or minous pneumonia can inflict widespread unthriftiness mortality within a wildlife population. These are item- and substantial mortality. ized as follows: Anomalies, Stress, Trauma, Suffocation, Neoplastic diseases.-Sigtlifica11t mortality among Neoplastic Diseases, Toxicologic Diseases, Nutritional white-tailed deer has not been attributed to neoplasms, Diseases, Viral and Rickettsia1 Diseases, Bacterial Dis- but quitc a variety of tumors has been reported. Occa- eases, Mycotic Diseases, Parasitic Diseases, and Senility. sional possibilities in this regard include: adenoma; While an investigative field team is en route, it is tumor of adrenal cortical type cells; fibroadenoma; fibro- routine to carefully critique the above potentialities. ma; lymphangioma; lipoma; osteoma; carcinoma; malig- Thus we usually narrow the disease probabilities from nant hepatoma; inesothel~oma;lymphosarcoma; and sar- coma. Skin fibromas of viral origin appear to be by far be difficult, since the causative organisms cannot be the most prevalent tumor affecting white-tailed deer in grown with ease in the laboratory. No doubt there are the southeast. One or two animals so affected can precipi- many viral diseases of white-tailed deer which have not tate much public concern, but from a practical point of been identified or defined. view fibromas are of academic intcrest only. Nucte~iuldiseuses.-Deer are vulnerable to an array Toxicologic diseuses.-Toxicities are capable of in- of bacterial diseases, some of which are capable of inflict- flicting substantial mortality in localized areas, and often ing extremely high morbidity or mortality, especially give rise to overwrought public sentiment. Thorough when deer density is high, facilitating spread of the investigation of the area for a source of poison is essen- organism. Infectious entities within this category are tial. Experiences to date in the southeast involving con- extremely numerous, therefore only a few major offend- firmed poisoning of appreciable significance among wild ers have been selected as follows: anthrax; blackleg; deer include: arsenical cotton dust; pine oil concomitant enterotoxemia; Ieptospirosis; listeriosis; various types with starvation; potassium dichromate plus copper sul- of abscesses; and occasiotlally arthritis. - ~ Of the above, fate; pesticide, including several chlorinated hydrocar- antl~raxhas tlie greatest potential for reaching epizootic bons plus perhaps one organophosphate; petroleum prod- proportions in deerherds of the southeast, since it is ucts of uncertain origin; and fescue poisoning associated particularly prevalent it1 areas of periodic flooding. En- wit11 adverse weather conditions. Poisoning often is diffi- terotoxemia, or overeating disease, also produces rather cult to dlagnose and must be approached with extreme spectacular mortality and is associated with the sudden caution It 1s not as common among white-ta~leddeer ;ivailability of a high protein 'carbohydrate diet following as often thought to be, which rnay suggest a word of a maintenance ration. For differential diagnoses of the warning to the investigator. many bacterial diseases to which white-tailed deer are Nz~lrilior~aldiseases.-Significant deer mortality in susceptible, careful cultural procedures are mandatory. the southeast seldom is due to starvation alone, but Mycotic diseuses.---Fungal infections do not rank a- malnutrition predisposes an animal to other mortality mong the leading etiologic agents affecting white-tailed factors. A direct relationship appears to exist between deer, and known occurrence in the southeast is practically nutritional level and the degree and intensity of para- nil. The followi~lg conditions nevertheless should be sitism, with associated unthriftiness ancl light to severe considered: actinomycosis; mycotic stomatitis; asper- mortality. Diminished reproductivity of a deerhcrd also gillosis; ringworm; and mycotoxicoses, such as ergot or seems related to the nutritional status. Under certain related toxins. Perhaps the most prevalent of the rny- circumstances, a low nutritional level predisposes deer coses of white-tailed deer is actinomycosis, which causes to excessive consumption of available toxic products. a distortion of the lower jaw. Most fungus infections History of the deerherd, management practices, and found in deer are reflected by a general unthriftiness environmental conditions prior to mortality should be and are restricted to isolated cases. Fungi often are carefully studied. Additional factors to be considered secondary invaders to other disease conditions, and care include: soil fertility; mineral deficiency; vitamin dcfi- should be taken in ascertaining the true causative agent. ciency; artificial food-patch planting; abundance of mast of the suspect organism will facilitate diagnosis. on area; climatic conditions; and others which would Parasitic diseases.-I-lelminth parasites are the most make food unavailable. The most direct approach for frequent cause of significant and widespread deer mor- studying nutrition of deer appears to be through actual tality in the southeast. Mortality due to parasitism ruminal analyses rather than field investigations of usually is associated with overpopulation and subsequent browse which can be rather subjective in nature. There malnutrition, although several nematodes are capable also are strong indications that available mast may be pathogens in their own right. At least 30 different far more significant for white-tailed deer than current helmintlls are harbored by deer in this region, but the opinion would have it. Certain aspects of many timber most capable pathogens are: lungworms, including adult stand improvement programs therefore appear to leave and immature forms; large stomach worms; medium a lot to be desired. A new look may well be directed stomach worms; and hookworms. Lungworms or large toward a "multiple-use concept," which is coming into bloodsucking nematodes often cause considerable fawn increasing prominence throughout the region. morta1it.y. Heavy stomach worm burdens usually re- Viml and 1-iclcetlsiul discuses.--11rfectious entities un- flect overcrowding and food shortages. Meticulous der this category are potentially devastating to deer parasitologic examination of deer carcasses is imperative populations and probably were responsible for the pre- since the most pathogenic forms are near-microscopic viously cited catastroplles of 1949. 1954, and 1955. Le- or nlicroscopic in size and are easily overlooked. Quite sions described from animals examined on those occasions often, the most spectacular-appearing parasites are in- strongly suggest epizootic hemorrhagic disease as a prime consequential to a deerherd. Perhaps it also should be suspect. Paramount consider;~tionsfor viral and rickett- mentioned that studies of parasitism among deer may sin1 diseases include: epizootic hemorrhagic disease; prove to be an aid to management. Evidence indicates bluetongue; and vc>sicular stomatitis. Epizootic hemorr- that certain helmintlls arc density dependent, and the hagic disease may prove to be identical to bluetongue. concept of "indicator parasites" may prove valuable. Vesicular stomatitis occurs in cir:er of the sor~thcast,and I11 this regard, both numbers of individual species and the lesions arc indistinguishable from foot and mouth the total number of species encountered vary with host disease. Constant concern therefore must be manifest density. Thus parasites may be used lo reflect carrying for early detection and differeiltiatioil of the above con- capacity of the range, or the nutritional status and feed- ciitions. Diagnosis of viral and rickettsia1 diseases may ing habits of the host. Senility.-Old age is not considered a significant mor- losses of these animals inevitable. Early cletcction fol- tality factor for white-tailed decrherds, hut when it lowed by immediate eradication constitutes the only occurs it is restricted to old does. Little stock is placed recourse for minimizing the anresome consecluenccs of in the "old barren doe" concept, however, since sufficient this type national emergency. observations have been made to demonstrate that oldcr Game and fish officials throughout the so11t11c:is~ern does can conceive, undergo successful parturition, and United States have fully recognized the ever-incrc,;ising raise healthy fawns. Hunting rc?gulations reflected by threat of foreign animal ciiseases, which at any time may proper management will rectify the few losses that are wreak havoc with the Nation's m~~ltibillion-dollargame attributable to senility. animal resources with concomitant impact upon the Upon arrival of a field team at the location of deer entire livestock economy and associated industries. mortality, a general conception of the problem usually Therefore, during October, 1966, the Southeastern Asso- has been established through the suggested processes of ciation of Game and Fish Commissioners adopted a rc2so- elimination. At least, the situation has been reviewed, lution which afforded a merger of efforts betwccr-i wild- conditions appraised, and the more likely possibilities life and domestic animal interests. A modified vcrsion chosen. Caution nevertheless must be exerted to avoid of the resolution is as follows: a "specialty bias," which can frequently creep into field "WHEREAS: At any time a clevastati~lgexotic animal activities. The chief investigator therefore must strive disease can be accidentally or purposefully introduced to be a diagnostician, and not a specialist in any given onto the Continental United States, which could icriously field. Also it should be recognized that seldom is every- jeopardize the entire livestock economy, one pleased or satisfied by the end results that are ob- "WHEREAS: Various forms of game animals can serve tained. If these prior reconciliations are not made, the in the capacity of unrestrained carriers of a foreign road to a successful field operation will be mudded with disease transmissible to domestic animals, i. e., whitc- tears! tailed deer as carriers of foot and mouth disease IFMD); An experienced and well-organized investigations1 "WHEREAS: An enemy of this nation could utilize team should arrive at a sound tentative diagnosis within various methods of introducing a devastating foreign dis- a matter of 2 days to I week. For many conditions, ease into wild deer, which could spread rapidly with considerable laboratory work is in order at the base eventual introduction into domestic livestock; of operations, but with proper facilities and adequate ingenuity on the part of supporting staff, a confirmed "WHEREAS: Tremendous expansion of white-tailed diagnosis no longer is shrouded by the mysteries of deer populations has placed a virtual blanket of these yesteryear. animals over the southeast, thus affording an unbroken chain of susceptible animals through which a foreign It is hoped that oversimplification of a major problem disease could spread; is not suggested by this approach in investigating deer diseases. In fact, as a result of drastically changing "WHEREAS: Early detection of a highly infectious socio-economic factors predicted for the next three dec- entity is absolutely mandatory for the continued welfare ades, disease problems involving white-tailed deer will of certain big game animals and domestic livestock; greatly increase, and demands will be intensified for "WHEREAS: Animal Health Division officials of the procuring information on the many interrelationships Agricultural Research Service (ARS), United States De- that exist between swelling dcer populations, human partment of Agriculture (USDA), offer to provide game health problems, and production of domestic livestock. and fish personnel with training necessary to participate The relatively new billion-dollar white-tailed deer in a program designed for early recognition of potentially industry of the southeast nevertheless will continue to dangerous diseases; thrive regardless of any disease or vectors thereof cur- "AND WHEREAS: In the event an exotic disease is rently existing on the continental United States. Al- suspected, the Animal Health Division (ARS, USDA) though there will be setbacks at local levels, and on will afford diagnostic services essential for early detec- occasions entire deerherds may have to be destroyed, tion, with control measures thereafter being delineated this big game animal resource will survive any condition in accordance with accepted procedures based on valid presently known in this country. scientific data; The major disease threat to white-tailed dcer and "THEREFORE BE IT RESOLVED: That the South- other Cervidae of this country therefore is not from eastern Association of Game and Fish Commissioners within but from without, in the form of a devastating support an exotic disease surveillance program in coop- foreign disease. For example, authorities in the field eration with the Animal IIealth Division (ARS, USDA)." no longer use the word if but when foot and mouth dis- This vital alliance between game animal and domestic ease (FMD) is reintroduced into this country. livestock interests was officially enacted July 19-21, Greatly accelerated military/'tourist~businesstravel, 1967, when the Animal Health Division (ARS, USDA) increasing demands for importation of meat and byprod- sponsored a regional Foreign and Emergency Disease ucts thereof, worldwide use of biologics, etc., pose an Surveillance Training Program in response to the above immediate likelihood of accidental foreign disease intro- resolution. The program was coordinated by the South- duction. The purposeful introduction of a devastating eastern Cooperative Wildlife Disease Study of the Uni- pathogen also cannot be ignored, which would be capable versity of Georgia's School of Veterinary Medicine and of exerting a tremendous impact on vital segments of held at the Georgia Center for Continuing Education. the Nation's entire economy. White-tailed deer subse- Participants included game officials, biologists, and law quently would be intricately involved, with staggering enforcement personnel from 15 southeastern States: Ala- bama, Arkansas. Floritl;~,Gc~orgi:i. I

" . . . to relate and cn-Iph:isize to Soi~tlieasterti Statos have cornl)lcteti or ;ire in the 1)rocess of planning Game and Fish Field personnel ti;<. full ramifications of similar trtiinil~gsessions at the Strite level. These coil- possib!e I'o~cigntlisctise outbreaks in this country; f'c.rencc.s encompass the full complement of technical and " . . . to describe the position whits-tailed deer law enforcriment personnel; which essentially adds from and feral swine tiow will occllj)y in tile event of foreign 200 to 300 trainecl l-ieoplc per State. These men arc be- disease introduction into thc southetist; rornilig well versed on the full ralnifications of foreign disease introciuction and the necessity for immediate " . . . to famili2rize Game and Fish personnel reporting of tiny suspicious case involving wildlife or with the elaborate n;~tionwitiec:mcrget?cy disease eradi- dorncstic animals. Thus conservation officials and game cation organization of the Animal I-Iealth Division, ARS, biologists are establishing direct communications with USDA, and specify the vital role wildlife interests hcre- Animal Health Division officials (ARS, USDA) and after may play in that program; excellent liaison between their respective State veteri- " . . . to train Game and Fish personnel to imme- narians and diagnostic laboratories. diately recognize and rcport evidences of a possible As a result of these cooperative efforts, southeastern foreign disease outbreak; wildlife interests now arc in position to make paramount " . . . to establish liaison bctwecn attending Game contributions in the eventuality of foreign disease intro- and Fish personnel and the Veterinarian-in-Charge duction. The only regret today is that these cooperative iANH, ARS, USDA) in their respective States for inau- activities thus far have been restricted to the southeast. gurating exact reporting procedures for all suspicious It is hoped, however, that within the near future similar cases; resolutions will be adopted by other regional game and " . . . to provide basic informati011 and visual fish associations, with wildlife and domestic animal aids with which attending Game and Fish personnel interests ultimately combining forces on a national can return to their respective States and relay to co- front. This will be invaluable insurance toward the workers instructions received during the trail~ingpro- preservation of countless thousands of big game animals, grams." millions of domestic livestock, and billions of dollars. After 2 days of intensive lectures with aecompallying Such investment today will pay unprecedented dividends visual aid sessions, a test exercise was conducted, in- tomorrow. Critical Factors in Habitat Appraisal William D. Zeedyk Division of Wildlife and Range Management Southern Region Forest Service, U. S. Department of Agriculture Atlanta, Georgia

In appraising habitat the uppraiser must go be- the impact of deer on the habitat itself. Ilis primary yond enumerating the factors limiting deer nurn- tool being the allocation of land and resources to deer bers and consider what factors control the habitat habitat purposes, he needs to know the relative value itself. Several criteria for judging habitat quality of various inputs in terms of habitat outputs. are presented. Hence, it is apparent that habitat apljraisnl is rather Before discussing the critical factors in appraising pointless without having first a clear statement of deer white-tailed deer (Odocoileus viriginianz~s) habitat in management objectives. How will appraisal data be the South, let us consider for a moment how habitat used? Is a sustained yield of deer the objective or is it quality is measured. Is there a uniform standard of simply the quantity or sport realized? Is the naturalness measure? The biologist will answer, yes, habitat quality or artificiality of tlie habitat of any consequence? What is measured in terms of carrying capacity. But does is the reason for making the appraisal? The answer will carrying capacity alone fully describe habitat value? help determine what to measure. Unfortunately, carrying capacity is not absolute, but SOME CRITERIA FOR APPRAISING HABITAT is relative. It fluctuates seasonally and annually with rainfall, temperatures, mast and browse yields, inter- At the risk of overworking a cliche, the white-tailed specific competition, and to some extent with the size deer is a highly adaptable creature. It thrives or survives of animals supported. Carrying capacity may be the in every forest type in the South. It would be virtually accepted measure of habitat quality, but it is not a impossible to enumerate precisely all the factors limiting very precise one. Appraising habitat on the basis of habitat quality over such a wide range of situations. carrying capacity alone is like valuating a house on the However, the most frequently identified criteria in ap- basis of square feet of living space without regard to praising habitat are: other considerations. 1. Inherent soil fertility. In appraising habitat, the appraiser must go beyond 2. The abundance and variety of palatable forage merely enumerating the factors limiting deer numbers, available to satisfy the seasonally changing dietary and consider what factors control the habitat itself. requirements of deer. Factors controlling the habitat are, for example, inherent 3. The degree of interspersion of food and cover com- soil fertility, floods, droughts, temperature extremes, ponents. the pressure of land use trends, physiography, forestry practices and other factors. Thus habitat appraisal in- 4. The nature and extent of escape cover serving to volves more than counting twigs, calculating acorn pro- reduce legal or illegal hunting harvest and pre- duction or plotting the location of year-round water dation by dogs. sources. Habitat quality cannot be computed, only 5. The resistance of the habitat to severe weather judged. stress. What are the critical factors in habitat appraisal? 6. Water. The word critical, as used in the title of this paper, Inherent soil fertility.-Inherent soil fertility is re- means decisive. What are the decisive factors an ap- flected in the nutrient quality of plant materials con- praiser must measure? Which can he ignore? sumed by deer and ultimately in the physiology of the What may be critical will depend, somewhat, on the animal( s 1. French et al. ( 1956 ) established minimum purpose of the manager-his goals and objectives, and nutritional demands for crude protein and phosphorus the management options open to him. One might ap- for growth and body maintenance of deer. These ele- praise habitat as a guide to managing either the herd, ments are often lacking in southern forage grown on the habitat, or both. infertile soils. Thus, soil fertility may affect such herd The herd manager, whose responsibility is primarily characteristics as population densities, productivity, to the hunter, will consider those factors critical which average weights, and antler clcvelopmcnt. alter carrying capacity, annual fawn production, average Thorsland ( 1967 ), reporting on a nutritional analyses animal weights, antler development, hunter success, etc. of seasonally selected deerfoods from sevcn areas in His primary management tool being harvest regulation, three physiographic provinces of South Carolina, writes: he may need to know how habitat quality varies with "There was a relationship between mineral contents herd density, what temporary conditions, such as in the soil samples and plant mineral content. which droughts, alter habitat quality, or what features of the was especially evident with the mineral phosphorus. A habitat influence harvest rates. poor soil was usually reflected by plants with low min- The habitat manager, whose responsibility is primarily eral values. to the landowner, may be concerned with increasing the "The analyses showed that tlie Broad River Manage- yield of deer from a specific unit of land or determining ment Area i Piedmont Plateau j has the most nutritious plants. In general, plants from this area contained the of southern deerherds. Is it wise to measure browse? highest mineral contents. 'rliis was supported by the If cteer take such a variety of foods (Lay 1967h based average live weight of deer killed on the Broad River on availability, qualily alld preference rather than form, area during 1965-1966 being higher than that of deer what should the habitat appraiser measure? The an- killed on any other area involved. Thc plant species swer is: measure those :isljects of the forage supply from the Reimunt area anti thc Francis Marion National which by their nature lend themselves to reliable in- Forest I both Lower Coastal Plain i were lowest in nu- ventory, which local focld habit studies indicate are im- tritionaI contents. This was substantiated by these areas portant indic;itor items during the sczison in question; having soils with the lowest miner:il contents and the iind which can be correlated with hcrd dynamics, pro- average live weights of clcc:r killed on tl-~cseareas being ductivity, alld animal and range condition. lower than those for deer killed on any othor study area." The following cxcerpts from the writings of biologists Trace elements typically deficient in some soils rnay working throughout the South testify to the above: influence productivity rates. Postulating on the variable Dunkeson (1955 I, Missouri, wrote: "The pattern of productivity of Florida herds. IIarlow and Jones (1965 I deer browsing in Ozark woodland had the following state: outstanding features ( 1 i Green forbs aupplled the "Evidence to date strcingly lnd~catesthat the low re- major part of decr food through the period from March procluctivt. rate of does from the flatwoods and deep to November and were an rniportant pclrt of decr diet sands of central Florlda are a result of the lack of iron throughout the year, ( 2) grasws, ihruhs, fungi, fruits, carbonates and or coii,ilt 111 the ioil and not from an weds and acorns were ~mportant seasonally, (3) a over-abundance of dcer or a lack oi av,nI,ihlc foods nuinber of y~lants. %ere consistently ~lnpaldtable and "This information does not emphatically imply that other5 were palatable for only a short ilme dur~ngthe deer populations, wbere reproductive rates are low, can- growing season. not reach densities approaching populations on good "Little winter use was made of deciduous woody twigs range, but it does indicate that herd increases will occur and evergreens were so low in palatability that these at a much slower rate " types were not good indicators for determining the condi- The inhibiting effect of deficient trace elements on tion of deer range during the winter. the productivity of cattle granng native ranges In the "Shrubs which were palatable for a long perlod during South is we11 documented Mineral dlei ~~~pplernents the grou,ing season appear to he the most sensitive indi- are wldely applied by cattlemen to increase calv~ng cator plants to show destructive overbrowsing. Three . . . rates and to reduce anemia. were browsed to the point of destructiorl on the same The above examplei indlcatc the crlt~calbear~iig that areas where forbs were not clearly affected by overuse." sol1 fertility has upon deer habitat. There are many such references In the literature Fertile, '~lluv~alsoils and soils derived from limestone and metamorpliic rocks tend to support more nutritious plants, a higher carrying capacity and more productive herds than soils derived from infertile, acid shales, sandstones, and co;istal marine deposits. A key step in any habitat appraisal should be to determine how and to what exte~itsoil fertility may limit deer management potentials. The literature is not con- sistent, however, on techniques for rcmcdying mineral deficiencies on an extensive basis. The c~bunclarlceand 7)nricty of pc~lntclbleforage clvail- able lo sc~tisfytise se(~s01ti11lycP~clnging rlic'f~ry req?tire- nlents of dcr~.-Probably thc rnost distinctive character- istic of babitat quctlity is the abrunda~icc,variety and nutritive quality of forage available to satisfy the se;ison- ally changing dietary requirements of dcer. Forage is defincd as all ihe unharvested plailt n~citerialsavailable for animal consumption. Deer were once cotlsidercd to be browsers-feeders upot~the twigs and shoots of shrubs, trees, and vines. But. alas, deer feed with (:qua1 relish upon grass, :trlni~al and ljcrenni~il forbs, fruits, flowers, fungi, ioliagc, and twigs with little regard to the fonn or pcirt of the plant cor~su~neda~ici with utmost regard lo pnIat;ibility, succulence, nvail:ibility, and nu- (li1amr:rd and 13ox ( 19fi8 i reporting the food habits tritional cotitciit of footis ingested, Deer ;ire frugivorous of decr from soutb Texas gr;rssiand-br~rshland complex and herbivorous ol~porlunists,not browsers. say : "Whitc-tailed decr were primarily grazers rather Untold m,ln hour5 ot t tine ,ind moncy 11'11 e been ex- than hrowsc,rs (luring the wintcr-spring pcsriods . . . . pended measurii~g l>ro\vse supplies, i. e., n>ootiy twigs Complcsity of llic rliot reducecl the iinport~ulceof any which furnished but a minor proportioii of the total diet onc or si(:vel,al sljccics in the tlict. Among high priority forage species perennials were more important than The degree of interspersion of foocl crnd covey corn- annuals. Deer food habits varied according to avail- poneizts.---Several authors have commented on the sed- ability and phenology of range vegetation, and were entary nature of southern deer-preferring to stay and further modified by forage preferences." starve than to migrate in scarcll of food. It is generally Harlow and Jones ( 1965) in Florida identified 23 accepted that southern deer do not migrate wit11 the major species or species groups in deer stomachs col- seasons althoiigli some slrifti~~gbetween types within the lected in fall and winter from the flatwoods and sand home range rnay occur. Recent telemetry and tagging pine-scrub oak types. The gamut of plant parts eaten studies have verified the limited range of southern deer. included acorns, fruits, berries, leaves, twigs, grass and beirig from ahout '4 mile to 1'. miles in r;idius and sedge stems, and blades, and the entirety of mushrooms. coveriiig from 200 acres in good habitat with high 11op- Yet in separate studies in the same types, Harlow showed ulations to 3,000 acres in poor habitat. Iieccnt investi- ~ - that in spite of the variety of foods eaten, a strong corre- gators have attempted to show a diminisliilig radius of lation could be drawn with the abundance of acorns and movement with increased herd density ( ;Vlarcl?inton palmetto berries and the weights of deer in the 115 and and Jeter 1967; Marshall and Whittington 1969). Appar- 21.5 year age classes, and the percent of 1% year old ently as populations illcrease, a higher degree of -inter- bucks harvested 2 years later. What should be meas- spersion is needed to compensate for diminished move- ured? ments.

Segelquist and Green ( 1968 ) reporting the progress The foregoing sililply mcnns that qiiality habitat must of the Sylamore stuflies of penned dcer in Arkansas oak- possess a high interspersion of food and cover compon- hickory types substantiated the inverse relationship be- ents, for not only is the daily range of deer small, but tween mast availability and forage usage reported by the variety of foods they require is great. These foocis Korschgeti (1962) and others. Wiien mast was avail- originate from a diversity of plants occupying many able, deer ate little else. When mast yields were low, different sites, types, soils, aspects, age classes, and forage use Increased and forage use way heavier where succesiional phases In the forest. green herbage and evergreen browse were mo5t plentl- Among thr cr~t~calfactors controll~ngthe degrec of ful Important unpubl~shedstudies undrrway at Syla- ~nterspers~onare more may yet link fluctuations in deer reproduction and 1. Physiography, or the distribution of intermingled survival rates with fluctuating mast yields. and the soil types, sites, ar~tlaspects (Ryrne and Zeedyk nutritional quality of native woody browse. What should 1966 ). be measured? 2. The complexity of land use patterns, i. e., agri- Various investigators throughout the South and south- culture, industry, etc. east have seriously cluestioned the value of woody twigs as deerfood while underscoring the value of fruits, a- 3. Forest management practices. corns, foliage, mushrooms, legumes, grasses, and pal- Is it not logical that, given a population at carrying atable evergreens. Woody twig tips arid buds remain capacity, a high degree of interspersion should result importarit only during early spring when growth is in the most efficient use of the low quality elemerlts rapid, succulent and highly nutritious. intermingled with the high quality portions of a range? Goodrum and Reid (1962) and others have dernon- The nuture and extent of escclpe coz.~rs~~ving to re- strated how critical food shortages may occur in late dzccc? legal or illegul hunting Izctruest and predntiolr, by summer when the nutritional value of available succu- clogs.-Little effort has been directed toward defining lent foods is submarginal for deer growth. At this time tllc importance of escape cover in southern deer habitat. the availability of composite flowers and fruits may he In many areas where protection from poaching and dogs highly important. is less than adequate, the presence or absence of satis- Lay ( 1967a sums it up succi~lctly: "A major lirnita- factory escape cover, such as dense. watery swamps, tion of browse surveys is that browse provides less tl~a~i may be a critical factor in survival. Excessive escape half and possibly as little as one-fourth of the deer diet cover is also vexing to deer management. Dense covcr on such fully stoclied ranges . . . . Fruits, including curtails legal harvest throughout much of the Coastal acorns, are more importat~t. Mushrooms ;Ire c:spccially Plain where high density pol,ulations tire common. attractive and nutritions. I-Ierbticcous green stuff is iin- In the Appalachian Mountain, Piedmont and Cumbcr- portalit. Browse, however, has the advantage of year- land Plateau I-'rovinces, insufficient escape cover makes to-year stability. It is more permailerit and more meas- dcer particularly vulnerable to dogging. Secondary urable." losses to disease and parasites and of fawns occur where What should the habitat apl~r:~isermeasure? wcokened deer successfully evade dogs, but do not sur- He should measure those characteristics which arc vivc the ordeal. most readily and most consiste~ltlyilieasur;tble with pre- 'I'lle habitat appraiser should decide wlietiier the cll:ir- cision. He should me;xsure those factors which are den- tictcr of escape cover ;id\:ersely affects either protection sity-dependent and reilect an upward or downward trend or h;irvest, and dcterminc whether protection, harvest in range condition as caused by foragirig pressure of rcgul;itions, or habitat management pr:ictices need to be the herd. fIe slio~~ldmeasure those density-indepen- modified accorciingly. dent itctns which can b(t most reatlily correiatcd with 1i'c'sistaric.c~ oj hnbitnt to sci?ere zuc'ctthel- stress.--Tlie populatiotl dynamics of the herd. He should metistire ability of habitat to withstand the stress of severe those items subject to n~anipulation througii methods weatlit~rarid provide adequate food arid shelter for the at his disposal. duration of need might be ternled its resistance to severe weather. The emphasis here is on abnormal stresses. Biologists of the Ouachita, Ozark. and Daniel Boone Throughout the South, tcinporary severe wcather in- National Forests assumed that artificially constructed cluding floods, hurricanes, snow, freezing, rain, drought. waterholes on the dryer ridges and above barrier cliffs prolonged heat or cold, and unseasonal frosts occasion- would favorably influence the distribution of decr. This ally create periods of stress. Its ability to resist such assumption is essetitially unproven except that such stress is an attribute of high quality habitat wilereas waterholes arc heavily used in early autumn when ridge- the lack of this capability is a Inark of poor hahitat. top mast crops ripen and free water is especially scarce. Usually, but not ~~ecessarily,the mechanism for survival Use is heavy where attractive food plots are nearby. is the cila~icedistribution of limited acreages of "key However, there is no evidermce of an overall increase in areas." A critical appraisal will isolate and identify deer numbers attributable to the presence of artificial those which lend severe wcathcr resistance waterholes. to habitat. Resistance to stress is a function of such divc:rse factors SUMMARY as the availability and nutritive quality of "stored" re- serve forage not otherwise used, elevated sites in areas Habitat appraisal is a complex task calling for an prone to flooding, a source of succulent vegetation in evaluation of many factors. The appraiser should not be areas afflicted by drought, aspects sheltered from killing satisfied with merely counting twigs or fruits producecl frosts, and so forth. and consumed, but should explore those underlying The Iiterature contains rn:u?y refererlces to habitat factors which control t,hc habitat itself. To be meaning- which was or was not capahle of carrying normal popu- ful, habitat appraisal sllould relate habitat condition to lations tlirough brief periods of severe weather, for herd dynamics and vice versa. The appraisal should be example: interpreted in terms of the variables the manager can manipulate or accept as controlling. Burnett ( 1959 ) reports, "The Tensas ( Louisiana ) deer herd was exposed to disastrous floods in 1912, 1913, Several criteria are listed above as critical in judging 1916, 1922, and 1927, especially was this true of the habitat quality. The list is not exclusive and may not flood in 1927; this flood almost wiped out the herd." apply equally well throughout the South. It may prove useful, however, in stimulating the deer or habitat man- Harlow and Jones ( 1965 1, referring to Florida Ever- ager to take a rnore careful look at what critical factors glades, state, "Due to prriodic die-offs of deer ( as a result of high water levels forcing deer on tree islands over really are. extended periods until the food supply has been de- pleted i from starvation the population of 'Glades' deer LITERATURE CITED fluctuates widely." Tl~eabove illustrates how a single limiting factor, lack of elevated areas with adequate food, Burnett, L. E. 1959. The Tensas deer herd. Southeast. Assoc. results in a low resistance to flooding and reduces the Game and Fish Cornm. 12th Annu. Conf. value of otherwise high quality habitat. Proc. 1958: 213-224. The importance of succulence as a feature of palatable forage is generally accepted. In a reference exemplifying Byrne, J. G., and Zeedyk, W. D. how a source of succulent vegetation fortified a habitat 1966. The application of soil survey information against droughts, Chamrad and Box (1968) state, "Deer to forest-game habitat management on the concentrations on the dry lake beds arc most 1,ronouncc:d Cumberland National Forest. Southeast. during periods of severe drought. Under such conditions Assoc. Game and Fish Cornm. 19th Annu. some green vegetation persists for longer periods in tl~ese Conf. Proc. 1965: 174-181. depressions than on the surrounding upland sites." Chamrad, A. D., and Box, T. W. Other exanlples could be shown to illustrate the con- 1968. Food habits of white-tailed deer i11 south ccpt of resistance to weather as a feature of high quality Texas. J. Range Manage. 21 : 158-163. habitat. Dunkeson, R. 1,. Water.--The literatrire is inconsistent on the import- 1955. Deer range appraisal for the Missouri citice of free water to deer. Strode reports little use of Ozarlts. J. Wildlife Manage. 19: 358-364. known water sources during dry periods ill the sa~tdliills of the Ocala National Forest. On the other hand, R'licl~ael French, C. E., McEwen, L. C., Magruder, N. D., and (1968),reporting on the drinking habits of deer at Welder others. Wildlife Refuge, Tcxas, noted the ljrescncc of open water 1956. Nutrient requirelncnts for growth and ant- ~tndor the distribution of succulent, green vegetation ler tleveloprnent in llle white-tailed deer. J. affected the distribution of decr during late surn~ner Wildlife Manage. 20: 221-232. droughts. Michael infcrred from data collected over a Goodrum, P. D., and Reid, V. H. period of 2 years that deer us<, any water available, 1962. Browsing habits of white-tailed deer in drink more in hot seasons than in cold, that some deer the western Gulf region. First Nat. White- lick dew from 1eavc.s and al,p;rrently never drink, and Tailed Deer Dis. Symp. Proc. 1962: 9-14. pregnz~ntdocxs drink rnorcXth;in nonl)rcgli;int deer. EIc noted that the tcxndency of deer to co~lcentrat,c near Harlow, 1% E' , and Jonc.5, E' I< , Jr succulent vegetation in dried up 1;ilce allti river Isottoms lOfi5 'I'lie whlte-tailecl deer in Floriti'i Fla Game rnigllt have a bearing on rnanagcment and influence ~ndFre5li W- 11ii11ts. Souf1ic~:isl. Assoc. (;:\me and Fisii n?cnt irri11lic;itions. J. Wiltilifc Manage. 26: Comm '2nd AII~IU.Conf. f'roc. 15168: 30- 164-172. 46. I,;~v.n. w. ?vIich;~el.E. 11. 19f57;i. 11c.cr rallge ulj11ruis;il in e:isttrrli Texas. .I. l!Jlii3. 1)rinltirl:; hahits of n-l~ite-t;iileci tlccr irl Wilrll~fehT:i11:1ge. 31 : 426-432. sll~ltllx1s Sol1tllc~lst.,\ssoc.. (;:llni. :illti T~LI~,1). W. Fish Corr~r~i.21sl ~~IIIILI.C'olif. Proc. 15367: 19fi7b. The it-n~~ortst?ceof viiriel y lo southern ticcr. 51-57, Sout11e;ist. Assoc. C;;irrie i~ndFish Comm. 18th A~IIIU.Conf. l'roc. 1964: 57-62. Scgc.lquist. C. A..illltl (;rei~ll,W. E 1 1l)eer foot1 yicltls in foiir Oznrlc forclst types. i\/l;irc.ilintoi-~.I<, i,., :inti ,Jctc>r;1,. K. .J. W~lcllifeXf;~i~:igc~. 32: :3:30-3:37, 1967. 'I7eIc.nietric study of rlrcr inovernen? ec,ology in the. So11t11c;ist. Southeast. Assor. Cainc '~~!IIII~SI:IIIC~,0. A, and Fish Comm. 20111 Annu. Corif. I'roc. I.N~~tritioii;il :lnnlyscs of scl'ctetl tlecr foods 1966 : 189-205. in Soi~thCarolina. Soutl~cast.12ssoc.. C;:.irnc Zil;irsl~;ili, A. I)., and Wl-iitlil~gto~i,R. W. and E'ish Con11l-1. 20th 12niru. Cotif. P1.o~. 1969. A tclernetric study of clecr borne ranges 1966: 84-104. Deer Habitat Quality of Major Forest Types in the South

J. J. Stransky ' Southern Forest Exper~mcntStat~on Forest Service, U S Department of Agriculture Nacogdo~hcs,Texas

soicil,c~).njcirc~sf t!jlic,,s. Yic>!rls; c,ort.c~latctl1cil 11. trrii- ~lsc\?:is eaten ~vl~ilct11c.y \vc\re ;~v:iil;ihl~I K~~rscl-igcsi~ f??t~(ri2tl1!ir<~~)t(~riis, 11f.1, intfict~iorsII~ c,(~r~.gii~cq (.!ITI~L- 1 l)(i:! I. 12 recc.ilt stutiy ill cast 'I'clxas slio~vetltihiit ljrc~wsc~ (,if!/, TIL(>J'OILI. rii!rjor forc>,st I!/I)I,S iliJ'jt>r ~ri(T~,!y. ;ivcr:tgotl ijnly ;iboul 30 ljc'rcent or the :iti~iuiildiet; ~icorns, 13otlorit-lalit1 iriit~lic.:i:~ds crrcJ ill(. lic3sl iitriiil(t1s !or miisht-oonls, fruits, gr;issc5s. alitl Sorbs matie up the rest i!~lti~c~-tciilccidcc~. j~ll~z~(~cl bg rl~c' ,sl~ortl~~uj-loi~l~ll~~ ( Short 1%.I,., ~i~i~jiiblislie(i~iii~nuscript i. 12ay ( 1967 I pi?ic. irccrt11~:ood.liic !cl)!nntf htrrdu.ooci. (lnd the? long- hiis :iiso q~lcslionctithe vtiiitiity of rntigc appl-:iisals based leiif-sltrsh pine jorc..sr t!jl~c's. ol11v or1 broxvse quantity. 'rile first stel] in effccstivc, man~ige~iicntof ilrcr 1iabit;it 'I'lic four major forest lypcs are: bottolii-land harcl- is totielcrlihine arrloi;rlt oi load uvailubic, ~~~~c;iyieltis, ivootis, loblolly-sltortlenf pilie-h~irdwoocis.uplancl hard- ,,vhc.lI eol.rclated Lvitlt allimiil rcquil.et,Ic. illt1ica- \vuo(is, nnd longleaf-slnsii piiie (lig. 1 I. They cover 220 tors of carryi,,g caljacity. ~~~~~~h the asscssmcllt of nhillion ;ic.res from Virginia lo Texas i Whec1t.r l!lfi(ii. prot~uctivityby forest tyi,cs w;is ;It In 1965, they harbored about 2 million wliite-tailed tieer, 250,000of which twre lcmlly harvesled. :;O ago ( s~~~~~~~~~1937 ,, rescnrc.ers rcccntly have been :ible to investigate foot1 yields tlior- ougiily. This paper reviews current inl-'orm;itio~~on cleer- Bottom-Land Hardwoods food yields in the four major forest types of the South. Bottom-land hardwoods occupy 38 million acres and neerfood consists of than llerbage alld browse, are the best of all southern forest habitats for white- thorigh most reports cnlphasize these two components. tailed deer. In nearly all Southern States, hunters have fruits and mast crops are also important, taken the most deer in this timber type (Anonymous For example, in a Missouri study, acorns made up 42 19'j8; Stral'sky 1967). - Deer prefer bottom-lands for a principal reason: the ' Thc author is on the staff of tltc Wildlife! I-Iabitnt and Silvi- fcrtile, well-wc~tercdsoils produce more food than upland cu1iur.c I,abor;ilory, whicl~ is n?aint;lincd at Nacogdoches, T~~~~,by tilc ~oLithcrrl~orcst Experilncnt Station in coop- soils. The superiority of bottom-lands to other forcst cration with Stcpiicrr F. Ar1sti11State University. types llas been dcmo~lstratedmany times.

GULF OF MEXICO

UPLAND HARDWOODS

SHORTLEAF-LOBLOLLY PINE-HARDWOODS LONGLEAF - SLASH PINE

Figurc 1.--SOZL~~LU~.TL ~OTIIS~~?/Pcs.

42 In I~oi~isiana,Collins ( 1ljf;l r re.jtortccl th;it bottom-lanci stzrlltls I Schtistcr ;i~itlIInlls 1963 I. 'I'hcsc to;:ils ;ii.i%Liigl?. oirks pro(iuce rnorcx :iror!is lxsr trcxc. i11nli ul11;iiitI o;ilts. Ix~~;iusci!i~ stantis ]];id iitii~suallytie~isc. t~irtl(~isio~~ic~.; (li' Sonic acorns were ;iv;iilahlc~every yciar from bcit11 rotl sliriil~s:illti young 1i:irti~vootis. It sii~~uitibix r,citctl tali o,rki c~~itl\r/hite onc'y Soot-l. Bottorri-lanct f~irc~stsiri ;in Al;ilxinia study contain'ri In tlic: OIeripine-iiartitvor,ti anti r11)l;iriti 1~;irtlwooii forc.st i (;;rinc>s (~f(11. 1054 I. ty~les. 'i'liey ~ioted,Ii~~nc~vc~~~. t1i;it tvilli lc>;if i'iill hrov;scx 12;ry 1 1965 I fount1 that east 7'cxns hottom-larrtl 1i;ibi- d(~c1irictlby 85 percent, to ;~bo~!tIll pouncls 1)er zicri:, 13 tats 11;ive more rn;~st-bearing trees kitid fruiting shrubs 1)ountis of ~vtiicliwere ~irc~fc~rreclhrox5-se. 'T1,c.y corrcititli~ti tiran up1;inti habitats. He c:oncludecl that r;inges with a t11;it low winter hrowsc~;ivtiilahiIity thcre ni;iy lin-iit tlt,ctr populations. Intcnirflc.tl mariClgc~mcntof the lobloliy-illor tlmf ~III('- 1i;irdwotjcl typc may increase the acreage ciC pint pl;in- In the Georgi:~ Piecln~ont,bottom-land hardwood for- t;ilions. 'I'hc pIa11t:itions Inily yield 11igli :in?ouiits of csts contiiined more browsc than lohlolly pine fcirtsts bro\vst. for the first fc,w yc;irs, but after tlie crowns i Moore and Manney 1962 1. In another Gcorgi;, study. close, little or no browse lvill be proilucetl for 15 tc) 25 hardwood types had significantly Inore brows(: t1i:in yclars i Str:~nskyaiid I1;ills 1967 I. Blair ( 196'7 I found sltisk and pine-hardwocid types 1 Moore et (~1. 1960 t. that lliiriliing increases production oE forage in planta- 111 South Carolln;~,Moore I 1967) found that bottom- tions. After planted trccs have reaclled snwtirnber sizc. land sitcs cont;iined many moro desirable browse species browsc is again pltkntif'ul I Sctir~stcr:inti 1l;ills 1963 I. than upland sites. Ife estimated carrying capacity at Retaining portions of tlie original mast-bc;iriiig htirdwood one decr to 13 acres in bottom-lands, as coml~arecito component would undoubtedly be bc~nc~i'ici:ilto (leer. 30 to 50 acres in It>blolly pine-hardwood, ;~nd78 acres in the longlcaf pine typc. Upland Hardwoods Even though the limher stand on it was dense, the Thc upland hardwooti type occupies 57 million .icrcs stream-bottom hardwood type was second only to open moitly along the northcrn bound,iry of the iouthrr~i cedar glades in tile protluctiorr of Ilc-rhage and browsc in forest belt. It is also found to some extent in the Caro- a study in the Arkansas Oznrlis (Segelquist anti Green linas and Florida. Principal tree species are oaks and 1968 1. hickories in association with southern pines, elm, gum, Unfortunately for deer, hottom-land hardwoods are maple, and redcedar. Product ion of cicer forage is usutilly being cleared rapidly for agricultural crops in some areas low, but in good mast ycars footl is abundant. ( Stransky and I-Ialls 1967 i. Through habitat manipula- 1)alko ( 1941 I reported that upland hardwood stands tion, the lobloly-sl~rt pine-11;irdwood type might in Missouri containcti at lcast 200 plant spc'cies of iri- be improved to offset the loss oi deer range in bottom- terest to decr: 89 in fall, 70 in winter, 78 in spring, nrrci lands. 115 in sumn1cr. Ileaviest browsing on woody stems took place in winter. I3c reported that the 11ost onlc- Loblolly-Shortleaf Pine-Hardwood blackjack oak association yiclded 140 pnuntls of forage This forest typc, which covers '79 million acrcs, is per ;icrc. ;,11d the black oak-lrickory ossociiitioti 110 dominated hy the two pine species; the proport,ion of pounds. hardwoods varios. Lay ( 1967 ) showed that, next to Ilunkeson ( 1955 noted that mushrooms in ul~laiid bottom-lands, this typc has the greatest variety of fruit- hardwoods of the Missouri Ozarks may m;lkc up us rnucli bearing plants. This variety is important to dcer, bc,cause as 25 percc:nt of tlie dcer tlict at times. He also found fruits and browsc are tll~lsavailable in different seasons tliiit deer browst:d heavily only when ;icorns were scarce. of the year ( I-Ialls at-id Alcariiz 1965; Lay 1967 1. ,%corns Scgelquist arid Greell ( I!)(it< I niadt: tlic same obsc:rvation are not as abunclarit ;IS in hottoni-lancis and arc 1~rob;ibly in Arkansas; forbs werc the major fooci itcins duri~igthe available for sllortcr periods t,hroughout the fall and growil-ig season. winter; mast failures ;Ire also more frequent. Elirenreicll and Murphy ( 1962 i rne;isurc,d Sorage yiclcis In north Georgia, Ripley a~ldMcClure i lllii3 I esti- of 155 pounds per acre in tile blnclijack o:ili-~>o~to;lli mated carrying capacity at onc deer to 39 ncrvs. Also association and 110 pounds in tile black oak-scarlet oak. in Georgia, Moorct c.t (11. I I!lfiO i noted that in r~irrying Associations 1 hat i~~cludetlcctiar liail much grass, ;I rhiir- capacity the type was bctwccrl the bottom-land h;rrd- acteristic also noted in tlie Arkansas 0z;irlcs by Scgcl- woocis and the sliish pine types. In Soul11 Carolina, esti- ciuist and (;rcc:rl ( 1968 I. r.n;ltod carrying capacity was one dcer to 30 to 50 acrcxs Springfictld 1'latc;iu forc~sts in Ar1tatis:is yiclticci 57 (Moore llI(i7). In North Carolinti, it was one decr to pounds of grass, 166 potintls of forbs, ancl 326 po~rtidsof 26 acres ( Moore and Strode 1966 1. browsc per acre. Eoston Mo~~ritninforests yieldcd 195 Ovcrstory density influences forngo yiolcl. For cx- jio~~ndsoS gr3sh 234 .j~cx~rldsof forbs, and 4-10 pounds ample, yic.lds from east Texas werc? as high ;is 1,600 of browse per acre. In ;in open nic~ntlow,grass 11roduc- pounds per acre on clearcut areas, 727 pounds undc:r tion was higher, 917 pounds 11or acre, hut forhs (99 shelterwood, 853 pounds under selection cutting, and ~~oundsi and brov>.s.sc I 84 pour~tls)were less than on the 426 pounds in uncut loblolly-shortleaf pine-hr~rdwood virooded sites ( fialls et nl. 1960). In iiortti <;eorgi;i. clcc.r l~i,owscyioltis \rlcxrcx only :3'3 ]i11~111clsper acre 111 iipI;iri~l Ii:i~~~l\vo~~~ls;inrI 42 p11~11cls ~ii~clc~r1)inc:s I l:i1,lc-y ;itiri Mc('1~ir.c: I!iii:i I. I!rowsc! yiclcfs ii,~>r(>;II.SI~ IOLV l7tj [)olin(I,sIII>I. ;IC,I,~,1 ill !hex O.<:~t.li,s1 S~*gi>l- (jiiis1 iiii(I C;~(Y,II19[i>l 1, ii~ilIiiglior ( I21 [IOII!I(IS I it1 tl;c, OIili~h(iiii~~Oii~i(~!-iit;~!i i S(~g(~Icjiiis1 ;iilcl 1~~~1111it1gloiiI$Hi21 I. Iri rc~c.c~nt1ythiiiiic9ti s1;ititls in 0~i;ieliit;ts?)i.c~\\-\c, yic31tls ~-r)s<.~IJ 171 i~oiiiitis 1x.r acr(,. 111 110tl1 tl~(: OL;irIi ;111i. Miss. (;;imc\ Longleaf-Slash Pine ;~iicj Yisli 27 i 7 I : 10-1 1. I3l;iir, It. .V. 15167. Deer Sorilge in :I lot~lollyj)itic> ~)l;i~it;ifion. ,J, \Vil(llifc~;\~l;iii;ig<~. :3 1 : 432-437, (yi~llitls,,J. 0. hest c;itli~.gr.;izii-ig iii southcri-i forcsts. I3rowso ;i~rclfruit I!)i; I. 'I'c.11 yc*;ir ;ic.oi~ntii;ist l>r.r~ri~ictioiislutly in .i~ic~ltls;ircl risiinlly low I (;i~cicirurn ;inti I'Lcxitl 1959 1, ;iiid 1,ouisi;itl;i. T,;i. Wiltl I,ife anti Fish. Cornn~.. tlci,r h;il>it;it ciu;i!ity is gcncrnily poor i Strarislcy and :3:3 pi>. Iialls 1967). 111 I~lc~rirl:~,II;irle)w ( 1951i i fourit1 oiily ;I sm;ill varic~ty r>;111<(!, iJ. 1). (if ~ootlyhlciuse p1,inti 'i~.iil.iblv 10 ticc.1 tluriiig tlic 1!)4I. 'I'lic irse at~tl;tv:riIability of lh' more com- ~~iiilcr111 li,it~tt,iti of loiiqle,if piirc, ttiriicy i,,ik, ,inti pine 1111111 wil~lcre1c~c.r hrowsc planls ill tllo Mis- il,iiu~oods1x1 the 1,onc.i C~~~ist'iII'l,ilil If,i~low,tntl Jon?< souri Oznrks. Sixth N. Alncr. Wildlife ( I0fi5 I r,iteti the c'ir 'ying c'ilj'ic3lty of iIcitwoods low Coi~f.'I'r;ilis. 1941 : 155-160. onc deer to 82 acrci 'I'hcy rated the c ~irI ying c~p~rclty I)unltcson, R. I,. of loiigle,ii plnc.-ocik ul)lands high, olic. tlccr to 34 acrc5. 11355. Ilcer range nl~praisal for the Missouri ~)rol~ablyhec,iuic of the I'rrge proportion of hardwoods Ozarks. .I. Wiltllifc ililanage. 19: 358-364. The abund,incc l)ei I 1,.1y 1965) Soc. Amcr. ITorest. Proc. 1958: 139-143. 'I'hc low protluctsvity of lonqlc.,tf ljtnc' I,rndi for dcer Ilalls, I.,. I<., atid Alcaniz, R. ti further tll~~str,rtcdby the ~ivcr,lgc>kill of one deer lo 5 2,000 or more 'icrri in Iol~qle,if forests ~n I,oursian,i Scasonal twig growth of southern browse plitlits. USDA Forest Scrv. Res. Note SO- r H,ty good 19(ib ) Simi1,ti cl,it'~ 11,i.v~ becn reportetl ?lie- 23, 5 P]). South. Forest Exp. Sta., New .ibhcrc in thc South (Str,inilcy ai~ciI1,ills 1!168i Orlca~is,1~. CONCLUSIONS AND OUTLOOK 'fulls, I-,. K., and Alcaniz, 12. IfBrowse plants yield bcst in forcst openings. F:vc,ti though tli~c1:il;i 011 Cooti procluclic~n aiicl tlccr J. Vi'ildlilc Mantigc. 32 : 185-186. ~~op~illatio~isarcx fr;ig~ncnt;iry, il appears that the major forc~stsdiSfcr ~videlyin carrying capacity. 13ottom-land I-l;~lls,1,. K., Rclatl, 12. A,,and Crawford, I-I. S., Jr. i~arti~voot1sare thc i.)clst hnhit~ilsi'or w1iitc.-t~iiicd dcctr, I NiO. Forage :111d ground-cover conditions in un- fi~llo\vcciin i1rtlcr hy the Iohlc~llypitic-liai-tlwood, upl;ind maiingcd Ozark forests. USL)A Forcst Serv. Ii;irtl\vootl. ;inti longleaf-sl;isi? pint, tyl)c>s. L'rilnc hottom- South. Fore,st Esij. Stn. South. Forest Rcs. l;it-iti acrcxngc?. lio~vc~vi~r,is (I\vi~icilillg by tll~(ii~y. In 1 : 1-6. iohlolly t~ine-liarti~voodst;lnds, it~tonsivetinibcr mariagc- Halls, I,.K.. and Stransky, J.J. lncllt nl;iy ridilc\c: bro\vsc iirltl m;ist sul~l~lics.Possible 1968. (:anlo food pI:intil-igs in southcrn forcsts. sc~li~tiotlslo tlrc~se prof)lcnis arcx rc.scrviiig ccr1;riti areas Thirty-'l'hird ?I. i2rner. Wiltllifc ant1 Natur. for gatne, intc~usiiying1i;iRitat til:iicigo~ilc~tton otl~c~rs Resourccls Conf. Trans. 1968: 21 7-222. by. s~s~jplollc~ritini:.- n;itrir;rl for;igc xritli footi plots, or ciicour;iging the growth of nalive hrowsc pl;tiits ill per- Fk~rlow,R. F. mnr~cntoljeniiigs i Iialis ;illci Str:insky 1!1613; 1I:rils anti 1956. Prc1itiiin;iry report oil a deer browscl census Alcaniz 1968 1. based on 100', clip~~ingmethod. Southeast. Assoc. Game and Fish Comm. 9th Annu. the Georgia forest survey. Sout11r:ist. Assoc. Conf. Proc. 1955: 134-156. Galne anti Fish Cornm. 14th Annu. Clolii. PrOc. 1!)6O : 98-104. Harlow, R. F., and Jones, F. I<., Jr. 1965. The white-tailed deer in Florida. Fla. Moorc. W. II., and Strode, I). 11. Game and Fresh Water E'ish Comm. Tccli. 196fi. Deor br,o\vse resources of the U~vharric Bull. $4, 240 pp. Xation;ii F'orcsst. USlIh Forcst Scrv. 1Ze- source Bull. SE-4, 20 pp. Soi~tlieust.Forest Harlow. R. F., and Tysoti, F:. L. Exp. Sta.. Aslreviiie, N. C. 195!4. R preliminary report on the effect of mast abundalice on tlie \veiglit and rei>roduction Riplev T H ,111ci McCllire .J P of deer in ceritral F1orid;r. Southeast. ilssoc. 196:3 Deer bro~viereiources ol i~ortli (;cwrqj,i Came ;irid Fish Comm. 13111 Annu. Conl. USDA Forc\t Serv 12csouicc I!ull SI+:-2 20 Proc. 1$15!): 62-69. pp. Southeast. Forest Exp. Sla.. Xsi~c~ville. N. C. Haygood. J. I,. Sciiiistcr, J. I,.. and 1I;ills. I,. K. 1966. Where are the deer in I,oiiisiana? La. Con- 1!163. Timbcr overstory dc1erl11iric.s deer forage serv. 18i 9&10 i : 8-10. in shorllc;if-loblolly pine-harti~voodforests. Korschge~~,L. J. Soc. Amcr. Forest. Proc. 1962 : 165-1 67. 1962. Foocis of Missouri deer, with some manage- Scgclqi~ist,C. A,. ;ind Green. W. E. ment in?plicntions. J. Wildlifo Maiiage. 26 : 1968. Deer food yields in four Oxark forest tyl~cs. 164-172. J. Wildlife Manage 32: 340-337. Lay, D. W. Scgelyuist. C. A,, and Peiinington, R. E. 1965. Fruit ~~tilizatioriby deer in southern forests. 1968. Decxr browse in the Ouachilu Forclst in J. Wildlife Manage. 29: 370-375. Okl~ilioma. J. Wildlife M~inagc?.32 : 623- Lay, 11. W. 626. 1967. The importance of variety to southern deer. Stcgemn~i,L. C. Southeast. Assoc. C:;imc and Fish Comm. 1937. A foorl study of the white-tnilcd doer. 18th Annu. Conl'. Proc. 1964: 57-62. Second N. Amcr. Wildlife ConS. Trans. Moore, W. H. 1937: 438-445. 1967. Dccr browse resources of thc Atomic Strnr~sky.J. J.. and IIalls, L. K. Energy Conilnissio~i'sSavannah River Pro- 1!3137. Timber and game food relatiolis i~rpine- ject Area. USDA Forcst Serv. Rc:source liard~vood SOI-ests 01' tile soull~eri~United Bull. SF-A, 28 pp. Southeast. Forest Exp. States. Fourteenth Congr. I'roc., Ilrt. Union Sta., Ashcvillc, N. C. Forcst. Res. Organ. Vol. 7: 208-217. Moore, W. El., and Manney, R. B. St~,anslry.J. J., and Halls, L. K. 1962. Deer browse plants in loblolly ;i~i(iwater 1968. Woodialid m;iri~igt~menttrends tli~ct ;iffect oak-gum types of Piedmont C;eorgia. USUh game in Coastal Plain i'orcst types. South- Forcst Serv. Soullieast. Forcst Rxp. Sla. tr:ist. 12ssoc. C;;ii~ie ;ilici Fisli Conim. Zlst Res. Notes 172, 2 pp. Annu. Coni'. I'roc. 1967: 103-1 08. Moore. W. kI., Ripley, T. H., arlci Clutter, J. I,. Wl~c~elcr,I' R. 1960. Trials to dctornlit1e rcl~itive (leer r;illgo 19(ili T~rnberregion5 01 thr. South Folc,\t E"ir-~n- carrying ccip;icity v~iluesin cor1nt:ctiolr Lvitli vr2517) 18-19 Agricultural Clearings as Sources of Supplemental Food and Habitat Diversity for White-Tailed Deer' Joseph S. Larson Massachusetts Cooperative Wildlife Research Unit Bureau of Sport Fisheries and Wildlife U. S. Department of the Interlor Amherst, Massachiisetts

Forage cleariilgs have been ~iseclin (leer arld tur- mowing once every 2 years to complcte rer?ewal 1p1ov1- lccy mcrnagc.~nent since 1935 and by I96:i over 30,000 ing, fertilizing, seeding) on an annu:11 basis. acres in 22 Slnles werr: dec?oterl to tlzis prclctice. Many early programs called for inst:illation on n grid The cstiinc~tcdreplacenzent i:c~lncnt or pattern reported cti117s of the prtlctice, IIIL~(In (:cologic(11 fact the topography would permit. base conzrnens~rratewith costs cannot be esfcthlisl~ed The cost of creating and maintaining clearings varies to sltpport this progrcrnz. More resecrrc.1~on tl~rc~nti- depending on topography and cost accounting practices. tnti7:e cincl qz~alitativectspects oj n(tf urcrl (leer fi~oiis, Slate wildlife agencies tend to utilize appropriated funds pop~tlntionestimation, and dcer cind I!z~nl,rr l)ehai>ior ;IS lump sums for man;igcnient of given areas, mnking is needed to slzow wlzetller such costly and int(~nsir:c cost accounting for specific practices very difficult. The n~r~nagernentprogrnnzs c~redesirable. Rcd~ic,tionin USDA Forest Scrvicc requires that depreciation of equip- c;cpenditz~resfor pasture-type clecrrings and ,nore ment, road time, mileage. aciministrative overhead and emplzasis 01% de7;elopiny better methods to c~sscss related costs be co~nputedand assigned to each project. the effects of habitat clzc~ngearc rc~co~nnlended. Their quoted costs usually exceed those provided by DEFINITIONS Statc.5 Installation costs run from a low of $15 per acre to a h~ghof $1,000 per acre The lower f~gurerepresents Agr~cultural food clearings are opcn~ngs In forest the mlnimum cost to rejuvenate an old f~eld51te and the hnbltat, e~thernatural or man-made, which produce a lirghcr amount, the cost to create a clearing In rugged natural or planted forage crop requiring some periodic wooded mounta~titerraln U.;ually costs In the Coastal agricultural management. Sprout clearings, seecicd roads, I'laln origin ol clearings lies in ;in attempt to provide 1 10 to 60 acres. A seccthcd is prepared, licne and fertil- diversity and food in the Sorest c~nvironmcnt. Wildlires izer added and seeds sown. About 75 :;pecics arid varie- originally ~~rovidvcln disruptive c~cologywhich favored tits of plti~itshave bi:r~r tried hut cslovcrs arid grasses deer. Fire protection :iritl the succession of old I'armlands are most 1)ol)1ilar. M;~intc.iia~iccpr~icliccs vxry arict mow- to 1x)le-slagc' forest 11;ts crc;ltcd habitat lacking diversity. ing is most conlmon. Intensity of n~~iirrtc~ianrc!rulls fronr 'rlic first cle;rri~lgswere hantl-cut openings wliicli f;ivored --~ . -. - ~ . ' Bascd l;~ryclyon c~:irlic~t.studic9s by t11~aritlior (1,ai-son l!)(jlia, sprout gro~viliant3 relc.:ised grountl-level vegetation. Tlie l96ib). a(1vetit ot' 110we1-cc1uipnient soon mticie it possible for U S Birrc~;rii of Sport Fisirt~rirsand Wildlii'c, Blnssac1~tisctt.s m;in;rgc~rsto cl~;itcpasture-lilzc openings which would i)ivision of Fisl~i~rii's:irrd Garnc, WiltiliS(. :\l;in:igci~i(~ntInsti- tute ;irrd University of hlnss;~elrtisctls(Coll~gc of Agriciiltirrc,, resist succc>ssion ;~lrdrc~t;iiri ~~redomintintlyherbaceous Expcritilcllt Station, and Extension Scrvicc) cooi~ci';riii~y. vcxget;~tiorifor ;I 1ongc.r tiine. Today wildlife managers believe that the major role We have no sound evidence of the effect of clear~ngs played by clearings is to provide supplemental food. A on wildlife production, movements, and harvest To my secondary role is lo increase "edge effect." Influence 011 knox~lcdgeno "befo~e and ,iftcin data ,ire avall,ible tc game harvest, public relatio~is,and ii1iinl;il distributio~i \uppoit lhc cornnloll Li,iu~nptiontkiat L~CJIII~Y\in~ieabt are other roles less frequently cited. Turkey and deer harvest and hunter success We know hunters use the are the species for which clearings are created. Pasture- access roads and clearings, but we do not know what like clearings are more commol-~than the infrequently this means in terms of man-animal contacts or how we mowed or brushed type. might manipulate hunter distribution in relation to harvest. PROS AND CONS W~thoutthis knowledge we cannot intelligently dis- Posifi7;e aspects.-Diversity of habitat is necessary for cuss the relative values of forage species we might plant, successful managetncnt of deer populations and where the proper fertilizer or lime applications, or optimum this is lacking a system of clearings can provide openings, sizes, shapes and distribution of clearings. Yet these are "edge," and diversity of food species. The most striking topics which today consume time and money in many evidence for this that I have observed is in the large southeastern and southern deer management programs. pine plantations of the Piedmont and Coastal Plain of The relatively new questions about possible parasite South Carolina. Here the clearing program probably is and disease exchange on clear~ngsare not going to be the key to maintaining turkey populations. Although resolved until we know how clearings, or any other we arc discussing deer habitat the example is still ap- hab~tatmanipulation for that matter, aifect distribution propriate. Where man creates an artificial monoculture and feeding patterns of deer. it is axiomatic that remedial measures to benefit wildlife must be artificial also. AVAILABLE AND NEEDED KNOWLEDGE Lush green clearings with abundant evidence of deer Current knowledge -The literature contains abundant use (droppings, tracks, and grazing) are tang~bleevidence references to the need for hab~tatdivers~ty and to tech- to the public that "somethlng is being done" for w~ldllfe niques for creating clear~ngs Leopold's (1933)edge effect on a management area Openings and clear vrstas In an concept, Stoddard's (1936) turkey observat~ons,Blakey's otherwise closed forest present good opportunities to (1937) Ozark turkey range recommendations, Mosby and observe deer frorn concealed vantage points Handley's ( 1943 turkey management monograph arid Where otherwise lacklng, clear~ngsand thclr attendant Wheeler's 11948) turkey work in Alabama have been access roads provide avenues of hunter u9e Row hunting part~cularly influentla1 in establishing forage clearing may be nearly excluded where clear openings frequented programs first for turkey and then for deer On the by deer are absent wholc these stud~esare based on broad hab~tatevalua- Negative aspects.-Installation and maintcnalice of tion In the f~eldand support the need for d~versityin clcarings are very expensive practices, especially when vegetative types Some encourage estabhshment of clear- carried to the point of operating a pasture improvement ings to achieve this, but none ~ncludedetailed studies on program. Maintenance costs can easily exceed installa- the effects whlch one might expect for such a costly tion costs by several-fold. Habitat diversity can be pro- practice. vided through a well planned revenue producing forest More recently a few studies have approached this goal. managemcnt program. Cuttings, log landings, skidroads, Lay (1957) and Blair and Halls (1968) found the southern millsites and the like can provide openings, edge and forests low in protein and phosphorus requirements for a rliversity of nat~zralfoods. Seeding for erosion control livestock and suggested that supplemental pasture pro- car1 be designed to add forage species otherwise absent. viding these elements for deer might be important. By These openings are less permanent than managed pas- and large these studies considered woody stems, shrubs, tures, but they are byproducts of a profitable program and vines. We have comparatively little knowledge and under forest liarvest rotation new openings can be about native herbs and forbs. McGinnes and Ripley regularly added with~nthe confines oi the management 11963)reported that inclusion of clearings in the long area ttrm managemcnt research program on Broad Run, Jef- With few exceptions an extensive clearing program ferson National Forest, Virginia, did not increase the cannot be j~lstifiedon an ecological fact base. If cleariilgs dccrherd. Englisll and Bramble (1919) showed that deer are actively supplementing the food base of any manage- use of clearings can be related to soil type and its subse- rnent area we cannot prove it becaiisc we do not know quent influence oil plant nutrient content. Added lime what is bring s~rpple?~~c:rrfrd.Quantitative measurements and superphosphate on deficient soils will attract deer. of forested deer raiigtl are largely restricted to woody Lewis (19671. in central Tennessee, determined by sta- browse. Qualitative measurements ;ire almost unknown. tistical analysis that turkeys were sighted significantly Con~p~trisoni,Ire frequently madc betweeti planted lor- clo5tr to clearings than would be expected ~f thev were age and woody stems, shrubs, and ~~inrs.Comparisons distributed system;rticaily over the forest. The average with native volunteer hrrbs and forbs, of the typc e11- nun~bcrof turkeys per observation was significantly couraged by minimum mowing ;uid fertilizer on South higher for ficlds 10 to 20 acres in size. What implication C;irolitia clearings, are lacking. Unless the surrounding this may have for doer is not clear, but certainly this fort.st range condition is known, quantit;~tivelyand qual- shows that statistical analyses are not too sophisticated itatively for all major food sources, \ye have no may of for evaluation of this mancigcment practice. knowing what suy~pleinenta1role clearings play or ought Research on "cdge effect" since Leopold 119331pre- to play. sentcd the concept has been sparse. Most work concerns sottghirtls, where ljositivc~cc~rreltitioi~s 1i:rvc~ ljc~liest~ib- 2. Oiir tot;il c~tiviroiinicnt is ch;ii~ginganti wildlife lislied hcl\vcoli "ctige" and riest :iI:iiiitl;ilic*c, ;ind quail mni-i;rgc~iiic~-ildccisic~iis once sriitablc for ;ipplica- \vlicrc call cou~ltsn~id f;ill csovc~ys;ire rc1:itcrl to cert;ii~ ti011 to wlit~lcS1;itc.s or I;irgc, portions of States ty])c~oS"c.tlge." 13:ii.ic.k 119501 found t11;it ;ill "edge" tnay he in;itic~c/~i:iteFor smaller arcxas becatisc we tyl)c>sarr-c.1101 ccjuiiily v;il~~;ii)lefor tieclr iit t11e Arjiro~t- ;ir~u~iable to nec~ir;itely nieasure deer numbcrs ti:i~lis.>it~tl C'rciss I l9(i:31 Co~ititidc,cxr Iraclis rii tl~i,Atliron- ;ilid ~)~~~jul:itic,lrcli;i~~ges utitlc~r eastcrii lorcst con- clacks ;ivoidcct clcariiigs it1 tile \viiiti,r i~tilessthey 11;iri dilio~~s.011ly large cIi;i~igcs iit 1joljul;rtioli num- soiitlicrly cx1)c]stirc.s. Currcnlly tl-,cxrc~is littic evitioicc to sul,lji~rttile cli~im1Ii;il "cdgo" crc:itetl hy c~lc~iirit-igspro- f;rctc,rs. c:it~ hc tictr.clcd using present tccliniqucs vitlrs ally 1)cisilivc~herrefit ti, clc~~r. to clstimate ariin?al nu!?ibers. Clearit-~gs it? most ;2l:ii1y workers havc es;imitic~rlprcicl~~ctio~r tiiitl i~tiliz;~- c;ises tire rc~lntivc~lysni;ill oi\~iroriment:~lchangcs. :iiiti if tlicir c~ffcc~tsarc1 ti, bo tlctectcd, tltcn very tion of food s~jc~cicsljlnlitccl it-, elcirriligs. 'I'liis tylx of .ivork h:is bccn rcvicwcd c~lsc~\r.hererI,;rrsoi~ I!l(j7bi. rlr;iclis, 1 dropioings, ;illti balcs of fcirngc, arc not c~ild-prod- :ircl ~ieotied.1 stiggcst that iiigh do1l:ir costs trsso- ciatetl with for:~gc.c1c:iring liian;igernciit slioultl intiicc>i u Iiich lt,i\c. yet to IIP rc~1,itc~tl10 ,triini,rI rttirnhc~rs hc liiild not be, meiisurccl oiily ;igc'~nc.rit pr;icticc~s, si~cll;is i'or:ige elclarii-igs, arc I1ecc.s- it1 tcxrli~sof iinimtil i~u~nbcrs.Mtiriipulati~ig ~cnit~inl stiry. 111iic11 1jc.tti.r iiil'c~rmtitir~~iwill be. iicc~icti iii four tlistril~utiottanti m;ili-arlirnal co~ltnctsc;iri he valid objecti\rc:s of ;i ~nti1i:igcm~1itpri,gr;ini. Telctiietry ;~ljl)(';u.sto offer th:. grcx;ilcst potc,nti:il Sor evalu- ati~igtiocr bch:ivii~rin rclaiio~tto habitat corn- cic~c~rfootirosoiirc.cs. Rc,sc;ircl~iii t lic past Ii;is con- l~(nieiits.Such sliitlies sl~oultipcrn-tit cv:lli~ation cc.iltratcti (it-, \vootiy bro\vsc~bul icx;iti tic~,rstiltiics of "ctlge cllect." :ii-iti tiefiiic optimum size, sliape, in Pclitisylv:iliia i Watts 1964; Ilcaiy l!)(i7! intli- dis1ributio1-i cilirl c.ffcct of clc;iril~gs oil tfeer be- c.;rtc> that tlot>r fc~~iIic~;ivily oli I-ic~rl~iiccoussljoeic~s. 1i:ivior. M;irchi~iton :ind Jcter (l!I(i7! found that S~~gclc~uist;111tl (;rc.c.tl I 1968I li;iv(' Foutltf t hat ;I corn1iio11 ritovcxmc111 p:itte~,iiof r;idio-t;iggcd tfcer in 1Zl;ibama ;iltd I'loritl;~involvett fccdi~lgin open ;rrc7;ls or fi~rtige~cletirings :it r-iight iiud a return to ~oocictltIrc;is :it tl;iwrr, but 3 liliiitcd nuinbcr of ;ilii~n:ils 1vivct.e tr:icecl. More ititc~tsivestudies tilrtcleci

.> L s~iulIrc~:ixt,A!Jiicii ir~ii)ro\.t~tii~,~tlc~rstti~~t-li~~g of ho\v 1. 'I'lic stutly oi' Itiit~t~rbelitivior, wit11 sljecific refer- eicxcr cxtract Eoc~ti ~ic~~isfroin ii;ititral fi~rc>strcit~ge circc to cltarii~gs,nl?],e;irs lo be vvcll s~iitedto the \vil1 tlc~vc~leip;I ri~C~rc~iic,c1)oinl for c3v:iluntii?g tlic tccl~liiqucsc~mploycti in recrealion rcse~irch. Tile triic, roI(1 of cleiiring.~;is .su~)plernc~~it~ilf'oi~dsources. rc~latioiiof bow hui~tiugto clearings iippcars self- evident. James cl (11 ( 1964) found that all deer LITERATURE CITED kills by all metliods in the North Carolina Pied- Bailey, R. W., Uhiig, 11. G., I3reiding, G., and Shrullk, mont were within 1,800 feet of a road or trail, and in the mountains nearly all were killed within W. F. 2,400 feet of similar access ways. Roads and trails 1951. Wild turkey lnan;~gemerit in West Virgirlia. appeared to serve equally in distributing hunters, W. Va. Conserv. Comni. Rull. 2, 50 1113. and they speculated tkat the difference in use of Barick, F. B. "off-trail" areas might be due to differences in 1950. The edge effect of lesscr vcyetatioii of certain the hunting habits of urban and rural sportsmen. Adirondack forest types will? particular refer- If this is the case, certainly there are many other ence to deer and grouse. N. Y. State Coll. hunter attributes and preferences of which we Forest. Roosevclt Wildlife 13~11.51, Itti pp. know little and which may have importance in Blair, R. M., and Halls, L. K. habitat management programs. 1968. Growth ;ind forage yucility of Sour souther11 I)t~o\vsc ipecic,s. Soi1t1ic~;ist..2ssoc.. (;;imc ;ind CONCLUDING OBSERVATIONS E'isii Con?ii?. Zlsl Xiiciu. C'oiiC, l'roc. llI(i5: 37-(32.

If the expenditure of nearly $8 million in creation and Blakey, H. L. maintenance of clearings has been an entirely satisfac- 1937. ?'he wild turliey on the Missouri Ozark range. tory investment then this discussion is academic. I sus- U. S, Biol. Surv. Lc;ifl. BS-77, 32 py. pect that this is not the case. Wildlife rnnnagers who Cross, C. W. contributed their data and the bcnefit of their experience 1963. Wildlife use of small forest clcarings in the to my studies were not in cigreemcnt regarding the roles Adirondacks. M.S. Thesis. State Uliiv. Coll. clearings played in their management programs. As Forest. at Syracuse Univ. N. Y. 122 pp. early as 1936 Gabrielson 11936)doubted the economics English, P. F., and Bua~nble,W. C. of clearings. Essentially the same questions were raised 1949. Kill 'em?----starve 'em?-feed 'em? Deer eat by Bailey et al. ( 1951) in West Virginia, Krefting ( 1962 best forage they can find. Science for the in the Lakes States, the Northeastern Forest Wildlife In farmer, pp. 7-8. Penn. Agr. Exp. Sta. Bull. 502, Research Committee and myself today. Suppl. 3. Wildlife managers at the 1962 Northeast Wildlife Con- Gabrielson, I. N. ference were reminded by Longwood ( 1962 ) that, "Con- 1936. The correlation of forestry and wildlife man- fronted with a task too big for our manpower and facili- agement. J. Forest. 34: 98-103. ties we have a tendency to dive into the physical part of the work without adequate advance planning. It is Ilalls, L. K., and Stransky, J. J. difficult for us to sit back and go through the processes 1968. Game food plantings in southern forests. Thirtg- of analysis and planning when so much needs to be done. Third N. Amer. Wildlife and Natur. Resources It is easier to get at the job and leave the detailed plan- Conf. Trans. 1968: 217-222. ning until later. This is where expediency overcotnes IIcaly, W. M. our better judgment." 1967. Forage preferences of captive deer wliilc free ranging in the Allegheny National Forest. M.S. With due regard for the need for "action programs" Thesis. Penn. State Univ. University Park. 93 in the early days of wildlife management, I suggest that PP the extensive use of costly clearings is the type of prac- tice to which Longwood refers. Major emphasis on a James, G. A,, Johnson, F. M., and Barick, F. B. single management practice in hopes of effecting n posi- 1964. Relations between hunter access arid deer kill tive response by deer ignores the adlliixture of variables in North Carolina. Twenty-Ninth N. Amer. which make up deer habitat and is a high risk venture. Wildlife and Natur. Resources Conf. Trans. 1964: 454-463. I recommend that we draw the line on adding more Krefting, Id. W. of the intensively managed pasture-type clearings and 1962. Use of silvicultural tcchniqucs for improving reduce the maintenance costs to a rninimum on existing deer habitat in the lake states. J. Forest. 60: clearings. Where diversity in habitat is laclting, coordi- 40-42. nation with forest management will apply costs to several goals and keep single-purpose wildlife expenditures to L,arson, J. S. a minimum. 1967a. Forests, wilcilife ant1 habitat manage~nent--a critical exan~inationof practice and need. USDA More emphasis should be directed to bettering our Forest Serv. Res. Pap. SE-30, 28 pp. Southeast. ability to understand and predict the effects of habitat Forest Exp. Sta., Asheville, N. C. change oli animals and man. The East has experienced one cycle of habitat destruction anti recovery. In all Larson, J. S. likclihooti we are approaching a period of rene\vetl Inan- 19671). Problems in n~easuringfood production on Sorest niade thrcat to wildlifc habitat as hi~rnan populations wildlife cleclrings. Southeast. Assoc. Game and L~IIC~demands on resources grow. Devclopnient of el'fec- Fish Comm. 20th Annu. Conf. Proc. 1966: 33-37. live neth hods for evaluating habitat cliange will find Lay, D. W. broad applications in environnit~ntaln1;tnagemc:nt. 1957. Some niltritional problems of deer in the sonth- ern pine type. So~itiieast.Assoc. Game rind Fish Mosby, H. S., and Handlcy, C. 0. Comtn. 1Otl1 Aunu. Conf. Proc. 1956: 5:1-59. 1943. 'rile wild turkey in Virginia: its status. life history and management. 281 pp. Richmond: Va. I,eopold, A. C'omm. Game and Inlar~dFish. 19:33. Game managcmerit. 481 1311 N.Y.: Charles Scribncr's Sons. Segelyuist, C. A,, ;ind Gre~li,W. E. ll)(i8. Deer food yields in four Ozark forest types. J. 1,ewis. J. C. Wildlifc Manage. 32: 330-337. 1967. Populations of wild turiicys in relation to fields. Stiteler, W. M., ,Jr.. ai~dShaw, S. P. Southeast. Assoc. Game ant1 Fish (lon~m.18th l!i(jtj. Use of \voocIy browse by white-tail decr in Annu. (lolif. Proc. 1Yfi4: 49-56. heavily ftrrested areas of northeastern Uriiterl St;rtes. 'I'hirty-First N. Amer. Wildlife and Longwooti, 17. R. 1962. The selection, design and ~inalysisof wildlife- Yntur. Resources Cotif. Trans. 1966: 205-2 12. habitat stiidics. Pap. ~.c,:rd ;it Northc~:isl Fish and StotJci;~rti,I-I, I,. LVil~llii'e(.'o~il' I1 11ij. ll)::(i. >la11agc1netit111 wilt1 lui.Ii

Jack A. Crockford Gcorgia Game and Fish Comm,si~on Atlanta, Georgia

A cli>clrly dcjinetl philoso~~h~joj deer11i:rd ~~lnn- have i~e\~crhc~lrl ill :igrct~rircnt on what \ve ;,re tryink

(~tlcnz(:t~flzas nppccrc~ltl!l. . liol g(:1 c~t:ol~:etI(1s rrjlccfctl to do and how. b,tj tlt~i:nriety of ILILTI~~T!~~~gi~l(~tic~ns. I)ii(: lo I~c As ;in avid dcer liu~iter,it is alu.:ry-s iny 11opc. t1r;it ii~c~hility to clppl?~II!IIIL(LCJ('~IICIZ~ tecJ?niqi~~s,(11 lzrr thcrt, will he a place \viictre I can c.r>joy what to rne ii t Jlt~izhiii~iiitg. on ~(IT~Cper(.i~ttt(~gos of (leer protliicir~g q~~alitytl:>cr hunting. We 11:ive aluirys tricvi to be surc la~ld,it is s~~ggcstedlltc~l ht~~tiiilg is tltt liz(~jori~tj1i~- we have suck opportut~iiiesill (;co~,gi;~,;i!ici have bccn cnce pr~sentlyc~~~ctilr~ble lo rr~c~nrcgc. flrc: lz(~~t1.It is forturr;iic so far in being pcrmitteti this iusury, if it is f~~rll~rrsirggrstcd that i~;rlei~sit~~?c:o?-l; oit lzc~ri:csling ii luxury. Quality seems iml~ort;i~itto mc. but irftcr mc1l~od.sby .u;ilcllije 1uor1ic:r.s is ?L(~CCSS(LYIJ fo (~fji- n1:iny discussions on tilis suhjcct I have found vcry cicnily ILse this tool. il plcci is rt~clrlefor 1rr~clcr.ship c111ickly nr~tloften vcry bli:ntly tirtit wliat is acccpt;lble by the wildlif~projession irl 111c c~doplionof ol~jcc- quality to me may be quite diSfcroilt from what is con- tivcs, fi~rttterst~rdlg of Ftcrri;t:.sfi~~ginell~orls lo refine sidered acceptable to other inrlivid~~als.or other agct~cics. rcg~~lalorilprocrsst?s. c~nrllo ~~~ninlnii?(1 high qltnlily I'm not evt.11 sure wc? put the same ingretiicnts illto thc of the hunting cnperiencc ilz rc>giclalion matters. matter for consideration, but bclieve it is irnl~ortantto When I accepted the inv~tatlonto dl~ussthe philos- establish some stantlards. I have serveti for some years ophy of deerherd management ~evcralmonths ago, the on the Deer Narvest Subcommittee. At one of our assignment seemed like a f'tlrly s~mplcone I-lowever, mectings we decidod to define "quality," as rcl:ited to it soon became more complex and ~t\ec1ms to me now that deer liunting; and we did, over anil over again all day the most I can do is to discuss two basic problems. I long and into the night. We defined it too, but I dorlbt havc come to bclicve that the philosoplly of deer man- if we evcr agreed privately on the accepted definition, agement is reflected in the regulations governing the but rather gave in from sheer exhaustion. At least I did. harvest. If this is tr:le, a review of tlie variety of The definition is as fol1ou.s: "I-Iunti~~gfor a maximun~ harvesting regulations indicates that a firm philosophy period of time with a minimum of lrunter contact and has not yet bee11 developed. This is the first problem. controls and a rcason;thlc opportunity to see rlocr or The second is, that given ;i philosophy or clear set of fresh deer sign." objectives, it is doubtful if we arc sufficiently knowledge- In the southcast. managcme~~tareas have become an able to set reg~llationsto effectively reach thkr goals. accepted fact of life. They continue to persist, probably Put more simply, I don't bclieve we are in :igrecmont, aftcr their original purpose as restoration areas has been on what we are trying to produce or how. If stating s:,rvc,d. Some might even be better off if abandoned. the problem is the first step in its soluiion, I hope then It seems to Ine that one of the things which permits that this discussion will bt: a contribution to this program. them to live on is tlris matter of quality hunting. trow- ever, hcre again, tlrese areas arc hunted in a variety of PHILOSOPHY OF DEER MANAGEMENT ways. It is the hope of m;iny of us that some guidelines For years we have met, with our department people, might (:merge if the expcriencc~sover the years on tiicsc and argued long, loud, and often bitterly, to sct regula- areas were ev;rluat,ed. 'I'he Forest Game Conrn~ittceat tions for the coming year. We are guidcd largely by orre time proposed n "circuit riding brain ~siclicr" to scientific biological data, but ;tlso by personal opiniofls, study this matter. One thing is certaiti, the appro;ichcs to some prejudice, often politics, anti always a fair amount lri~ntingtl~csc arc,as havit bcen v;irictl c,rlough to give some of tradition. The same traditions, whicii sacred :is t11i.y intcrcsting i~lform;ltion,and to indic;~tc there is some are, were at one point instrurnentai in clirninatil~gdeer cc:nSusion on how thry should be handled. 111 discnssions from all but a few deep swamps and private 11rescrvcs on this subject, I have hcartl that "1 day is enough for in Georgia. Somehow we come up with successive scts anyone to hunt deer; get as many hunters as you can of regulations which have permitted a steadily increasing in 1 day; kill as many deer as necessary, and you're decrherd and harvest. Thc9se regulations, at present, through wit11 it for pr no tiler ycar." Sornc. rc~cornnrc.nd con- vary from 3 days with control of hunter numbers in trolling hunter numbers by permits sold prior to tile somc areas, to 80 days with dogs in others; from bag stason. Still others suggest making the regulations com- limits of one buck to a buck and a doe or two bucks 1,licated cnough to limit hunter numbers and harvmt, season long; to some areas with either sex the last day anti many other appro:iches. In our State, we have prob- and other variatiotrs. Somehow, it has worked, but I lems lieclping some irrras open as long as wc, could, so personally don't feel that dcer man;igen~entis cjrlitc that far as dcer populations are concerned, simply frtrni the sophisticated, nor are we that artistic in setting c:ffcctive standpoint of administrative problems. Pertlaps then regulations. Looking back over 1n;lny sessions, it seems we arc "ovcradministratit~g'. tlic. hunts. On thc. otlrer to me the cause of much of the controversy is that we hand, and I know opinions vary otl this, we believe that in muriy situations cieer can easily be overlrarvcstecl. We of a farmer ;ind so far as I know, I was never able at have one very popular area in (.;ec:rgiti on ~vhichbuck any tinlc to motlify in any \vay his tlesirc for a "clean" hunting h;is l.jc~ctnne largclly a 11;iturc hike, we hclievc. farm. I tlii~ikwe are in tliis same positiol~today with from ovi.rkuiititig ljucks. I~tcitlentally,this is acconi- the various 1;iiiclowric~rships tncritio~ic,tl, without tho atl- l~lisltetiin 6 clays a year. Sonie yoars ago, :I notcci c1ec.r vantage of the close f;iii~ilytiw, ant1 I think llic resr~lts s11eci:iiist gave some s;ige ;itlviec lo gofrtl fricl-iti of niine. may ho soinecvh;it similar.

,3 IIis :itlvicc \v;is. "Son, yitilr only ljrohlcnt \villi (Icier is lhc varititicjli in I~iritlowiic~rshipsis on^ of our ])rob- killing cliongh of tiie~n." IVc. 1t;ivc.li't Sot~~itlthis to bc lc~iris,botlt fr-on? titi, s1:iiidpoint of ~iroclucticii~;il~tl h;il.\-cst. eiitirc,ly so in Gcorgi;~,and onc3 c;in im;igine tlici resillis I t lrink \vi. siio~~ldc,xpect morc intc~tisivc~jirt~rluctioli priic- if \ve shot "either sex" or? the ;irca mc~~itiorictlitt the 1ict.s ijli public Inlitis t1i;ili on intlustrial fol.cst iantis a~id spirit of that :itlviec~. still less on 1jriv;itc l~rirtls.It may also vc.1-y likely tlc~vc~lo~~ that ;is harvesting tecl~iiir(uesare rcfined the three classes Scver;il Scars ;igo 21 1iiilili:ig clul-, in tl~c'l) south (;ixi~rgii~ of o\&-r-r?ersiiips\vill rcyiiirc s?parnte reguliitiotis. 'I'lic rcportecl ii kill of seven bricks tlurir?g tlie IiO-tl;iy sc3ason on 100.000 ucrcs. Thc eluh mcrnlx,rs n.cxrc, cjiiite Ii~il~py. j~:rttcrn is 1)ro))abiy son~e~vhatsiniilar ovcr most of the their only eoiiccrii lxirig to have r:tioiigii tlcer to finti soutlic~ist.It would sccm clear to me tltc~ithat dccr are one Src~siitrack to have a chase with rltrgs. Such rc~siilts ;it prcXsciit,aiitl are tiestirictl to reriiuin lor the most part. a hyprotluct ctf the land. 'Sherefore tvc ~vill11ot 1t;ive can colnnionly he coritrastc~ti111 other areas oi' c~cju;ilsize which wc~ultlprotiiice an iinn~ialh;irvclst of 400 to 700 the c1i11x)rturiity lo al~plyIi~ibitat prticticc.~ prim;irily to hi~rlis. Ohviorlsly. t1ic.re is a witie gap in rcsults hcrc. produce dccr. controlietl :iltnost i3ntirely by hunting prcssurc, irt this Wc arc left thcll xfith liuntilig as the one major influ- case motiviited by tratiition. ence \viih \vkich to 1n;tnage rlrer. To t~cconi]~lisltthis. \vc meet 1 (lay a year ~~~itliour comti?ission. iitid 21s statctl. Most of 11s in the southcast hav(x for years been in- "fi~~itlcdby somc biological cl;ita, pctrsoilal opinions. volved in dcer ucxitoratioli progrunis, alitl think \\7ith I ~irejudicc,politics, ancl tl,atlition," we cxstablish rcgula- outstanding success. Most of the i~-ivtstig;itionsand re- tioits govc9rninji thc harvest. I thirik it coulri be said searck projects have been tione in line ~vithprotiuction; here that lve are attenipti~tgto produce deer sciontif- i. c., rcproclucti.r.e rates, ccirryilig capacity, i~ubitatn-ian- ically. and arc harvcstil-ig them "artfully." I am iiot agement, inventory systems, and others. This is all good sure what impact colnl~letcfreedom to rnatiipulate the and valuable i~iform;itionand the work must, course, of "l)~>ductiontlemclits" would have from year to year be continurd. However, now that we 1i;ive the dcer oli ricer in thv hag. if that is our goal, hut I think it is to manage, I think it is imporntivc that more thought clear tlt~it in a rc\gt~li~tionssessio~i in \vhic11 one of the and stridy be cxpcncled in the directioti of hiintiltg, wliich above lingretiicntc, becomes don~~nant,that a n?,ijor in- is no doubt the most effective ~~ic.thi;d\re ti:ivc wit11 which llue~icccan be 'ilfcctcd, often quite by acclclent One we can rapidly and c~ffectivt~lyapply cotttrols to the herd. of t11o5e ingredlcnt5 often does become ~Iomitlant,prob- would c:xpect somc critirism on this position, hut first I ,ibly hccauie we ,ire larqcly gutdrd by

Dean A. Murphy miss our^ Department of Coriservatio!i Columbia, miss our^

Deer h~cnfingregnlntions. brcg lirtzits (111~1ltirnting 50 States. The total harvest of white-tailed deer in the inetlrods in the sotttl?eccster?t United Stc~teswere southeastern Uriited States increased from 129,034 (leer .stndiecl thro~tgh(L SILT^?^^ of deer ir~rnfit~ginformcc- in 1955 to 559,645 in 1967. tion pcrmphlets. The following summary of methods, lirnits, and regu- Killing tleer by lise of clrtificiccl light nt r~ight, lations is b;isc~Itrri copies of 1968 deer hunting regula- sllooting tlcer in lakcs or S~TP(LI~LS(~nd ~~~rsniitg dper tions received from the 15 southeastern States il~volved. zuith nzotor ?:chicles were gc!rterally ~.rrohil)itc~cl.Legal There was a wide disparity ili completeness of the regu- rrtelhods included: still hl~nfingon Joo: and from (1 lations received. Some of the pampl-ilets were complete, stirnd, lzlcntirtg frolrz cle~xrtcdplntJoi-~ns, ilrit*ing deer clear. and concise. Some wtrc incomplete as to iriforma- by osgrrnizc.d gro?rps of hitntrrs, crncl c~crlltizg bzicks tiotr on legal wcaijor.is. methods or other factors. In by ~(xttlingof czntlers. c,tlicrs. complete information could be found only after Driving deer 111ifh dogs u~clsl?gcrl in 10 of the 15 searching through the entire it-iforrnation folder. The Stc~1e.s. greatest corrfusion occurred when attempts were made Rifles, sirotgztns, ntuzzle-loctders and longtjolus to i~rcorporateall hunting iand even fishing) regulations were legal teec1p3n.s in (111 15 Stc~lcs. Sez:en Stcites into one pamphlet. As an outsidcr seeking specific it~for- had sonze arras lin~itedto iise of shotg~insonly. Most rnation, I cerlajnly appreciated the pamphlets which of tl~eStates attempted to set restricfions on rifles. were devoted solely to regulations for hunting dcer or One State h(~dno reg~tlutionson firoar~ns. Shot- closely related species. guns loaded with bnclcskot were lrcgal in the 10 States which pc.rmitted driving deer with dogs. METHODS Daily bag limits were generally set at one dcer What should be the criteria for delermining legal per day but four Strrtes permitted cr nrltlliple daily methods for harvesting deer? The rnethods permitted hag. Secison li~r~itsvaried from one deer to 67 per should be humane, sporting, and el'fective, but not so season. effective as to give the hunter ali tinfail- advantage or Length of the firetrrnls deer sc.ci.sons vcl7-ied from endanger the resource. Killirrg of dcer by use of an 5 days to 120. Secrsons begun A~rgust15 cincl con- artificial light at night is an exaniple of a method which tinued througl~Janunly 19. fits the latter critcrioti. This mcthod is just too effec- The basic season wcts CL "bucks only" season with, tive, and is illegal itr all of the southeastern States. A iiny-deer being legctl in I-estricted arecls. Nine States swirnming deer is also very vulnerable and rnost of the pern~ittczdeitkc~r-sen. dcer I~clrvest in 1068. States in our region prohibit killing deer in any body Archery deer scc~sonsciverc~getl 58 drc?js. of water. Pursuing of dcer by any motor drivon convey- ance on 1;lnd or water is almost universally prohibited. Co~~trolof the harvest is the most rfjectir>e IrLan- cxgetrzent ,nethod c~vctilahle for white-lnilecl deer. Most of tile acceptable methods of hunting are legal Deer regnlations shonld be 7urilten lo p?.o~?idc'the it1 the southeastern United States. The most common ~rn(~.rinz~trni~1rtot~nt of rr~crc'c~tio~zccl oppor t nrtity for the method is probably stillhunting, by which I rncnn stnllr- ilrg deer or1 foot or waitirlg for them on starrds. It has grt!utest rtl~nrljerof h~tntersu~itho~tt er~d(r~zgerir~y th(! resource. The czirrerlt deer rey~tl(~tioitssecrrz to br been my csperietrcc~that pursuirig deer 011 Soot is the r~cconrplishingflris objeclilic.. hower,er, tl~~reis dan- least cfi'ective of a11 hunting methotis, c,sl,eci;rlly in coun- try where snow solclorn occurs. Waititig on a stanci is ger of overpop~ilationas 11 result of conlirtnc?tl hlcclcs- only Iz(LTI>~s~. usu;illy more prociiictivt:. ?'he deer hunting stand is gerier;illy more effective My subject, "IIunting mcthods, limits, and regula- if it is elevated. Platforms iri trees arc common, ;tntl in tions," actualIy collccrns control of the Iiarvc~st. It is Texas I ~~licicrsta~rtithat spccii11 Ilullting towers are built. cornmoil k~iowledgethat sil-icc the lsrgc native predators In Florida, dcler are Iiulltcti from towers t?roctcd on have been eliminrrted, harvest by man is the single tirost rnotor vehicles, 'I'he rc9striction that these towers could effective control on doer 1)o~rul;itiorisi~r the United States not be lriglier tharr 8 feet ilbovc the lo\vesl bottom surface ( Allen 1954. 13. 140 i. of siich vehic.les was tile otily l.cstrictioti which I fount1 Deer populatioiis wer-t! t~c'ariyeli~nir?;iletl from niost 011 tho LISP of c1ev;itc.d sta~itisCor l~ulitingdc'c'r. of the Ur~itedStates by unrc~gulaledhunting (luring the Ilrivitig of dctr by org;inizcti groups ol hunters is at1 pioneer era, with a low point in rnost St:rtes ill the early even motx, cffc:(:tivtx h~iritit~g111cllrod :itid appe;irs to be 1900's I Allen 1954, p. 135 I. Through eo~~trolof l~tirvt:st, accc:pt;rhlt> ti~rirugl~oiitiirc. rc,giort. ~isuallyaccompanied by restocking. ticcr have been rc:- Ilrivitig tlt,et will? (logs is ;I nlctlioti altnost urliquc stored to cvery Statc and deer art, tro~v11~1tit~d in ;ill 11) Ihc soiiIlic~;~sl~~~~nrc'gio11. l)ogs~~~:iyhe ust~i lc~gnlly in the pursuit of deer in 10 of the 15 States. The sportillg BAG LIMITS cti~icsof using dogs to ll~lntdeer appears questionable What is the purpose oL a llmit oil the daily bag a11d to me. However, it is contended by the advocates of "dogging" that this is the ollly method which can be possession of deer? The purpose should be to prevent overharvest, spread the harvest more evenly among the used to harvest deer from swamps of the southern coastal hunting population. a11d prevent "game-hogs" froni areas. The use of dogs may be justified in the swamps, slaughtering more game than they can utilize, thus wast- but what is the excuse lor use of dogs in upland pine and ing the resource. and hardwood forests'? Daily bag limits are generally set a1 one deer per day. Calling of deer by the "rattling" ol antlers is apparent- However, in Florida, two deer (bucks) per day may be ly confincd to thc State of Texas Other types of call5 taken. The regulation pamphlets from Georgia and Texas 'ire banned there and in most olher States Baiting and indicated multiple bag limits but did not indicate whether salting also are generally banned. these limits could be taken all on the same day. Season possession limits varied from one deer (buck) WEAPONS per season in six States to a limit of one deer ( buck) per Related to hunting methods is regulation of type of day in Alabama which had a 67-day season. Four States permitted two deer per season and hunters were per- weapons which may be used to harvest deer. Both rifles and shotguns were permitted in all 15 States in 1968. mitted five deer per season in parts of South Carolina Only Oklahoma had an area restricted to use of rifles. and Texas. Seven States had some areas limited to use of shotguns. FIREARMS SEASON HUNTING REGULATIONS In our region, the mail] reason fur limiting llunters to shotguns seemed to be related to the theory that shot- A well written deer hunting regulation should be con- guns are safer than rifles. I-Iowever, in Michigan, certaill cise, as simple as possible, but complete. It should pro- areas are restricted to use of shotguns as a means of vide a maximum of hunting opportunity with a minimum reducing the numbers of hunters. Tradition dictates that of restriction compatible with the best utilization of the deer should be killed w~tha rlfle, therefore many of resource. This objective apparently is very difficult to their decr hunters will not hunt with a shotgun and avoid obtain. Deer populations, range conditions, and patterns areas which are restricted to shotguns of hunter distribution are not constant even within the boundaries of a single State. These factors result in a What is considered an adequate deer rifle? The Sport- multiplicity of regulations designed to compensate for ing Arms and Manufacturing Institute recommends cart- the variations. In addition, sociopolitical factors also ridges in caliber of .23 or larger, developing a minimum complicate the setting of regulations in some areas. of 975 ft.-lbs. of energy at 100 yards. The Ammu~iitio~i Division of Canadian Industries Limited recommends that a decr rifle should have 1,500 ft.-lbs. of muzzle energy. Most States in the southeast have attempted to set restrictions which approximate the above recommenda- tions. Rifles of .22 caliber, rimfire ammunition. full- ,jacketed bullets and automatic weapons are generally prohibited. Only Mississippi has no restriction on tile type of firearms wh~chcan be used. Shotguns smaller than 20 gage and larger than 10 gage arc gerlerally prohibited. In the 10 Slates which permit the use of dogs, shotguns loaded wit11 buckshot are legal weapons. Shotgunners in other States are re- stricted to use of slugs. Muzzle-loading rifles ;ire legal in all 15 States. Most of the States set some minimum requirement as to caliber or grains of powder which are legal, but again the stand- ards vary greatly. Longbows are legal weapons In all States in the reglor1 Somc States set minimum standards for archery equip- ment but the standards vary wiclely. Crossbows are legal weapons in Kentucky, Missouri, Oklahoma, and Arkansas. Arkansas is tile only Stat,_. ~~hichhas a special season for crossbows. Legality of the crossbow was not clearly stated in regulations which 1 received from five States. Both rifles and shotguns are legal weapons. statewide, in Missouri. We find that 91 percent of our hunters use riflcs and 9 percent usc shotguns. A few hunters use pistols which are legal in .38 caliber or larger. Dec;iqnation of Ienqth of season and sex of deer which August through December. it would bc difficult to find rnav bc taken arc cornmoil to all rt'sulat~on\ I,c~lgfh oi trine fol a spccial archery seasori. 111 thosc States with season varied trcmendously in the southeastern region relatively long archery seasons, the archery and gun in 1968. The average number of days of firearms deer seasons usually overlapped. hunting per State was 47 days. Total days of hunting Seasons allowing only the longbow began as early as ranged from 5 in Kentucky to 120 in South Carolina. The September 14, in Florida, and continued as late as Janu- total season in South Carolina encompassed 140 days, ary 31, in Arkansas. Avcrage length of the specified but hunting was not permitted on Sundays. Maryland, archery seasons was 58 days. Four States provided an North Carolina, and Virginia also did not permit hunting archery season over 90 days in length. Archers were on Sunday. A similar law cxists in Georgia but evidently permitted to kill deer of either-sex in all open arcas in is enforced only on a local option. Seven States offered eight States. They were restricted to harvest of bucks less than 30 total days of hunting with rifle or shotgun, only in at least some portion of the open territory in one State offered 30 to 60 days, six States offered 60 the other States. to 90 days and one State offered morc than 100 days. Deer hunting begail as early as August 15, in South MANIPULATION OF REGULATIONS Carolina, and cont~nuedas late as January 19, in Flor- Control of the harvest is the most effect~vcmanage- ida A dyed-in-the-waol deer hunter i mith plenty of money and time) could flnd 158 continuous d'iys of deer ment tool available to the manager of white-tailed deer huntnlg in the 15-State area Range research and improvement are useless if over- harvest reduces the deer population far below the carry- The majority of States provided several seasons dif- ing capacity Conversely, knowledge of nutrit~onand ferentiated by season dates. Kentucky and Oklahoma reproduct:on are wasted if underharvested decr popula- confined all their deerhunting to a single statewide sea- tions exceed thc carrying capacity son. At the other extremity, Texas had 18 distinct sea- sons differentiated by season dates. What then should be the objective in using this tool? Other divisions of hunting territory in the southeastern It should be to provide the maximum amount of recrea- States were achieved by areas closed to deer hunting, tional opportunity for the greatest number of hunters differential bag limits, and restrictions as to sex of deer, without endangering the resource. use of dogs, and weapons. Many case histories have demonstrated that under- The entire harvest in six States was restricted to harvested deerherds will expand beyond the capacity of antlered males only. The remaining nine States per- their range. Adequate harvest is needed to control this mitted harvest of either sex in at least some portion of expa~lsion.With the large numbers of hunters available the State. Alabama and Louisiana prohibitcd the taking today, deer managers can control expanding deerherds of spotted fawns during the either-sex seasons. and could in fact practically eliminate deer from any given area if sufficiently liberal hunting regulations The definition of a legal antlered male during bucks- were set. only seasons also varied between States. Six States re- quired only that antlers be visible above the hairline. That's where the trouble starts. The setting of deer Seven States set minimum length limits ranging from hunting regulations has generated more discussion and 3 to 6 inches. Antlers with at least one fork on one just plain argument than regulations for any other game antler were required in the other two States. In Texas, species, with the possible exception of waterfowl. Deer the definition of a legal male deer varied for different managers almost invariably recommend more liberal areas. harvests than the public or even personnel of their own department are willing to accept. Greater similarity existed for shooting hours. Eight States had regulations which could be interpreted as For a well documented and thoroughly interesting meaning from sunrise to sunset. From ''2 hour before analysis of the biological, social, and political facets of the "rhubarb" which can result from attempts to liber- sunrise to 1/2 hour after sunset was the legal shooting period in six States. Missouri was most definite, setting alize deerhunting regulations (in New Jersey), I recom- the shooting hours from 6:30 AM to 5:00 PM. mend that you read Doe Day, by Tillett (1963 1. He concluded, "This little tale highlights the importance of Hunters in five States were required to submit their professional competence and skill in the direction of deer for checking at stations authorized by the game government bureaus and the administrative need for the department. qualities of persistence and courage along with quasi- political skill. The moral seems to be that the profes- ARCHERY SEASON REGULATIONS sional game-administrator should be a leader rather than Archery seasons provide maximum recreation with a captive of his constituency." a minimum of danger to the resource. Therefore, archery The quasi-political skill is of most importance in those seasons should be as long as possible without conflict States in which game departments must still answer in with other interests. All southeastern States permitted some degree to the legislature for their hunting regula- the longbow as a legal weapon during the regular fire- tions. Bartlett (19491 stated, "The answer apparently arms deer season. However, because the archer cannot lies, at least partially, in the fact that game managers actually compete with the firearms deerhunter, a special often do not have the power to manage deerherds prop- season for archers only is usually provided. erly. Such regulatory powers often are vested in an un- South Carolina does not provide a special archery deer responsive legislature-unresponsive possibly because season but, with the regular deer season open from game men may be poor educators." The exact powers and authority needed were outhned without endangering the resource. However, the re- by Hunter t1957), In dlrcusslng methods used to obta~n source can be endangered by underharvest, even morc hunter distr~but~onIn Colorado, "To obtain proper hunter than by overharvest. distribution it is necessary that a game and fish com- From my limited knowledge of conditions in the south- mission, through its Director, have actequate autliority eastern region. it appears that present deer rcgu1;itions for effective administration and management. Powers are adequate for controlling deerherds in all States exccpt of the cominission should include the authority to estab- Texas. Even there the problem seems to be one of eco- lish season lengths and dates, bag limits, shorten or nomics and landowner acceptatice rather than proper extend, or close seasons on any species of fish and game hunting regulations. within two 48-hour periods. designate the sex, species and What of the future? There seem to be two routes. The the number to be taken, the manner and have the right first leads to problems with overpopulations of deer in to open or close refuges to hunting." those States which continue to shoot only bucks. The A common theme was found in most ciiscussions of tradition of the bucks-only season, once established, is deer harvest regulations (Allen 1958; Bartlett 1949; difficult to eliminate and it is not too soon to begin the Longhurst 1957; Ruhl 1956; Swank 1962; Swift 19511. educational process necessary to prevent establishing This theme was a plea for more authority to set morc the tradition. The second route relates to the human liberal hunting regulatior-is and for more understanding population increase. The number of deer hunters in the of the problem by department personnel, the public, and United States increases annually while the total deer legislators. However, most of these papers were writtoi range shrinks because of changing land uscs. Tliis route by deer manager-adniinistrators of tlie Northcrn and leads to problems with hunter distribution and control Western States where deer populations had increased of the harvest. to the level where range destruction and wholesale star- Allen 119581 reported that only about 10 percent of vation of deer were occurring. They were stuck with a a deerherd is harvested under bucks-only regulations and buck law and couldn't get rid of it. All agreed that nearly 20 percent is harvested under any-deer regula- education of the public, of lawmakers and of their own tions. Illegal kill also becomes excessive in Statcs which personnel was of prime importance. The authority to have nothing but buck seasons. It would seem that under hold either-sex deer seasons was the major goal of ]nost a multiple bag limit of bucks, s~~chas exists in several of their programs. Longhurst (1957)found that the major States in our region, that the harvest would exceed 10 factors preventing control of big game herds by hunting percent. However, Teer et al. ( 1965) reported that des- were: shooting bucks only-18 States, poor hunter distri- pite longer seasons and inultiple bag limits the annual bution-13 States, public opposition-12 States, inacces- harvest in the Llano Basin in Texas averaged only 14 sability-11 States and legislative restrictions-10 States. percent of the standing crop. With a three deer limit, How do the statements and condltlons outl~nedabove one of which must be antlerless, the harvest took only apply to the reglon we are d~scussiirg? Some of the~r about 20 percent of the herd. I3;rrvest of more than statements regarding basic needs still apply However, :3O percent of the standing crop is needed to control a the overpopulat~onsof deer wh~cinconcerned the carller healthy deer population. authors do not exist throughout most of the southeastern Hunter distribution can be a problem even with auth- reglon Deer populat~onsIn much of the reglon are still ority for setting liberal seasons. In Missouri, we tried recovering from lows of the early 1900's and any over- to move hunters from an area near St. Louis by publicity populations are generally confined to local situations. to the effect that the area was overharvested and hunt- As Gresham (1969) said, "It is difficult to see how deer ing success was decreasing. The publicity was ineffec- hunting can continue to improve in the South year after tive, but when we changed the regulation to bucks-only year, but it does The ~inswer,of course, 1s that In many the hunting pressure was reduced by about 50 percent. States of this region it had a long way to go. Deer had We have also found that hunting pressure was affected been eliminated from vast areas of the South during by tlic day of the week on which the season opened centuries of poor managetntnt, and now good manage- iMurphy 1966). ment is putting them back." Are the current regulatioiis adequatc or is there still Coloracio has made an intensive effort to achieve good room for improvement? Swank 119621 concluded that hunter distribution. Hunter t 1957 1 reported that the conscrvative seasons with no provisions for postseason opening date, le~igthof season n~idtype of season (e.g. hunts, multiple bag limits, or either-sex seasons stamped either-sex, antlered only, antlerless only, two deer on a State as being antiquated in its big game program. a license, multiple licenses, two deer multiple licenses, Progress apparently is bci~lgmade to remove the south- pre-, extended and post-seasons I all were useful regula- tion manipulations to i~lfluencehunter distribution. The eastern States from tlie antique age. 111 1949, only Vir- ginia had an any-deer regulation (Bartlctt 19491. In multiplicity of regulations which I mentioned in review- 1957, five States had any-deer rclgulations but 1,onghurst ing current regulations indicate that deer managers of (19571 concluded that only two States were actually the southeast region are well aware of and are using controlling the growth of their deerherds by hunting. In these techniq~~eswhen applicable. 15168, nine States had any-deer regulations. Wit11 rcla- O~ictechnique which app;irently has not been neces- tivcly long seasons, lltrrvest gener:~Ilyrestricted to bucks sary, as yet, in our rcgion is zoning and quotas. This anti multiple season bag limits, it \voulti seem that tlc,cr system usually is necessary when tlic herd needs har- hunting regu1;ltions it) the soutl~easternrcgion are accom- vesting but hunting pressure is so great that unrestricted plishing the objective of providing maximum recreation numbers of hunters would damage the resource during an any-deer season. Several States in the midwest have 1968-1969. Winchester News Bur. Olin Mathie- adopted this system. We may see similar types of seasons son Chern. Corp. appearing iri the southeast rc:gion if the nurr~bcrof hun- IIuntcr, G. N. ters continues to incrctisc :it the present r;ite. 1957 'rl1(\ t~cllrriyiies rrsc-tl in Colorado to obtain I woulci )jreriict als~jthat tlic i~scof dogs For hunting hunter tlistrii~iiiiori. 'I'uer~ty-Seco~idN. Anier dcir will eventually be prohibitcci. 'I'r;itlition is hard Wiltilifc C'onf. 'I'r;iris. 15157: 584-593. to overcome, but in my talks with deer mantigers of ll~esoutheast 1 c;iri tielcct that cyvcL~lnow some of them I,ongI?urst. W. 13. would like to makc this restriction. I believe that re- 1957. The clfectiverress of hulltirig in controlli~~gbig striction on use of dogs will be brought about urllen game populations in Nortli il~ncxrica. Twenty- ntimbers of hu~~tersfar esceed the availahlc~resource. S~CY)II~N. Amcxr. Wiidlifc Corrf. Trans. 1957: 'The north-coritral States. wliich do not allow the use 544-5651, of clogs, have high numbers of hunters 1v11icl1 iri ~icxarly iaurphy, n.'4. every case caii control tlie dcerllerds vi-ithout tho use of 1966. Eflccts of various opening days on deer harvest clogs tRuh1 1956). and hunting pressure. Southeast. Assoc. Gan~e The Soutlicasteri~States have developed several fine a~idFish Corn~n.19th Annu. Conf. Proc. 1965: cooperative programs and coulci 1)rovitle ti~iotherexa~nple 141-146. by standardization of decrhui~tingregultitiotrs. A region- wide adoption of uniforn~ regulatiolls or, no~lrc~sident Ruhl, 11. D. license fees, dcfinitiotl of ;I legal dclcr riflc ;11lc13 reyuire- 1956. Huiiti~igthe ~vhitc~iail.In The clccr of North rnent that guns be unlondeci ;ir-td cased during transport America, 1,~.261-331. Harrisburg. Ponn.: Stack- are some possibilities. pole Co. Swank, W. LITERATURE CITED 1962. Manipulation of hunting regulations as a man- Allen, D. L. agement measure. J. Forest. 60: 30-32. Our wildlife legacy. 422 pp. N. Y.: Funk anti 1954. Swift, E. Wagnalls Co. 1951. Deer herd control rnetliods and their results. Allen, D. L. Forty-First Conv. Int. Assoc. Game. Fish, and 1958. How our game laws waste deer. Wildlife in Conserv. Comm. 1951: 126-133. N. C. 22:112): 4-7. Tillett, P. Bartlett, I. H. 15163. Doe day. The antlerless deer controversy in 1949. White-tailed decr resources, United States and New Jersey. 126 pp. New Brunswick, N. J.: Canada. Fourleenth N. Amcr. Wil(1life Conf. Rutgers Univ. Prrss. Trans. 1949: 543-552. Tecr, J. E., Thomas, J. W., and Walker, E. A. Grcshatn, G 1!)65. Ecology anti manaycrnent of white-tailed tlcer 1968 Reglonal report Tl~cSouth Wlnche\tcr wcst- in the I.,lano Basin of Texas. Wildlife Monogr. ern huntlng compendium for the lilllted States, 15, 62 pp. Managed Hunts by State Agencies A. Gordon Spratt Game Management Divis~on Florida Game and Fresh Water Fish Cornrn~ssi~,~~ Tallahassee, Florida

State game agencies conduct mcinclged deer hunts Colitrolled deer huilts are conducted on ~ipj~roxi~natcly on 15 nlillion acres of public and nri~xttelands in 15 million acres in Al:ibani;i, Arkansas, Florida. C;iv)rgia 12 Sozcthcast~rnStales. Managed hunts are csscntial Kentucky. Louisiana, Missouri, Mississippi, North C;lro- in the management of ujildlifc on recrcatioizc?l aruas. lina, South Carolina, 'ret~nesscc:, and Texas. The acre;igi The pl~hlichc~s accc.ptt~d the concept of controlled includes apl~roxirnately10 niillioil ;:cres of pri\~atc>lands hunting. Allho?~ghparticipating 1andowncr.s are and approximately 5 niilliot~acres of jiublic latitis. Uur- benefited, fa?~orublep~~blic relations and monetary ing the past 18 years there have hccn few significant returns must be continued and rnc~intc~int~dat a high- changes in the amount of pitblic lands available for this er Ic7)el in the future. Szcper~~isecihunting programs use. As an example, there were 4.2 million acres of are costl?], but are not US diffict~ltto justify as otl~er public land open to hunting in 11 Soiltheastern St~itesin programs and practices freqztently employed by 1951 iFrye and Swindell 1951) as coiiipared to the cur- Stale game agencies. rent estimate of 5 million acres in 12 States. The numbcr of supervised hunting areas on public lands has increased, State agencies are legally responsible for the reguia- but the data provided by questionnaire respondents do tion, control, and matlagement of resident wildlife ial- not permit a valid estimate of the increment. In 1951 though there are current disputes questioning this juris- there were 50 separate areas in 11 Statcs. There are diction ). Therefore, State agencies must provide the now over 60 in seven Statcs. In 1951 eight of the 11 leadership in maintaining wildlife resources in the best States required special permits arid six of the 11 States public interest (Berrymarl 1961). Effective leadership charged for the special permit (Mosby 19521. During the requires the development of programs which insure the 1968-69 hunting season 12 States reqilired special per- public interest. Increasing recreational demands, popu- mits. Six of the 12 issued free permits. lation growth, and habitat destruction have, however, made the task increasingly difficult to accomplish on MANAGED HUNT OBJECTIVES private and public lands which are available for public use. "Controlled" hunts are insuring a level of hunting The primary purpose of managed hunt programs is to opportunity not otherwise afforded. This form of hunting provide hunting opportunity consistent, with cominercial has become especially necessary and popular in white- and other recreational land uses. 13utttcr control, super- tailed deer management. vision, distribution, and deer mitrtc~gementare secondary For about 40 years supervised hunting has been used on objectives according to most of the States responding to specific areas wl~ereneeded as a management technique the questionnaire. liunting opportunity is being insured in order to provide public hunting at the same time. The and highcr levels of sustained use arc realized on areas practice has increased in popularity, especially for forest where managed hunt programs arc. implemented. game species, during the past 10 to 15 years and it is The degrees of control imposed on the hunter vary a subject of widespread discussion. widely and may be more varied than the objectives on Managed hunting is "any form of huntitig during which cach area where hunts are conducted. Controls often controls more rigid that1 those imposecl by general hunt- include special permits and rclstrictions on the number ing laws are exercised over the hunter on his method of hunters, h;trvests, methotls of t;iking wiltllife, carnj)- of taking various species of wild gamc!" iMosby 19521. ing, use of intoxicants, vehicle types. days and hours of Managed hunting is employed to manipulrtto hunters hunting opportunity, types of firearms, use of ct~tnl)- while managing wildlife. fires, and other activities. In additioil, hunters are often My literature review on the subject verified Mosby's restricted to specified zones withill open hunt areas and 11952) opinion that managed hunts are not reported in are frequently required to chcck ill and out and report the literature to the degree they should be. To obtain their kill. background material, we submittclci questionnaires to wildlife agencies in 15 Southeastern Statcs. All of the MANAGEMENT IMPLICATIONS States responded to the inquiries enabling the following White-tailed decr management is. as it is being prac- discussion on the philosophy and current status of man- ticed, depoident on current data regartling populations, aged deer hunts in the Southeastern United Statcs. herd dynamics, animal condition, and hunter prt,ssure atid success. The maintenance of tl~isinfortnation, par- CURRENT STATUS ticularly in the cxtensivc forcsted areas of the soullieasl. Approximately 30,000 decr are harvc.stcd annually on provides a reasonable base for man~~gemetittRipicy anti controlled huntitrg areas in 12 Southec~stcrnStates, and I-Inlls 1966). Sul~ervisedhunts provitle a good ol.)portull- an estimated 1.2 million mail-days of cieerhuntitig are ity for the collcctioil of lriallagement data. Checking provided. stations arc the most common means of obtaining the data (Thompson 1951). Supervised hunting is both desir- State administration and control of hunting in itself able and essential for intelligent wildlife manage~nent benefit5 the owner Ir~dividualhunting arrangements, of I-eci-eatioiial areas according to Mosby I 19521. lluntcr supervisioi? and eliforceineiit are responsibilities hat fe\r owners wish to accept. Thc majority of coop- PUBLIC ACCEPTANCE AND HUNTER SUCCESS crating individual and corporate landowners are willing to be relievecl of these responsibilities as well as those Although managed hunting means greatc,r reginicnta- involving wildlife mnn;~gement. tion, the concept is not as distilstel'ul to the hunting Liability relief is a niajor benefit and a prime incentive p~~blicas sotile resource acin~iilistrators may believe. to landowners in leasing land for managed hunting. Sonic people oppose everythii~g,but most sportsmeri Liinited liability relief is available to landowners who will abide by and accept the degree of control in~posed make their lands available to the public for recreational when they understand the reasons for the controls purposes in 10 of the 15 States surveyed [Fowler 19671. ~Mosby1952). Florida and North Carolina were the first to adopt legis- 'rhere are hunters who p1,efer supervised hui~ts.Their lation of this kind in 11163. It has been a clefinite help reasons include greater success, safety, quality hunting, in acquiring managerl hunting lands. more favorable hunting conditions, and the existence of Although thc relationship between State game agencies public use facilities (toilets, wells, developed campsites, and landowners is usually favorable, public hunting boat ramps, etc.). creates problems for both. The hazards of wildfire, van- The qucstion of public acceptance is of little conse- dalism, da~nagc,theft, and littering are diffi- quence in a few States where managed hunt areas pro- cult for owners to ignore. Consequently, private owners vide the only form of hunting opportunity. A significant are in a position to demand certain compensations and number of the hunters who visit the areas are satisfied State agencies arc often required to make additional on the basis that thcse areas afford tllcnl their only op- concessions when agreements are renewed. Cor~ccssions portunity. Ten States expressed unqualified public ac- may be costly and involve such things as damage pay- ceptance to managed hunt programs and two expressed ments, services, increased protection, and higher lease mixed public feelings. In addition, nine States indicated rates. that hunter succcss was greater and three States felt Problems on public lands are usually a result of con- that the same level of success is realized on uncontrolled flicting land use practices Itlowever, general land use areas. pollcies govcrnlng the use of publ~cland negatcs the need to discuss the few rnstances where publlc agencles LANDOWNER ATTITUDES are competing In the management of renewable resources Cooperative agrecincnts and leases with public, pri- and land use. Multiple use policies arc providing more vate, and industrial landowners are the foundation of access and additional huntrng opportunltles A review many recreational prograins (Frye 19671. Few State of managcd hunt regulations in the southeast indicates game agencies own an appreciable amount of public that fewer hunter controls arc cxerclsed on public lands hunting lands. Private landowners must benefit and and free access is more common than on private lands. receive compensation for the use of their lands. Their coopcration is necessary. The occurrence of a situation MANAGED HUNT COSTS where an individual owner feels a moral obligation to The cost of conducting supervised deer hunts varies allow hunters access to what is traditionally public prop- with the intensity of management and the degree of erty (game) is rare. There are, however, land holdings controI imposed. in the hands of realistic, civic-minded concerns or indi- viduals who recognize the problem of supplying hunting Supporting data provided by nine States indicated an and are willing to do their part (Frye 1967). This is average expenditure of 8 cents per acrc per year. The particularly true when the landowner retains complete cstimatcd costs providcd ranged frorn less than 1 cent control over his land and its particular resources that are per acrc to 18 cents per acre. The respondents indicated unquestionably his private concern (Frye 1967). that approximately 1.2 million man-days of deer hunting Although occasionally a few landowners become disen- are provided at an average cost of $1.08 per day. Cost cllanted with managed hunt programs, they are generally estimates provided by individual States ranged from 20 satisfied and all are benefited according to the 12 States cents to $3.25 per man-day. which rcspoiidcd to the questionitairc. Managed hunts are expensive to conduct. However, Better public relations and greater finaiicial returns State game agencies are making a direct contribution are the primary benefits to private ou7nc,rs. State agen- to the hunting public and the invclved costs arc often cies are sometinies ablc, by virtue of normal developmei~t easier to justify than other prograins and practices fre- and maintenance activities, to offer scrvices equivalent quently uncfertakcn, as pointed out by Mosby (1952). to financial support iFrye 1967). These services include fire protection, controlled burning, posting of perimeter CONCLUSION boundaries, fence repair, assistance in maintaining capi- The demand for supervised public hunting areas and tal improvements, and technical assistance involvillg intensive use of existing areas is increasing. The degree general land managcmrnt practices. Routine patrol by of huntcr control and the number of rnanaged hunts are State personnel provides a degree of control over, and expccted to increase proportionately. Managed hunting protection from, fence cutting, vandalisn~,rustling, lit- programs have been successful in providing opportunity tcring, trespassing, and theft. and controlling use. The ability of State agencies to maintain controlled hunting on private lands. Twenty-Sixth N. hunting in the future will be increasingly important, Amer. Wildlife and Natur. Resources Conf. particularly with regard to the acquisition of additional Trans. 1961: 285-292. areas. Large industrial landowners will continue to make their lands available if recreational use can be controlled. Fowler, K. S. The extent to which private owners cooperate in future 1967. Obstacles to the recreatiot~aluse of private for- programs is also contingent on the ability of State agen- est lands. Practiculn for M.S. Univ. Mich., Ann cies to regulate hunting. Favorable p~tblierelations and Arbor. 89 pp. the respect for private property are contributing factors and will require considerable attention. Frye, 0. E., Jr. The problem of providing participating private owners 1967. Cooperative progranis for public hunting on with additional financial return is critical. Private leas- privately ownect lands. In Forest wildlife man- ing is steadily removing managed hunt arcas from coop- agement, pp. 56-61. La. Strite Univ. 16th Annu. erative management programs. Public financial support Forest. Symp. Proc. 1967. La. Slate Univ. Prctss. is necessary ancl it can be justified. Public benefits in- cluding associated outdoor recreation opportunities arc Frye, 0.E., Jr., and Swindell, D. E. products of managed hunt programs that could bc sup- 1951. The southcastern public hunting program. Pap. ported by a larger segment of the public. To date, Ihc presented at Southcast. Assoc. C;nn?c, anti Fish hunter has shouldered the majority of the financial Conlm. Fifth Annu. Meeting. 11 pp. burden. Future programs will require a nlore equitable distribution of the financial burden. Mosby, II S State agencies would be renliss if they did not recog- 1952 Somc results of supervised foreit game hurlti nize the importance of a supervised hunting area as a in Virginla Pap prtsentcd at Eighth Annu proving-ground to dcmonstrate sound managcme~ittech- Meetlng Northedit Sect Wlldl fc Soc 8 pp niques for private enterprise. Private invcstment has already established a number of succe~sfulmanaged deer Ripley, T. H., and Halls, L. K. hunt areas In the southeast which are open to the public 1966. Measuring the forest wildlife resource. 111 on a fee basis Their continued success will encourage Measuring the southern forest, pp. 163-1!14. them to contlnue to voluntarily provlde for the fuller La. State Univ. 15th Annu. Forest. Symp. Proc. use of their lands "In the publlc lntcrest " 1966. La. State Univ. Press. Thompson, W. K. LITERATURE CITED 1951. The value of checking stations in wildlife re- Berryman, J. H search. West. Assoc. State Game and Fish 1961. The responsibility of state agencies in managing Comm. 31st Annu. Conf. Proc. 1951: 138-141. Effects of Timber Harvest and Regeneration on Deerfood and Cover

V. E. Carter and E. A. Dow Bureau of Sport Fisheries and Wildlife U. S. Department of the Interior Atlanta, Georgia

Forest manaqement activities on major", forest STUDY AREAS areas of the National Wildlife Refuge System demon- Primarily through timber harvest techniques, it has strute how sound si1z:lculture managrment tech- been possible for the Bureau to develop high-quality niques can be modified to prouidc timber, deer, and wildlife habitat conditions for deer and other species of associated b~nefitsnear oplinzz~rnlevels. wildlife on major refuges with sizable forested areas. A inultiple purpose forest environ?i~entproductive This group includes White River National Wildlife in timber, wildlife, and public rise is in great public Refuge, Arkansas; Noxubee National Wildlife Refuge, demand. Combined management efforts of both for- Mississippi; and Piedmont National Wildlife Refuge, esters and wildlife biologists are required if maxi- Georgia. The timber quality 011 these areas, although mum forest .nalzies and benefits are to be achieved. responding favorably to intensified management efforts during the 1960's, remains somewliat under par; the Timber cutting versus no timber harvest is no longer scars of mismanagement in the 1930's, prior to Bureau the baslc question concerning habitat improvement on acquisition, are still evident. Refuge forests are com- forested lands Now the prlmary concern of land man- mitted to multiple-purpose objectives; primary emphasis agers 1s more related to the effects of various methods is to develop a suitable forest environment for the broad of timber harvest on wildlife habitat Planned programs spectrum of wildlife, with special concern for migratory of management both In forestry and w~ldllferequlre a and rare or endangered species. Shallow winter close working relationship between forestry and wildhfe flooding of bottom-land hardwoods at White River and technicians When this exists, an enlightened land-man- Noxubee usually attracts more than one-half million agement and multiple-use program results The end ducks annually. These same lands produce deer and objective is to develop and to perpetuate a forest envlron- other wildlife and timber products near optimum levels. ment sultable for harmonlous product~onof both plants Management techniques on refuge lands are also modi- and animals fied to favor wood ducks and species like the red- Timber harvest and subsequent forest regeneration cockaded that are highly specific in their profoundly affect deer food and cover Lay (19671, habitat requirements. Goodrum ( 1949 1, Harlow and Jones ( 1965 1 Aldo Leo- pold et al. ( 1947) went so far as to say "brush in the sun" was always a necessary condition for deer irruptions in SOILS, CLIMATE, AND VEGETATION the East." A variety of soil characteristics, climate factors, and The other extreme is demonstrated on park-like areas numerous cover types makes it impossible to generalize where timber management has been limited to continu- on forest and wildlife recommendations tl~roughoutthe ous protection. Under this form of management, deer southeast. habitat eventually fails to produce enough food for The White River National Wildlife Refuge in eastern wintering herd requirements. Arkansas is representative of the Delta flood plain and In the past, research in the respective fields of timber demonstrates a rapidly growing forest of mixed hard- and wildlife resources has supplied general management wood species. guidelines for one or the other, but rarely has research Noxubee National Wildlife Refuge in castern Mis- provided specific technical recommend;ttions for com- sissippi represents a combination of forest types. Delta bined management. Considerable knowledge exists re- hardwoods, Appalachian hardwoods, pine and mixed garding the necessity of providing diversified food and pine-hardwood timber types of the Piedmont. cover requirements within the normal home range of deer and other wildlife. The 33,000-acre Piedmont National Wildlife Refuge This paper describes the timber management approach is typical of the pine-hardwood forest common to the being taken on national wildlife refuges to benefit deer Piedmont region. Generally, hardwood sites are on the and other wildlife. Special emphasis is devoted to the more productive soils which flank the streamcourses. even-aged management opportunities on one national About 20 percent of the refuge is suitable for commercial wildlife refuge of the Lower Piedmont. hardwood production; 75 percent is in pine; and approxi- mately 5 percent is maintained in fields or openings free of timber growtli. ACKNOWLEDGMENTS These refugcas arc widely distributed and include most The authors are indebted to refuge managers, foresters, of the major forest types. This makes them excellent and biologists who manage the iiational wildlife refuges proving grounds for research and offers tren~endous and provide annual reports on the renewable timber demonstration possibilities for niultiple-purpose manage- and wildlife resources. ment. TIMBER AND DEER YIELDS of the Piedmont Plateau. This pine, pine-hardurood, ailc hartlwood forest is managed under an 80-year rotatioi Thc multiple uic fo~c,itm,i~i,igcmcxnt progr~iln nou 111 effect on White Riber, Noxi~hcc,,rnrl I'ietimorlt reitiqcs with an :?-ycar cutting cycle. Regeneration units average aljproxirnately 50 acres, with a maximum cjf 100 acre! ii repre\cnt,itlve oi se\clial in~ijorforest type5 'lnd gco- gr,iphic condilioni of the iouthe,iit Theic combirretl iriiti :i n-Iiniinum of 10 acres. Cnder this inodil'icd even program5 w~llprovide ionic, iniight into the scope and aged iipproacii, 1 80th of the forest area is bein!< regen succci\ of timbcr ,inti wiltilifc nl,in,igernt>nt ,ii it relates crateti in small units oil tin ;inr-i~ialbasis. to timbrr harveit arici ricer yielrls Forest types are ftivoreti according to site cal,;lbilities 011 this basis. hardwood cotnposition objectives are tc Theie three major forcit

Wl~iteRiver Regenerat~ontechniques usually vary with each major Noxubee Piedmont forest type with~neach geographic area Foresters can Total pred~ctwith a reasonable degree of accuracy what regen- (.ration method is best suited for each local s~tuat~on Current timber harvest operations remove: only about Regenerated stands usually provide excellent teed~ng three-fourths of the annual growth. Once the timber arcas for deer and other wildl~feduring the early years stands reach optimum stocking levels the annual allow- of ctevelopment Once the young trees grow beyond the able cut will be balanced wit,h annual growth. These height suitable for fced~ngthey still provide excellent operations can best be described as improvement thin- cover for dcer ning~,which are tiesigned to increase the quality of Cromer and Smith 1 1968 I, reporting on the results of the forest environment for wildlife, timber, and people. a deer browse study in Appalachian hardwoods, indicated These periodic partial cuttings will fit into the broad that "any type of economical timber harvest apparently criteria of either all-aged or even-aged silviculture man- produced more than enough stems of preferred species agement. to support a population of at least 13 decr per square Prescr~btdburning, long rccogrii7ed ai ,In excellent mile without appreciable damage to reproduction of liab~tatmanagement tool 11-1 5outherrx p~neforcits, ii desirable tree species. Thus, the need to produce deer iitilized under the even-aged managemclit concept with- browse is not of vital importance to timber manrigers out conflict with forest regeiier,rtion icq~iirementi IIard- in this arca a~idtheir cutting practices can be oriented wood units located adjacent to pine are excluded from towards other objectives. Under situations wherz thc prescribed burning because of their susceptibility to fire objectives of game management envision a large deer- tiamage. Prescribed burning makes its greatest contri- herd, it is obvious that deer density will re,ich a point bution to forest management when utilized in conjunction when increasing available browst: would be a valid goal w~thperiodic th~nningssp'iced throughout the rotation of cutting." period Rased on the results of refuge forcst management prac- The 33,000-acre Piedmont Refuge is typical of the tices throughout the southeast, deer popu1;ltions of one flora and fauna found on approximdtely 26 mllllon acres dcer to 17 acres do not prevcnt regeneration from de- vcloping into quality stands. Where the forest is man- Artificial regcneration, tisually with intensive sitc iigecl iincier lo~~grot:itioris ;inti frccjuent cutting cycles, preparation. is utilized rather exteiisively in the Cciastiil cieer foocl is usually ;ihuiitlant, and ~ilthoi~glideer utilize 1'l;iin Stodtlarci 11!)371, Strodc tlil521, and r3~ck~ith rcgcneration ;ireas, the tlcgrec of use is conil~atil~lewith 119671 rei)ixted c~xcellc'iit tioer iise follo~~irigsitc prep- thc rlcvc~loprnc~itof ri~gcncrntion for conimercial pur- ;tr;ition by rnc~.li:inic;il ~nethotls. lxlses. 130th l>liiiiting ;iiitl sceiling 11;1\~ebeen usc~tlextolisively Scve~ity-l'ivc. i)c'rcc>l>t of the 100.000-;icrc h;irda.~ioti on l)r~p;11'~~(1~it(~s. Si~c~tlillg may hc.nefit decr niiiri, thari forest of the Wliitc, Ri~.crSatioiiii! Wiltllife Ii<~l'ii,qein 111:irititlg si1ic.c. inorc c~])sc rinfli~otlctl iircas sometimes csce~tione tlcor to 5 Itlr iiiiil~c~r~jrotluction. acrcs. Tlrcsc. floorled conciitions 11i:iy j)crsist for only ii fern t1;lys or rc>nlain for sc~yeralmonths. WILDLIFE RESPONSE TO MANAGEMENT Prior to hcginning timbcr h;ir.i~cstopc~r;iticiiis, this con- 'rhesc once, ticpletcd refuge lands were restocked with centratcd ciccrhcrd dcpletcd its foot1 siil~plyaiid hcgiili lvild 1~1rkc.yanti tlccr. Given ;i high degree of protection. to destroy ;igricult~i~.:ilcrops oil ;irij;ici~nt f';irml;incis. tliise species and other nrildlife responded very favor- Following Iigllt thinnii~goper;itions over ;i p;irt of this ably to ul,plietl habitat m;lnagement techniques. Im- iinflooticd nrca, hnI3it;it conditions inll~roveti;inti dcl~rc- provcments rcsultcd primarily from timber thinning of ctation of agricultural crops tlcclitic~ti. Although these both the pitie anef hardwood tinlher types. Prescribed initial thinnirig ol~cr;itii>nsimprovcci feeding cotirlitions, hurnil~g,in a:idition to timbcr cutting, in the refuge pine the large deerherd continued to dcstroy regeneration. forcst proved extremely valuable in improving forest Ileavier thinnings were cxtendcd over larger ;irciis, thus floor habitat contiitions. producing arlequate browse and sufr^icient regcneration survivecl for timber production. This situ;itirjn, although Expanding deer populations overflowecl to adjacent ownerships. and some animals were removed through somewhat unique, points out that corisidcrable latitude exists whereby the forc,st can be successfully regenerated trapping for transplant purposes. Finally annual hunt- ing programs were initiated to maintain population- under high deer population densities. h;ibitat balance. Regeneration practices should and most often are Approximately 25 percent of the deer population is selected primarily for reestablishing a stand of timber narvested annually. This percentage will increase con- for commerci;il purpciscs. The practice selected should siderably when crippling loss and unreported harvested be modified when required to fulfill wildlife or other deer are considered. In addition to the deer, other wild- multiple-use objectives. life have also benefited from intensified management. Partial cuttings and frequent cutting cycle intervals Turkey hunting is pernlitted on each of these areas, and are needed to develop and n>aint:iin high cjuality, con- annual squirrel hunts are conducted at White River and tinuous deer habitat conditions. Seed tree or shelter Noxubee Refuges. White River Refuge alone yields a wood regeneration tcchlliques at m;iturity are nothing harvest of more than 30,000 squirrels annually. more than a partial cut, and a ~nodificatinnof thcse prac- tices may be included in the rcgencration proccss of clearcutting. SUMMARY E;rlancei( ant1 continuous se;isonal habitat conditions Resource managers are now confronted with a greater urithin norni;il cruising range of deer arc of gre;rter sig- ecological, economical, and social challenge than at any nificancc to them than the rogc~ricrntion method used. other period in our history. Single-phase forest man- p..ittlal - . cuttings arc employed on wiltllifc rcfugcs to ;igcment should yield to multiple-purpose management. provide the continuous habitat rcyuircd. Evell-agctl man- Sound silviculture principals permit considerable latitude agcmcnt---now the ~~rc,don~inatitsilvicillturc: system-- for developing habitat for wildlife--especially for decr. call hest be pcrpc,tuatod through c1c:ircuttilig ;it rotatio~l's The merits of the even-aged forest management sys- cnd. Species composition, distributiol~,and iinit size tem for timber production have been proven and ac- control of even-aged stcintis may bc the best methods of cepted by boll, tile USDA ~~~~~t service and industry. 1n;iintaining habitat h:iliinco withili nortnal deer range. ~l~~ future Opl,ortL~liitiesfor forest management Seasonal variety call best br provitied Ily illnit clear- rest with this silvicultural method. cutting uliclcr tlie even-aged ct)ncept. Interspersion of The multiple-purposc forest ~nanagement program smt~llst;intis of scil't\voorls ntid h~rrtlwootls along with on National wildlife rcfugcs has demonstrated that both pnj~erage ciistribution car] best he ~jrovidcdunder this timber and wildlife can be produced near optimum levels system. Prescribt:d burning is more acccy>t;ible ~i~ider under sountf silviculture practices without serious con- the theory of even-ageti managenlent than untier the flict. Refuge forests are managed under long rotation and coliccl)t of a11 aged nianngi~metit. Fire can be more read- frequc.nt cutting cycles. Aside frorn keeping the forest ily ai)l)lieci 111 thc softwooti type ;inti excliidcd from the in a vigorous growing condition throughout the long kard\vood stands, which is highly susc.el)tible to fire rotation, thinnings provide an economic rttulqn and damage. Even-ageti units should be large enough to proper crown spacing allows sufficient sunlight to reach be mantiged economically arid small cno~ighto insure the forest floor enabling growth of food plants for deer habitat variety. and other wildlife. Even-aged forest management on the Piedmont Na- to wildlife. M.S. Thesis. Sam I3ouston State tional Wildlife Refuge, as illustrated in a recent brochure Teachers Coll., Hutitsville, Texas. "Piedmont Wildlife and Timber" offers one example of how harvest and regeneration techniques of even-age Harlow, R. F., and Jones, F. K., Jr. silviculture can be modified to hcnefit deer kind other 1965. The white-tailed deer in Florida. Fla. Game wildlife. ;xiid Frcsh \rv',il<~i.Fish (Iriniiii Tcc,h. I31111 !I. 2-10 pp. Sound silvicultural priictices are vital to our Nation, as are wise use and enjoyment of the renewable forestry Lay, D. W. resources. Timber regeneration in small units at the 1957. Some nutrition probier~isof deer in the southern end of a sawtimber rotation can provide a s~~itableforest pine type. Southcast. Assoc. Game and Fish environment for optimum deer management. Colnm. 10th Antiu. Conl. Proc. 195fi: 5:3-58.

LITERATURE CITED Lay, D. W. 1967. The importance of variety to southern deer. Beclcwith, S. L. Southeast Assoc. Game and Fish Comm. 10th 1967. Effects of site preparation on wildlife and veg- Annu. Conf. Proc. 1964: 57-62, etation in the sandhills of central Florida. South- east. Assoc. Game and Fish Comm. 18th Annu. Leopold. A,, Sowls, I,. K., and Spencer, D. L. Conf. Proc. 1964: 39-48. 1947. A survey of over-populated decr ranges in the United St;ites J. Wiltiiifc M;in;ige. 11: 162-177 Cromer, J. I., and Smith, C. li. 1968. Sufficient deer browse produced by a wide Stoddard, If. L. range of cutting practices. Northeast. Sect. 1937. Use of mechanical brush-cutters in wildlife Wildlife Soc., 25th Northeast. Fish and Wildlife management. J. Wildlife Manage. 1: 42-41. Conf. Trans. 1968: 25-33. Strode, D. D. Goodrum, C. W. 1952. Experimental habitat improvement. Fla. P R 1949. Eight methods of tirnber harvesting In relation Proj. W-32-R Job 1-E, :3 pp. How Size adDistribution of Cutting Units Affect Food and Cover o"Feer

Burd S. McGinnes Virg~niaCooperat~ve Wildlife Research Unit ' Bureau of Sport F~shertesand W~ldltfe U.S. Department of the Interior Blacltsburg, Virginia

Decisions relating to the size and distribution of on deer than other cuts, determines stand distribution, cutting units are dictated by several considerations. and is crucial to habitat composition for future deer Multiple Use Guides in use by some southern forests populations. are presented. Shape of clearcuts is discussed as a Before optimum size and distribution of cuts can be necessary element in the planning for cutting. A determined, several other factors should be considered. hypothetical situation is offered which indicates the The average size of a deer's home range must be taken maximum number of deer home ranges which are into account. Cuts so large they remove all timber from contained in clearcuts of three sizes and three dif- the deer's range may be detrimental. Deer will be forced ferent widths. Generally, long and narrow clearc?~ts into new habitat. Very small cuts will have little if any affect more deer thc~nlarge square or circular cuts. beneficial effect upon the animal, although this condition Smaller cuts of 25 to 50 acres are more desirable. is difficult to envision. Clearcuts up to V4 mile wide appear to be reasonable. The home range for whitetails is smaller than was Cover for deer in southern forests is not considered believed a few years ago. Zeedyk (1968) reported that to be a particular problem, especially when unre- deer use only 100 to 300 acres. A study by Petcher (1967) generated strips are left on each side of clearcuts within a 2,400-acre enclosure in Virginia revealed the and where cuts are small. average home range size of adult female deer to he 172 ACKNOWLEDGMENTS acres. Wohlgemuth (1968) using telemetry reported that a doe's range was 140 acres, and a buck's, 340 acres Thanks to my colleagues who read the manuscript, during the rut. The Multiple Use Guide for National provided literature, and criticism: R. H. Giles, R. L. Forests in Texas suggests the average deer range to be Downing, R. F. Harlow and M. L. Dahlberg. Special approximately 1 square mile (640 acres). thanks are due graduate students R. B. Hamilton, M. C. Perry, and J. L. Sandt for drafting the figures and assist- ance on the hypothetical problem. DISTRIBUTION AND SHAPE An obvious fact in studying the question of size and Cutting units should be distributed to provide maxi- distribution of cutting units, is the paucity of literature mum benefit to the greatest number of animals; this on the problem. Where research facts are lacking, man- implies a wide distribution through space. Shape of cut agement practices may be based upon opinions evolved areas can have a marked effect upon deer habitat. Al- from experience and dictated by economics. One point though the question of shape was not part of the assigned should be evident from this paper, research is needed on scope of this paper, it cannot be overlooked. Any dis- the question of size and distribution of cutting units. turbance to the forest which alters the vegetative pattern It is imperative that facts be forthcoming since even-aged produces an edge-effect. Edge is a valuable byproduct management is a proven forestry practice. of cutting. Long narrow cuts will provide more edge The South contains vast acreages of many forest types; than will square or circular cuts. Table 1 shows the however, this paper will not be specific for types but peripheries or edge which would be provided by area will consider only the broad aspect of the problem. cuts of different shapes, each 1 square mile in size. The It is my intent to provide guidelines from representative southern forests, philosophies behind management pro- cedures, a review of research reports, and an examina- Table 1.-Perimeters or "edge-effect" in different size tion of hypothetical situations into which are inserted cutting units each 1 square mile known factors. Hopefully, from this synthesis will come Shape and dimension Edge stimulation for further research, new philosophies, and Mile critical evaluation and reorientation of existing forest wildlife management policies. Circular Attention will not be given to intermediate cuttings. 0.56 mile radius This does not mean, however, that intermediate cuttings Square have no value to deer; they can enhance the forage sup- 1 mile x 1 mile ply. I assume, however, that the forest management is Rectangular even-aged; that the regeneration cut has a greater impact - 3/4 mile x 1 mile ' Supported cooperatively by the Bureau of Sport Fisheries ?/z mile x 2 miles and Wildlife, Virginia Polytechnic Institute, Virginia Com- mission of Game and Inland Fisheries, and Wildlife Man- % mile x 4 miles agement Institute. ?hmile x 8 miles narrower arid longer the cut, the greater edge ~t provides chain strips uncut between the clearcut strips. The [In- The 5hape least procluct~vcof edge ir the c~rcul~ircut treated strips will be cut 20 to 30 yetirs 1;iter. Openings, natural or crcated, should be examined for Virginia's Jefferso:~ National Forest policy rcquires their contribution to the problem of size and distributioi~ that not ovir 25 pcrccnt of a comp:irtmeiit ~villhe re- of cuts. Clearcuts are essel1tially like openings for a gcneratecl in one cutting cycle 1h1~ilcolm;iii I E:iw.irds, few years and the optimum size of these openings (cutsi personal commuiiic;rtio~~~.Regeneration 3rc.a~sh:,~~itl bc call be determi~~cdwhcn we lcnow how far dcer will less than 20 chains wide, wlieri possible. IS they must venture from covcr into open terrain. be wider than 20 chains, key wildlife areas of 1 or more acres are to be retailred as iriclusio~is. An attcmpt FOREST SERVICE GUIDELINES is miicie to retain 50 percent oS cach cornpart!-nelrt in mast producers. This forest has the rule-of-thumb that How large shonld 21 cutting unit be to provitic the maximuan footl and cover for deer? This is a major for exrcry 40 acres of area \rihich is cut, 1 acre must he question on which thcre is little information. The USDA Icft ilncut to pro\ idc ior do1 trce5, cover and n1,iit Forest Service, however, has gc,nernl guirlelines for their Although the Guide for the N:itiorial forests in Texas operations. I 1967bi statcs tliat the norn~alrange for tlcer is approxi- The Forest Service i 1;inribciok for Region 8 I l9GHa mately 1 square mile, no size or distributicin patterti for directs that in plaiining ;ind conductirig stand regenera- cut areas is defined. They do, llo\t,ever, prescribe a tlis- tion projects, work will be directed toward the creatiotl tribuiion pattern for hzirdwood areas that will provide ol' even-aged timber slarrds of 30 to 200 acres. R(,gc>nera- hardwooti mzist within the ~iormtilrange of deer through- tior? of large lover 200 acres) contiguous stantls in any out the forests; hardwoods u-ill be rctained anci dc- 10-year cutting period is to be a\,oided where possible, vcloped. Maximum allourable distance between liard- and two or more rc.gc!neratcd arcxus in a comp;irtn~cntor wood areas in pine stands is ' 2 mile. Th- I~arclwoocicorn- in adjacent compartments are to be separated by unrcgen- ponent should be 200 square feet per 40 acres, either in crated areas of stand size. Regenerated stands as small rlurnps or intlividual trees. as 20 acres are permissible where justifiable, but should The Big Levels W~ldl~feManagement Area on the average not less than 5 chains wide. There is no limita- (;eorge Wdihington N,il~onal Forest In Virg~nia15 re- tion on the maximum regenerated area, but a decision ceiving the even-agt>d system of tirnbcr manager~cnt to regenerate any stand larger than 200 acres must be where cle,~rcutting ii done in block5 of 20 to 201 acrcc; supported. il'hornton 19691 When tlmber 13 clearcut 111 relat~vely Multiple Use Guides for a sample of soiltlierii National large hlocki, logg~ngor pulpin? is econom~cal The num- forests are given here. The Daniel Eoone National Forest ber of sales is incrcnsing annually and tile acreage cut in Kentucky specifics that rcgener;ition treatments is provitling much-xrccdecl browse for deer and cover for should work toward creatit1g even-age stands of 40 to 200 othcr game species, ciccording to Thornton. acres ( minimum 20 acres 1. Cuts in excess of 200 acres require approval. Tlre guitleline of this lorest provides no SIZE OF CUTS restrictions on shape of cuts since starid boundaries norm- ally conform to topographic, site, soil, cultural features, So little work has becn done t,o ascertain ol~timunisize and other practical limitations. The Daniel Roone Forest and distributioti of cutting units for white-tailed dcer Guide (1968) further states that regenerated stands will that ;I study in spruce-fir forcsts in Arizona takes on be separated by and interspersed with urrregenerated significance and may provide useful knowledge for man- stands preferably 40 acres or larger in size ! 20 acre mini- agement of white-tails in our southern forest ! Reynolds mum) Normally, not more th'ln 30 percent of a com- l966b1.Reynolds measured accumulated dropping groups partment need be reger~cratedIn the same cuttlng pcr~od from mule deer, elk, and cattle to evaluate use of natural Cutt~ngin adjacent comp,lrtmcnts In the i'ime 5-year iil~cicreated openings. ?'he lattrr wcrc blocks of 10 to 30 per~odIr to be avoided At lea\t one itand 111 c'ich com- acres that Ilad been clrarcut about 6 years previously. partment should be regenerateti during each cutting Relative use by cleer cleclirleti sharply as size of oljening period where stands qualify for regc~tierntiorr cutting. iiicrcascd. Areas Iargctr than 20 acrcs were little its'd; National forcsts in Florida 119661 spcciSy that the as openings becair~esnialler, relative use i~lrreasetl.Rey- riolds co~icludeti that theoretically. circular openings manager will leave uncut sm;rll ( 20 acres or less i isolated stands of oaks and all hardwoocis along ponds. bays, lakes larg~rthan 20 acres (526 feet radius) woulti be little or streams. These uncut areas should contain 200 square ilsed, except next to forest borders. Tliis study implic!d feet of mast-producing stems per 40 acres. Live oaks that for best 1labit;it eflects from even-tiged tirnber 111an- up to a stoclring of 200 sqil;irc feet 1)er 30 acres arc to be agenicnt, clearcut arcas shoulri be widely disperseti, less left uncut. Tlle It'lorida guitlelinc is specific iri dirccting thnii 20 acres in size if ir~patchc>s, and lcss t1l:iii I 5 riiile across if in strips tliat the land m;inager rcgerlornte 110 more than 160 acres 1320 acres in sand pinei iri 1 square mile in one cirtting Rcyrrolds il9Gfiai reported ariother Arizona study in period, scrub oak excoptctl. This is ir significant figure lioritlorosa pine which 11ns a direct bearing on size of bec;iuscx if the hon~cx ra11gc of a white-tail cxncornl~:isscs ciitling tinits. Dropping groups again ~vercused to com- 1 square mile, the niasirtiiirn ;illowsl)le cut wotilcl he pare usiLof tirnbcreti arcas ;ind 11:it~iralopcl~iiiigs ~vithiri one-quarter of thc deer's rang(:. This a1,pe:rrs to he ;I a cutovcr i'orest. ilc,er droppings Lvere slightly higher rensollablc portior-r to bc, rcy,clierated from lkle vio\vl)i~int in orw~rii-igstIl;iri witliil~the forcst for ahollt ti00 feet of the deer's welfare. 'i:irtlwood tylies are to be trcatt~d from the forost 1)ordi.r. I)ec>r droppirrgs Lvore absent by clearcuitilig strips 20 cIr;ii~is wide nud lc!aving 20- beyond 1,200 feet irrto the oponitlg. Rcynolds thought that these forest border relations suggest that distance more than the uncleared plots. The investigator con- across an opening influences use by deer, i. e., they do cluded that "complete clearing of tracts as large as a not like to get too far from cover. Reynolds stated that square mile, is therefore, detrimental to white-tailed "evidently deer and elk do not use openings to any extent deer, at least during thc early successional period. Leav- that are more than 2,200 to 2,600 feet across, respectively. ing from 25 to 50 percent of the native vegetation on For openings to be used as high or higher than adjacent the plots makes them more attractive to these animals forest for both deer and elk, distance across an opening than uncleared areas." We could interpret these findings should not exceed 1,600 feet." The management implica- to mean that a clearcut of 640 acres would be relatively tion from this study is that openings less than 1,600 feet unattractive to deer. However, if 1 1 to 1 r of the square across would best coordinate dccr habitat improvement mile were left in uncut strips, we could anticipate dcer with timber management when clearcuts are in strips, use to be double that of the larger clearcut. While these blocks, or natural tree groups. data relate to planted clearings, one fact emerges and Harlow and Downing (1969) studied the effects of size that is, a 1 square mile clearcut appears to be too large and intensity of cuts on some deer foods in the Pisgah for dccr. National Forest. Although comparisons were made be- A clearcut or any disturbance of the overstory sets tween clearcuts and heavy selective cuttings, the clearcut back ecological succession and greatly increases seedlings, data gave some indication of optimum size of cutting sprouts, grasses, and forbs. Forest openings are gener- units. Stands of 1, 21, 51, 54, and 55 acres, nearly 1 mile ally created by clearcutting and inay or may not be apart, were clearcut in an area of high decr density. An planted. It is during the first few years following clear- index of deer activity was secured during winter by cutting that these areas are similar to created openings pellet group counts. Vegetative transects provided data in vegetative changes and attractiveness to dcer. Studies on seedlings and sprout numbers 1 and 3 years after of openings in the forest can help us to resolve the ques- cutting. Three years after cutting, the three areas of 50 tions of size and distribution of clcarcuts. Wisconsin or more acres contained such a tangle of sprouts and vines biologist McCaffery (1967) reported on a study of deer that deer could not penetrate into the interior. Browsing use on more than 100 openings. Data consisted of counts declined significantly in the I-acre clearcutting because of droppings in fall, spot-lighting from spring to fall, of disappearance of desirable plants due to intensive util- and road track counts. The openings were used inten- ization. The 21 acre clearcut appeared to be superior to sively during spring and fall but little during midsum- either the 1 acre or the 50 acre clearcuttings. The work- mer. As many as 125 to 150 deer per square mile of ers concluded that when too large, clearcuttings produce openings were spotlighted during May, September, and overabundant woody growth. After 2 years the cuts October; surrounding forest densities were 25 to 40 per became dense and unattractive to deer. When clearcut- square mile. Areas of several square miles in size with tings were too small, the more desirable browse plants few or no openings had less than 10 deer per square mile, were soon reduced in numbers. Perhaps the best clearcut while 25 to 40 per square mile werc found in more favor- size is between 20 and 50 acres in the Pisgah area where able areas. These observations would indicate that open deer are abundant. land is necessary for good deer range. Significantly, Zeedyk's ( 1968 ) appraisal of even-aged management McCaffery reported that small openings, less than 5 acres asked the question of how big a clearcut should be. He in extent, were used more intensively than larger open- answered that "cuttings should be big enough to insure ings. regeneration of desirable timber species yet not so large If a clearcut were 5 acres and square, it would seem as to waste the valuable browse resource." Zeedyk points reasonable that deer would readily traverse and use the out that since a stand is regenerated only once in a rota- 466-foot width of open terrain. The greatest distance a tion, we cannot afford to squander the rich supply of deer could travel from the forest edge in such a situation food created by clearcutting stands that are too large. would be 233 feet. A 5 acre clearcut may be too small to He warned that the bigger the clearcuts, the more of the be economical, except in unusually productive situations; resulting stands there will be back to back. I-Ie urged however, there is evidence that deer make intensive use that clearcuts be scattered, be kept as small as is econom- of these smaller breaks in the forest canopy. ically possible, and leave some clumps and patches of Since decisions relating to size and shape of clearcuts unregenerated stems within stands. are not based on data, I devised a hypothetical situation. Beckwith's (1967) study in the sandhills of central Assumptions were made that deer were evenly distrib- Florida, an evaluation of the effect of site preparation uted on three areas with annual ranges of 640 acres, 160 on wildlife and vegetation, offered data on deer use of acres, and 40 acres. Widths of clearcuts were 1% mile, treated plots which were large enough to compare with !:% mile and $5 mile. Sizes of cuts werc 50 acres, 200 cut units. Three replications of four plots, each 1 square acres, and 500 acres. Each rectangular cut of established mile in size, were studied. One plot of each replication width had its length determined by acreage of the cut. was completely cleared of vegetation and planted to Templates scaled to the size and shape of the cuts were slash pine. The second plot was three-fourths cleared placed on a grid representing home ranges of the three and planted; the third plot was one-half cleared and the sizes. The question to be answered was: What is the fourth plot was left untreated as a control. Uncleared maxinium number of deer ranges encountered by a given portions of the second and third plots consisted of strips size and shape of clearcut? Table 2 represents the re- of natural cover 10 chains wide. Deer use was estimated sults of this test. Figures 1, 2, 3 show how the clearcuts from track counts. Partially cleared plots received about of different widths and acreages intersect different size twice as much use as the cleared plots or 40 percent deer ranges. Figure 1 -Hypothetzcal sztuatzon of nzazzmum number of Figure 3 -Hypothelzcal sztuc~tzon of ma-rznzum izunzbei deer ranqes (one deer per 640 acres) en- of deer raitqes (one deei per 40 acres) en- countered by 50(Ai, 200(B), 500(Cj acre counteied by 50iAi, 2001B), 500/Ci acre clearcuts, each '4,'3, ci~td'2 mzle wzde clearczcts, each i,' ,, and '2 n~zlewzde

Annual ~-p a.rcs a.rcs 500 acres rrrnges r t1s 111111 + n~illztn!--I~h rnl.!l I !n1112 !:!I. ls~n71-11t 1,,!1;~I:,]- 640 ;rrrrs 4 4 4 H 6 41610 7 160 acres 6 4 4 13 8 6 28 16 10 40 acre 8 7 ii 22 14 14 5.5 :32 24

The narrower and the longer the cut, the greater the number of deer ranges affected Where widths are nar- row, in all probab~lltythe clearcut would not drive the decr from its home range, howcvei, a clcarcut strip '2 mile wide m~ghtconceivably dlirupt a deer'\ home rang[, p,ittern and cause abandonment of the ~rllmediate.lrea The '5 milc wide clearcut affected the largest number of decr in each case, however, the length of this narrow strip becon~esimpractical when large dcreages are in- volvcd cspeclally in rugged teir,~in The 200 acre cut 'h mlle wide 1s 2'2 miles long, the 500 acre cut of th~s width is 6' 2 miles in length From the foregoing, either the 1 h or I mile w~declear- cut appears to be a suitable width If ~istrip is 1 u mile wlde (660 feeti, decr could ncvcr get more than 330 feet from cover In the case of the I milc wide \trip, a deer would be 6fi0 feet from cover 111 the mlddle of the clenr- cut Thcie wldths would trot dcitroy a great percentage of any one deer's range, rather the total edge would bc great DEER COVER Harlow, R. F., and Downing, R. L. 1969. Tile effects of sizc and intensity of cut on pro- Tho subject of dcer cctvcr was not strcxsscd in this duction and utilization of some deer foods in ls;ij)er. WIiilc cover docs not a11pe;ir to bc as important the Southeril App;il;ichians. Twenty-Sixth to soutllerri tic,er as it is to cieer iii the North, tiicy need Northc:rst Fish anti Wildlife Conf. Trans. 1969: it to escape from cold winter wiritis and Ili~rlters,and 45-55. for shade, fawning, anrl resting. Usually, cover is suffi- cient over most southc.rl1 dcc,r r:rngc?. Creek bottoms, McCaSfery, K. sm:ill hollows, 2nd coniferoi~sst;iiltls o11c or more acres 1967. Bargain openings. Wisc. Conserv. Bull. 32151: ill size provide gooci cover, li' cutting is done in strips 21-23. anti if utlregenerated arc,i\ of it,inti wirith arc left urlcut, I'etcher, R. L. thc.rc1 sllould bc no 1,ick oi ,rdcyu,rtc~coxrei Furthermore, l!Xi7. A polpul;rtio~r study of a confined deer herd. >lash left fioin cutting opcr,itiori\ rrr'iy jiio\ ide immed~,itc M. S. Thesis. Va. Polytccl~.Inst. Blacksburg. covcr Where hard\+ootls iprout rc><~ti~lyor seedlings 77 pi,. gx)w r'ipidly, cover will be provided shortly after the cutting Reynoldi I1 G 1966,i U\c of ponderoi'~ pine forest in Ar~zonaby CONCLUSION clccr, elk, aiict cattle USUA Forest Serv Res Note RM-63, 7 pp IZocky Mountain Forest and Even-aged manageinent of forests is the, order of the RL~ng('Exp Sta , Fort Collins, Colo day. To br most cffcctivc in dcer maliagement, long range plani~ingwill be required to give a good distrihu- Rt:ynolds, Ii. G. tion of stands in all stages of succc>ssion. Cuts nlust be 1966b. Use of openiiigs in spruce-fir forests of Arizona big enough so that some rel)rocluc~io~iwill survive deer by elk. deer, and cattle. USDA Forest Serv. Res. browsing if deer arc numerous, but riot so large as to Note 1334-66, 4 pp. Rocky Mountain Forest and permit a waste of food. Tlle rcgc.ncratio11 cut is the im- Range Exp. Sta., Fort Collins, Colo. portant one. Mast producers must be considered and provision made for a constant supply of this important Thornton, J E wildlife food throughout the area utlcl<.r management. 1969 I3ig I,evel\-a game managemelit laboratory The wildlife mai1agc.r must take into account the density Va Wildlilc 30t2) 4-5,23 of deer in his area, tllc, type of vegc>tation, the terrain, 1JSDA Forest Service. anti the economics of logging bcfore decidiilg on what 1966. Forest Service Handbook. Multiple Use Guide, size, shape. and tiistribu1,ion of' cuttillg rulits to en?ploy. Nat. Forests in Fln. No two situations will be alike. Gcncrnlly, the smaller cuts 125to 50 acres) will be most dtsirablc. Long narrow USDA Forest Service cuts give greatcr cctgc and bciici'it more deer than widc 19(i'iC1 Fore5t Ser vice Ii,lntibook Multilde Use Guide, cuts. or large square or circu1;ir cuts. 0ptim;illy. a clear- Reg 8 cut should be no more than twice tile distsilce whiclt USDA Fore5t Service a dcer will move from the Sorest etigc; this appclars to 1Yb7b Iqore\t Service H,indbook Multiplc Use Guide, be 600 to 800 feet. Clearcuts up to i mile wide appear Nat Forest5 In Tex to be reasoilable. The suggc!stion to have an unrclgencra- tcd strip on each sidc, as uritio 21s the clcorcrrt, ofiers USDA Foreit Servi~e. cxcellcnt distribution of cuts, provides cover, and may 19fifl Fore5t St,rvlce Ilandbook Multiplc Use Guldc, contribute to the vital mast supply. U,~n~elBoolie N,rt Forest, Ky

LITERATURE CITEQ Wohlgemuth, I) 1968 W1rc.d tor iountl Ga G,ime arid Fisll 3t214-6 I3eckwith. S. L. 19ti'i Effect of site l>rcp;rr:ition on ~\.iltilii'c~ailti vege- Zcc tiyli W U t;ition ill tht, s:~ntii~illsof rc.r>tr:il E'loritl:~. So~ltii- 19(i!l Wrldliie untle~ c\ elr-'igc.tl n?,rn'~gc~ment P'rp cast. Assoc. C;;,me :inti Iq'isli ('oil~m,1 llt 11 AIII~LI. pi cscntcd .~tIiy -'l'c1111 Sec Soc Amer Forcit Conf. Proc. 19fi3: 39-48, 23th Arrnu Meeting 16 pp Short and Long Rotations in Relation to Deer Management in Southern Forests

Phil D. Goodrum Bureau of Sport Fisher~esand Wlldlife U. S Department of the Interior Nacogdoches, Texas

It seertbs eviderlt frotn t.esec~i.cl~IJL(II jot.est ttl(oli1)- Tile trend in forest rnan,~gelneilt luwLll.ti ol~esljecies lati ti on through long-rotation mc~nagemcntwill pro- stands strongly indicates a need for rn:iriil~~il;itionsth;it duce a better deer habitat thun short-rotc~tionn~an- would produce better deer habitat. ugenzent. Long rotations offer greater flesihilifg in the ztse of manclge~nenttechniqzces for producing ad?- REVIEW OF LITERATURE quale amounts of nzctritionul forage, nzc~st,rind other In a young pine plantation in Louisi:rii;l that w:is sub- foods over a long und co~ztinz~ingperiod. 1,ong-rotcz- ject to short rotation, Blair 119631 fo~itid that heavy tion ~nanngementalso perrnits prcseruirzg and nznin- thinnirig j-iroducetl 52 percent more palatt~blcbrowse th:iii taining a variety of plunt spccies and stand condi- light thinnii~g,but that unpalatable browso was about tions wlziclz are essential to u good deer Izcrbiitrt. In the slime in light, mediurn, and heavy thirrnii~g. 'Tile short-rotation manugelne?zt, browse rna y be prodz~ccd plantation containcd no mast-bearing trclcs. in great ab~cnduncefor short periods of lime. Krefting t 1962) cited a study in a mi:itci coniler swarnp Deer management involves two main considerations : in Michigan that shoured clearcuttillgs in strips ant1 shel- the manipulation of the herd and the manipulation of terwood cuttings produced 161 and 184 pounds of browse the habitat. per acre, respectively, while no cutting and light sc?lec- tive harvesting produced 46 pounds per acre. Block anti Manipulation of forest habitat for deer will be dis- diameter harvesting yielded 130 and 134 l~ounds,respec- cussed in this paper with emphasis on short- and long- tively. Ilis study of deer use of these ciittings intlicatcd rotation harvesting, the two prevailing systems in for- greater use in the 66-foot-wide strips adj;iceilt to urlcut estry. Short-rotation management may be described as woods because of better cover in the untreateti area. the harvesting of forest trees at comparatively early The experimental area containecl no hardwood mast- ages, ranging primarily from 15 to 40 years, depending bearirlg trees. E-Ic also said that "uneven-aged st;inds upon the species. In long rotation the trees are harvested produce a larger variety of habitat conditions favcjrable when larger and older. in the range of 40 to 100 years to deer than large even-aged stands." or older. Size and height at a given age will depend on the fertility of the site, which may influence the Bryan 119501, writing on the rcsults of experimcnt;il length of the rotation. cuttings in the Appalachians of North Carolina to im- prove deer browse, observed that clearcutting produced There are four criteria in evaluating a forest habitat. too much browse and other growth, anti that light selec- for deer: tli Abundance of palatable forage. (21 Produc- tive harvt.sting produced too little. IIe concluded that tive capacity for mast, including soft fruits. 13) Variety selective cutting somewhilre between these extremes on of vegetative species and mixture of types. (4)Condition small ilcreages appeared to be tlie best procedure. of vegetative cover. Zeetiyk tlY68i suggested short-rotation cle:ircuts for browse production with not more than 10 j)c.rcent of an area being harvcstecl in this way. I-Ie also cml~hasized that under even-aged mnnagcnlcnt, harvesting of trees should be done on a long-rotation basis, 80 to 100 years. so that hardwood trees coulci reach the age of s~ihsttintial r-i~astproduction. Zeedylr recognizeti the problem of pro- viding mast in pine types anti suggested that patches of hardwoods be preservc:d or estahlishcd within such stotltls or be maintained as an adn~ixturethroughout the stands. Farrand 119631noted that ht~rvesting selected trees Crom a stand in Pennsylvania resulted in an increase of browse and improved species composition and variety of stand conditions. McKnight 119661 favored the long-rotation system. ;ippliecl within the structure of uneven-aged silvic~~lture. in southcrn Ilardwood forests. EIe emphasized that cut- tings should be made it1 patches or groups of trees for regeneration, with careful attention bcirlg given the condition of individual trees in the groups selected. He said that this variable selection system would result in a good hardwood forest and provide tile maximum range forage may be produced for a period of from 3 to 5 years of stand conditions in contiguous patches for wildlife in the short-rotation system of a 30-year cutting cycle. and recre:ttion. This system woulci also take advantage During the regeneration phase of the short-rotation of good, current growing stock. management ol pine, therct is usually a great surge in Summarizing a discussion of southern hardwood man- the growth of palatable browse, along with other secon- agement, Miller I 1957) said, "The principal requirements dary growth, provided a seeti or coppice source is present. of hardwood forest habitat for multiple wildlife species In a few years, however, tile pin(: crowtis out most of then are as follows: (ai a mature overstory providing the browse, and that wl~ichpersists is of poor quality mast and dens, tbi a thrifty understc~ry of palatable anti vigor Frirtl~erniore.tlie ticnsc 1,iirc precludes any broivse and desirable r~production,! c) forage and game significant production of lni;lst, cithel. liard or soft, in food plants in the ground cover, and (d) available water." future years. 1Inl.dwoocis sucl~cis oak and gums, even 1-Ie suggested that this could be achieved by making im- if their height growth Ictleps pace wit11 the pine, cannot provement cuts that would open the tree crowns to reach seed-proclucing age by the time of the next timber approximately 60 percent of closure. harvest, when the site is again prepared for planting. Schuster and Halls (1963) found that palatable deer Thus, for the long term, or over several rotations, it is browsc could be produced in adequate quantities (853 difficult to predict the e\lentual abundance and \velfare pounds per acre by selective cuttings every 5 years in of browse species. Each site preparation will likely a near-mature pine-hardwood forest in eastern Texas. destroy or reduce the regeneration potential of browse Forage production was closely related to the midstory species so that eventually the stand will be largely lim- crown cover. In this ~nstancethe niidstory had a covcr- ited to those species that were planted. Thus, in short age of 37 l~ercetitwhile the domirlant ovc,rstory was 32 rotation the variety of forage plants will be limited, and pcrcent. no h~rrdwoodmast will be present unless trees of seed- bearing age are retained in the stand at the beginning Stransky and Halls ! 1968 i. ;inlong others, have pointed out the nced for pc:rmancnt openings in pine-hardwood of the rotation. forest deer habitat. If short rotation must be used, wide spacing of planting Mixeci stands of conifers and hartiwoods proved to be stock and small cutting units within the home range of better habitat for trees as well as game in Russia deer should he better than close spacing and large cut- ting units. (Pogrcbniak 19621. Forests composed of several conifers, conifers-hardwoods, or mixed harrlwoocis werc healthier and grew faster than unmixed stands. Mixed stands also Long Rotation created more lime, nitrogen, phosphorus, potassium, and Long-rotation management usuaIiy involves some form magnesium in the surface soil. Furthermore. the soils of selective harvesting. The proctuction of palatable for- contained greater populations of microflora and meso- age and mast for cleer cfepends upon tree harvesting pro- fa~uia.It is nott>worthy that Pogrtbniak recorded simi- cedures and other manipulatioil techniques such as pre- lar results by research foresters in Germany, Czecho- scribed burning. From available data it appears that light slovakia, Sweden, and Poland. selcctivc~removal of trctes will not produce the needed Perina I 1962) ccrlled attention to the fact that the lack browse, tree regeneration, or mixture of vegetative spe- of suitable food in the all-coniferoi~sforests in Czecho- cies and typc. Light tree removal does appear to favor slovakia caused big game, presumably red deer (Ccrvus mast production in overstory trees but may keep yields cluphitsj, to suffer severe malnutrition, so that they of understory mast at a low level in old forests. Instead damaged the forests. Ile said that efforts h;ltl I)een ~~nder- of light removal, the current trend is to make clearcuts way there for many years to reconvert the pure stands in strips or patches. Such cuts should be comparatively to mixed stands by planting beech (Fugus sylvc~ticuL. I, small and not cover more than a limited part of the area. linden iTilin ~?~LTO~(L(VLL. 1, and carpinus i('cLrpinus betu- In making clearcuts it seems wise to retain some of the 1it.s L. ). best mast-bc'tr~ng treci becauic of the long periods ot I'erina crlso said that timber yield hati greatly declined time rey~~iredfor some specit~s,especially oaks, to reach in pure st;rnds of Norway spruce iPicc~ct rsrlsu L. ) and the age of nlaximum seed l~roduction.The long-rotation Scotch ~sirle iPinl~s.s?jli?estri.s I,.) in Czeel~oslovakia. system lends itself to thinning operations, which give This reduction was attributed to reduccd soil fertility. greater flexibility in maintaining a stand str~~cturesuit- insect infestation and disease, anti decreased water re- able to deer and other w~ltll~feShort cutt~ngcycles tention capacity of the soils. It is interesti~lgto note that ~Schusterand IIalls 1963) in p~ne-ii,ircIwoocl forests will this condition, according to per in:^, has crcatcd a very ma~ntainrclttt~vcly good forage procluctlon as well '15 unsatisfactory water flow dowilstream in other central good m,ist y~eldsthroughout the long rotation European countries. A nlajority of central European rivers rise in the country of Czecl~oslovnkia. As of Varietv and Cover 1960 only 11 percent of tllc iorc5ts in C"rc~cho>lovakia were fully ~ctlvehydroloqic,illy One of the prtnc lp,il ot)ject~v~sof for(l\t wifdl~fehabl- t~tni,~n,iqc>mc'tit 15 to rn,llnt,t~n ,I v'ir~ety of vcgetat~on RESULTS AND DISCUSSION in the h:ibit;it anti I bc~lievethis t~riticiplc,in general, is intiisputablc. Short Rotation F:xr)c~ric~ncctin central Euril~e;111<1 Russi;~has shown From the work that 11c1s bcvn done, it sccnis clt;ir that i-tiist~rlforcxsts result in hc>ttor growtll ;111d health that adcctuate amounts of pa1;it;rble browse and otlicr of tii(~trc3c.s ;1i1t1 thns :I srll)c~~.ic~rh;ibit;it for game. For more than a century foresters in these countries have Farrand, E. P. been reconverting all-conifer forests to a mixture of 1962. Deer and forestry. In Deer management in hardwoods and conifers to achieve a variety of species Pennsylvania, Sect. 2, pp. 5-7. Pcnn. State Univ. and mixtures. and Penn. Game and Fish Comm. Deer will eat a great variety of forage food if it is Goodrum, P. D., and Reid, V. H. available. However, among the many species of woody 1962. Browsing habits of white-tailed deer in the browse a comparatively small number is palatable to or western Gulf region. First Nat. White-Tailed preferred by deer. The average habitat contains only Deer Dis. Sym. Proc. 1962: 9-14. 15 to 20 percent first-choice browse species, while 35 to 45 percent is medium-choice and the remainder low- Krefting, L. W. choice or starvation food (Lay 1967a; Goodrum and Reid 1962. Use of silvicultural techniques for improving 1962; Lay 1967b). This led Lay (1967b) to conclude deer habitat in the Lake States. J. Forest. 60: that the quality of the forage is more important than 40-42. the quantity. He summed up the importance of variety Lay, D. W. by saying, "Forestry practices, which favor variety and 1967a. Deer range appraisal in eastern Texas. J. Wild- not just browse, may be expected to increase deer life Manage. 31: 426-432. carrying capacity . . . . The forest with the maximum mixture of stand types, ages, species, and treatments, Lay, D. W. together with well distributed clearings, will produce 1967'0. The importance of variety to southern deer more food for deer than even-age pine stands in large Southeast. Assoc. Game and Fish Comm. 18th blocks." Annu. Conf. Proc. 1964: 57-62. Small clearcuts within a dense forest are as desirable McKnight, J. S. as variety in the vegetation because they provide open 1966. Application of uneven-aged silviculture to or semiopen fawning areas for deer as well as a place southern hardwood forests. In Proc., Symp. on to loaf and play. Such areas also "green up" a little Hardwoods of Piedmont and Coastal Plain, pp. earlier in the spring, thus providing green food in the 61-64. Ga. Forest Res. Counc., Macon. form of forbs and young grass before browse species begin growth. Left alone, however, these open areas will Miller, H. A. be short-lived because of quick recuperation of the forest 1957. Capabilities and problems in the management vegetation. For proper management, some of them of hardwood forests for timber and game. In should be kept permanently free of trees and heavy Special problems in southern forest manage- brush. ment, pp. 91-99. La. State Univ. Sixth Annu. Forest Symp. Proc. 1957. Sch. Forest. and CONCLUSION Gen. Ext. Div. It seems evident that forest manipulation through Perina, V. long rotation management will produce a better habitat 1962. Changes of forests in Czechoslovakia. Fifth for deer, as well as for other wildlife, than short-rotation World Forest. Congr. Proc. 1960: 476-478. management. The advantage comes from greater flexi- Pogrebniak, P. S. bility in the use of management techniques for producing 1962. Creation of mixed stands as a method for raising adequate palatable forage over a long and continuing the productivity of forests. Fifth World Forest. period, maintaining mast yields, and preserving a variety Congr. Proc. 1960: 483-486. of plant species and stand conditions as well as openings in the forest. In short-rotation management browse may Schuster, J. L., and Halls, L. K be produced in great abundance for short periods of time. 1963. Timber overstory determines deer forage in shortleaf-loblolly pine-hardwood forests. Soc. LITERATURE CITED Amer. Forest. Proc. 1962: 165-167. Blair, R. M. Stransky, J. J., and Halls, L. K. 1960. Deer forage increased by thinnings in a Louisi- 1968. Woodland management trends that affect game ana loblolly pine plantation. J. Wildlife Man- in coastal plain forest types. Southeast. Assoc. age. 24: 401-405. Game and Fish Comm. 21st Annu. Conf. Proc. 1967 : 104-108. Bryan, P. H. 1950. Deer range improvement by modified timber Zeedyk, W. D. harvesting. Pap. presented at Southeast. Assoc. 1968. Wildlife under even-aged management. Pap. Game and Fish Comm. Fourth Annu. Conf. 12 presented at Ky.-Tenn. Sect. Soc. Amer. Forest PP. 25th Annu. Meeting. 16 pp. Timber Stand Density influences Food and Cover

Robert M. Blair ' Southern Forest Experiment Station Forest Service, U S Department of Agr~cuiture Nacogdoches, Texas

Forclge and fruit production for deer on forecst OVERSTORY DENSITY AND FOOD PRODUCTION lands is generally related in7)ersely to timber stand Forage Yield density. Growing timber nt (I minimurn density that will yield an acceptable rconomic return can in- Production of grasses, forbs, and browse is generally crease light penetration to the t~nderstory,thus im- inversely related to the overstory density. The relation- proving the enziironment for growth of food and ship is often expressed as a curvilinear function where cover vegetc~lioiz. increasing stand density in square feet of basal area or percent of canopy cover restricts the weight of forage Within a forest community, growth of food and cover produced (Halls et al. 1956; Ehrenreich and Crosby 1960; plants for deer is primarily a function of timber stand Halls and Schuster 1965; Blair 1967). density. As trees develop from regeneration to harvest, Ilerbage production in the Soutll ranges as high as they and the plants beneath compete for light, moisture, 3,000 pounds or more of dry matter per acre on cutover mineral nutrients, and growing space. Availability of forest lands and among newly established pines (Duvall these factors and the physiological tolerance levels of and Hilmon 1965). Beneath a forest canopy yields de- understory plants determine the ecological development cline as stand density increases (Rhodes 1952; Gaines of the forest. et al. 1954; Ehrenreich 1960; Schuster 1967). Production This paper emphasizes food production. It is more may average less than 100 pounds of dry matter per acre critical for deer than cover, which seldom is a limiting in fully stocked plantations of pole-size pines or natural factor. And the same plants that furnish food generally pine-hardwood stands with a basal area of 100 square provide adequate cover. feet or more per acre (Rhodes 1952; Blair 1967). Browse plants respond to stand density in much the LIGHT AVAILABILITY AND PLANT GROWTH same way as herbaceous plants. Dry matter production Of the environmental factors influencing composition, of 1,440 pounds per acre has been reported in forest growth, and vigor of plants, light is the one most easily openings in Florida i Strode and Chamberlain 1959). modified and controlled by stand manipulation. It may Beneath a tree canopy woody forage declines with in- become the dominant factor if the canopy is dense enough creases in stand density. In east Texas, browse in a to reduce light intensity below 20 percent of full sun. mixed pine-hardwood stand increased from 300 pounds Be~leattluneven-aged hardwood stands in the Central of ovendry materlal per acre where the stand dcnslty States, light intensity varied by stand density (Minckler was 96 square feet of basal area to over 700 pounds on 1961). In stands of about 40 square feet of basal area an area with 26 square feet (Schuster 1967) per acre, light transmission at midclay varied between Beneath a canopy the current growth of browse gen- 25 to 30 percent of full sun. The trees, pole size or larger, erally exceeds that of herbaceous plants, especially were in full leaf. As basal area increased to 70 square where fire has been excluded or is seldom used (Rhodes feet, the intensity declined to about 17 to 22 percent, 1952; Halls rt al. 1956; Blair 1967). and to around 9 to 13 percent at 100 square feet. Timber In most southern pine-hardwood forests the understory size had no apparent effect. growth 1s strongly ~nfluencedby a hardwood midstory Within openings, average light transmission was re- beneath the dominant pine canopy. As the stands develop lated to the size of the openings. Even those as small as much of the woody understory grows beyond reach of 18 to 30 feet in diameter received two to three times more the deer and forms a dense, multilayered midstory of light than beneath a hardwood canopy (Minckler 19611. young hardwoods and large shrubs (Schuster and Halls Pine stands generally let in more light than hardwood 1962; Blair 1967 1 . The combined influence of overstory stands of comparable age, basal area, and crown closure pine and midstory hardwoods governed browse produc- (Lull and Reigner 1967; Schomaker 19fi8). But even in tivity in a 30-year-old loblolly pine (Pinus laedu L.) young, dense conifer stands, light transrnissioti may be plantation that had been thinned at 20 and 25 years of less than 5 percent of full sun--under which condition age (Blair 1967 1. The rnultilayered midstory grew pro- understory vegetation tends to disappear (Shirley 19451. gressively denser and when crop trees were 35 years old it pritnarily determined browse growth. The higher and Because many species of forage- and fruit-producing less dense pine canopy exerted little inlluence. plants can grow and reproduce at low light intensities, Effects of dens~ty'Ire lessened in older plne stands they can be managed as a component of the forest under- wliere a micistory is sparse or absent. Although basal area may be greater, trees are fewer and taller. Consid- of the Wildlifc II;\bitat and Silvi- erably more light rc:iches forage plants than in younger h is maintainc'd at Nacogdochrs, Texas, by the Southern Fortxst Expcrimcxnt Station in cooper- stands with low, dense canopies or in older stands with ation with Stephen F. Austin State Univcrsily. a dense midstory. Heavy litter accumulation is often associated with I11 central Louisiana, Hastings 11966) found that fruit reduced plant growth, and leaf cast is directly related yields ciecIi~redslowly, but progressively, as the basal to basal area density of the overstory (Gaines el al. area in a pine-hardwood stand increased. Yields were 1954; Blair 1960 ). As pine needles and llardwood leaves avtrage or better in stands of 60 to 80 square feet of accumulate, many seeds iiever reach bare soil to ger- basal area but dropped sharply at higher ovcrstory minate, and young seedlings are often smothered. Mod- densities. erate litter accumulations, on the other hancl, are bene- He also reported that stand structure, c;rriopy form, ficial as they i~~sulatethe soil and minimize temperature and composition appreciably zrffected the fruiting of extremes and moisture losses. shrubs. Yields were highest and most consistent in forest openings. Below canopies yields were affected Floristic Composition least in stands with a pine overstory and most beneath a pine canopy with a multilayered midstory of hard- Floristic composition in the uriderstory is also a func- woods. tion of stand density. As light is reduced shade-tolerant plants predominate. Many browse species palatable to deer, such as .Japanese honeysuckle (Loizicerc~jc~ponica IMPROVING FOOD AND COVER ENVIRONMENT Thutrb. I, yellow jessamine fGelseir~i~lmnsenzperuirens Commercial forests are gener,~llymanaged to prov~de (I,. ) Ait. I, and arrowwood iVib~~rnz~mdenlaturn Id. ! are tolerant of shade, and are often more abundant under a favorable environmei~tfor Ire<>growth and regenera- a canopy than undesirable species as waxmyrtle iMyrica tion They can alro be marlaged for production of deer- food and cover cerifera L. 1 and persimmon (Diospyros virginznntr. L. ) (Schuster and Halls 1963; Blair 1967 1. Cultural practices that manipulate ovcrstory density Development of the overstory may cause undesirable and minimize midstory formation are important it1 pro- changes in both the growth and composition of herhacc- viding adequate light to the unclerstory. Reclucitig stand ous vegetation. As stand density increases, desirable density may also reduce competition for miner;ils and grasses often give way to less desirable species (Martin moisture among forage- and fruit-producing plants, as el ul. 1955; Halls and Schuster 1965). Growth of most well as among crop trees. Removal of excess litter would composites, legumes, and other forbs declines as light Irelp provide a seedbed conducive to the establishment decreases 1 Blair 1960; Ehrenreich 1960; Schuster and of understory plants. Halls 1963 1. Reduced light intensity may have greater To achieve the desired benefits for both timber and influence in decreasing the number of forbs ant1 their deer, management plarls must be based 01.1 sound ecologi- vigor than in determining the composition. cal principles. Factors such as the species of food plants desired and their growth requirements, age and structure Mast and Fruit Yields of the titnber stand, and type of forest products being grown should be carefully evaluated. Timber density sigtrificalitly affects production of fruit and mast, which are importaiit for dcer in seasons when Adequate food and cover can grnerally bc produced other sources of energy are scarce. beneath the dominant cai~opyif there 1s enough light Acorn y~eldsvary according to timber deiisity and Crop trees thus must be grown at constdcrably less than species (Goodrum and Reld 1956, Shxp 1958) South- maxlmum dens~tyand the inidstory must be I~initcdto ern oaks with small crowns appear to tolerate a denier treei and large shrubs that bcdr fruit and tnast Timber- I,md owners must be w1111ng to matiage their stands at overstory than large-crowned ipccles, yet yield much a mimmum density t11~1t will yield an acceistable eco- mast Sound, uncrowded trees growlng 111 a tlominant canopy position or In openlngs gener'tlly are the best nomic return arid will also prov~dea favorable envlron- meilt for growth of pcilatable deeriood The nutrient proclucers of acorns I Sharp 1958) Average yields per qu,llity of the food must al\o be adequate tree for turkey oak IQz~erct~~laevls W,~lt) in Florida were h~glrestin atands w~th15 to 80 stems per acre, but y~eldsper acre were greatcit whtxrc there were 150 to 160 LITERATURE CITED acorn-bearing trees ( Harlo~.iiand Eikum l9fj5 ) C),ik> growlng In deep shadc are gcner'11ly poor prorfucers Bl;tir, R. M. 1960. Deer forage increased by tliinnings in a Louisi- Amount 2nd freque~reyof fruit l,rocluctio~~by under- ana loblolly pine plairtation. J. Wildlife Man- story shrubs appears invc,rsely related to timber stand age. 24: 401-405. density. Fruit yields of several browse species in east Texas, such as yaupon il1e.r ~:ort~itorinAit. I. A~nericarr 13lair, It M beautyberry fCnllicc~rptri~t~~eric,arrc~ L. 1, and floweritig 1967 Dee-r forage ill a ioblolly p111c p1,intatlon .J dogwood (Co~r~ttsfloricla I,, i, were supprc.ssec1 more by Wtldlife Matiagc, 31 432-4317 the overstory cover than forage yielcis Croln tlie same I>uv;~li.V. I>.. and Hilrnon, J. B. species I H;ills and Alcatiiz 1968 ). Yields of fruit from 1965. New grazirig research programs for southern r~llaritsgrowing in tlie opc,ri were several tirries jiri?;ittlr forest ranges. J. Range Mallage. 18: 132-136. than Crrjm those herrcatli an overstory of 70 square fcet of basal arcs per acre. The hiirdwood tnirlstory, charat:- Ehrenrc~ch,J H tcristic of these st;tntls, had been olimit~;it.ecl. Presence 1960 li'sdble forage utltler pitic \tands9 USDA Furcst of this midstory wctuld have f'urtlier restricted light trans- Serv Cent Statci Foiest Exp Sta , Sta Note mission and yields. 142, 2 pp Ehrenreich, J. H., and Crosby, J. S. Martin, S. C., Dunkeson, R. L., and Baskett, T. S. 1960. Herbage production is related to hardwood 1955. Timber harvests kelp offset forage decline in crown cover. J. Forest. 58: 564-565. Missouri's managed forests. J. Forest. 53: 513- 516. Gaines, E. M., Campbell, R. S., and Brasington, J. J. 1954. Forage production on loligleaf pine lands of Minckler, L. S. southern Alabama. Ecology 35: 5:)-62. 1961. Measuring light in uneven-aged hardwood siands. USDA Forest Serv. Cent. States Forest Goodrum, P. I)., and Reid, V. I<. Exp. Sta. l'ech. Pap. 184, 9 pp. 1956. Wildlife irriplications of hardwood and brush controls. Twenty-First N. Amer. Wildlife Coni. Rhodcs, R. R. Trans. 1956: 127-141. 1952. Timber and forage production in a pine-hard- Halls, I,. K., and Alcaniz, R. wood stand in Texas. J. Forest. 50: 456-459. 1968. Browse plants yield best in forest openings. Schomaker, C. E. J. Wildlife Manage. 32: 185-186. 1968. Solar radiation measurements under a spruce I-ialls, L. I<., Hal?, 0.M.. and Southwell. B. L. and a birch canopy during May and June. Forest 1956. Grazing capacity of wiregrass-pine ranges of Sci. 14: 31-38. Georgia. Ga. Agr. Exp. Sta, Tc:ch. Bull. in.s.1 2, Schuster, J. L. 38 pp. 1967. The relation of understory vegetation to cutting Halls, L. K., and Scliuster, J. L. treatmerits and habitat factors in an cast Texas 1965. Tree-hcrbagc relations in pine-l~ardwoodforests pine-hardwood type. Southwest. Natur. 12: of Texas. J. Forest. 63: 282-283. 339-364. flarlow, R. F., and Eikum, R. L. Schuster, J. L., and I-Ialls, L. K. 1965. The effect of stand density on the acorn pro- 1963. Timber overstory determines deer forage in duction of turkey oaks. Soutlieast. Assoc. Game shortleaf-loblolly pine-hardwood forests. Soc. and Fish Comm. 17th Annu. Conf. Proc. 1963: Amer. Forest. Proc. 1962: 165-167. 126.133. Sharp, W. M. I'lastings, E. F. 1958. Evaluating mast yields in the oaks. Penn. Agr. 1966. Yield and chemical an~tlysesof fruit produced Exp. Sta. Bull. 635, 22 pp. by selected cleer-browse plants in a loblolly- shortleaf pine-liardwood forest. Ph.D. Thesis. Shirley, H. L. La. State Univ. Baton Rouge. 247 pp. 1945. Light as an ecological factor and its measure- ment. 11. Bot. Rev. 11: 497-532. Lull, H. W., and Reigner, I. C. 1967. Radiation measurements by various instru- Strode, D. D., and Chamberlain, E. B., Jr. ments in the open ancl in the forest. USDA 1959. Wildlife habitat management in Florida Na- Forest Serv. Res. Pap. NE-84, 21 pp. Northeast. tional forests. So~~theast.Assoc. Game and Fish Forest Exp. Sta., Upper Darby, Penn. Comm. 13th Annu. Conf. Proc. 1959: 38-50. Effects of Hardwood Control on Food and Cover

E. B. Chamberlain, Jr. Bureau of Sport Fisheries and Wildlife U. S. Department of the Interior Atlanta, Georgia

The importance of hardwoods in providing food pounds for the four types combined. On the Jefferson and cover for white-tailed deer in the southern forest National Forest, Virginia, browse production averaged habitat is documented by pertinent literature. Hard- 37 pounds per acre, ovendry (Stiteler and Schrauder wood control is, therefore, a matter of much concern 1967). Forage production by trees in north Arkansas in deer management. The most common types are ranged from 254 to 1,525 pounds per acre, green weight burning, chemical, and mechanical. In one or an- (Halls and Crawford 1960). In Florida, Harlow (1959) other form, the practice is well established. There found browse production running from 52 to almost 900 are direct relationships between hardwood controls pounds per acre, airdry. (and timber management practices generally) and The importance of hardwoods in providing cover for production of deerfoods. Effects depend upon inten- white-tailed deer in the southern forest habitat is not sity of treatment and on size and distribution of so well documented as is the case in food production. A treatment blocks. report by Davis and Winkler (1968) on brush versus The importance of hardwoods as food for white-tailed cleared range as deer habitat in south Texas shows that deer in the southern forest habitat is well established. while deer used cleared areas extensively, especially at The works of Pearson (1943), Korschgen (1954), Lay night, they depended upon the hardwood brush for cover. (1957, 1961,1967), Segelquist and Green (1968), Halls and A report by Krull (1961) on a study of deer use of clear- cuttings in northern New York indicated that during Crawford (1960), Stiteler and Schrauder (1967), Stiteler and Shaw (1966), Collins (1961), Harlow (1959), Harlow severe winter weather the greatest deer use was in uncut areas. Main timber types were hardwoods and hard- and Tyson ( 1959), Reid and Goodrum ( 1958), Goodrum (1959), and Strode and Chamberlain (19591, among others, wood-conifers. In an area of longleaf pine-turkey oak in show the great importance of acorns and other fruit, as Florida, it was found that clearing all woody vegetation from plots 1 mile square caused a pronounced reduction well as browse, to this species. in deer use. Removal of one-half to three-fourths of the Acorns are heavily sought after by deer and comprise native cover resulted in increased use by deer, again a very important part of the diet, even though production indicating the need for cover (Beckwith 1967). In gen- varies widely from tree to tree, year to year, and species eral, hardwoods do provide the cover required by deer to species. Individual trees may produce as much as in the South, though probably not to the extent that 45 pounds of sound acorns some years though averaging they provide the food required. It is safe to say, at any 3 to 18 pounds (Collins 1961). Average annual expected rate, that in the southern forest habitat both food and yield of sound acorns per foot of basal area has been cover required by white-tailed deer are furnished by found to be 4.90 pounds for post oak. 6.59 pounds for hardwoods of various species. white oak, 5.90 pounds for blackjack. 1.98 pounds for Hardwood control is, therefore, a matter of much con- southern red oak, and 8.19 pounds for water oak (Good- cern in deer management. In one form or another, it is rum 1959). an established part cf land management practices in Aggregate production of fruit by small trees, shrubs, the region. The most common types are burning, chemi- and vines of the understory may exceed acorn production cal, and mechanical (Walker 1956). Burning has long per square foot of basal area (Lay 1961). Production of been used for control of hardwoods, and has been well 20 to 65 pounds of fruit per square foot of basal area was reported by Lotti, et al. ( 1960 ), among others. The next found for five species in east Texas-dogwood, fringe- speaker on this panel will discuss this subject. tree, blueberry hawthorn, flatwoods plum, and sweetleaf. Chemical control using any of many available herbi- Utilization of these foods is shown by Lay's report cides may be carried out by use of sprayers, injectors, (1965) of the examination of the woody seed content of or airplanes. Aerial-spraying and mist-blowing to remove 2,295 deer pellet groups collected over a 7-year period undesirable hardwood competition from pine sites are in east Texas. Thirty-one species or genera were identi- in general use throughout the southeast by many wood- fied, the most frequent being oaks, yaupon, American land organizations. In 1960, approximately one-fourth beautyberry, blackgum, and hawthorn. Some species of of the total area treated chemically for hardwood control fruit were found during every month, though availability was by aerial and mist-blower application (Chamberlain and occurrence were highest in fall and winter. Acorns and Goodrich 1962). This practice developed over a 5- were present in 14 percent of all pellets examined and year period, for prior to 1955, only a few small experi- in all 49 stomachs examined in November and December mental plots had been treated with selective herbicides of a good acorn year. by aerial application. In 1955. Hiwassee Land Company Average browse production on four forest types in conducted the first, large-scale aerial application of the Ozarks varied from 76 to 131 pounds per acre, oven- 2,4,5-T in the southeast. In July of that year it sprayed dry (Segelquist and Green 1968) with an average of 101 2,000 acres of low value, inferior quality hardwood stands on the Cumberland Plateau in east Tennessee. In the showed some lag in mortality of undesirable species but years following, additional work has been conducted by also generally poor persistence of sprouting and relative- Hiwassee Land Company and a number of other indus- ly quick recovery from crown dieback in the desirable trial and private timberland owners in the southeast. species. The most desirable species was killed by all Helicopters are more commonly used for this purpose treatments. than are fixed-wing aircraft, due to their greater maneu- Mechanical methods of hardwood control include gird- verability. The helicopter passes back and forth over ling, cutting, and use of heavy eq.uipment with chains, the area being treated, in adjacent flight lines 45 feet brush cutters, and various blades. A study of resprouting wide, and at a height of 25 to 50 feet above the timber. from cut stumps by turkey and hluejack oaks in north- Rates of application vary from 1 to 2 pounds of 2,4,5-T west Florida indicated that at least two additional treat- acid per acre in a total volume of 3 to 5 gallons of solu- ments with heavy brush cutters 6 to 10 weeks apart tion per acre. A typical formulation consists of gallon during the main growing season were necessary to obtain of ester (containing 2 pounds of 2,4,5-Ti, M gallon of good kills (Woods and Cassady 1961). In another study diesel oil, and 4 gallons of water. The work is normally in pine-hardwoods stands in southern Arkansas, dense conducted from the latter part of May through early hardwood thickets were bulldozed clean. First-year July. sprouts were few, but after 5 years, one-fourth of the Mist-blower application of selective herbicides to weed area was overtopped by sprouts. After 7 years, half the forest stands was first used in the South in 1957 by S. F. area was overtopped by sprouts averaging 6 feet tall Potts of Crawford, Mississippi, who developed a light- (Grano 1961) A similar study in bottom-land hardwoods weight compact blower for mounting on the back of in west Mississippi showed that in 5 years sprouting a small crawler tractor. The technique involved with produced about 7,500 stems per acre, 1% inches d.b.h. th- mist-blower is to move across the area to be weeded and 15 feet tall (Johnson 1961). in adjacent lanes 20 to 40 feet in width. The herbicide, The direct relationships between timber management rate of application, and formulation are the same as in practices and the growth and production of deerfood aerial spraying. plants have been demonstrated by many studies. Sehuster Aerial-spraying may be used to remove either an over- and Halls (1963) showed that forage production was al- story or an understory, while the mist-blower is designed most four times greater in a clearcut and two times great- to weed out the smaller, understory vegetation. In either er in a selection cut than in an uncut control area. Thin- case the purpose of the weeding treatment is to remove ning of a loblolly pine plantation in central Louisiana at a sufficient number of hardwood stems to allow estab- age 20 and again at age 25 to approximately 100, 85, and lishment and release of the pine seedlings that will form 70 square feet of pine per acre resulted in yields of deer the new stand. browse ranging from 90 pounds per acre under light thinning to 137 pounds under heavy thinning (Blair Aerial application of herbicide is widely used to control 1960). Halls and Alcaniz (1968) reported that at age 5 oak sprouting in mechanically-cleared site preparation years open-grown plants of seven browse species aver- areas. In such cases, oak control may be 85 to 90 percent aged 32 times more fruit and nearly seven times more effective and control of other vegetation 75 percent effec- twig growth than plants beneath a sawtimber size stand tive. of pines. In mixed oak-pine forests of western Virginia Aerial application of herbicide is frequently used to improvement cuts that removed 30 to 80 percent of the convert a commercially worthless type, such as swamp basal area increased first-year browse production by titi, to pine. With titi, approximately 95 percent of the three times for the lightest cut and 15 times for the plants may be killed back to the ground, so that subse- heaviest. Four years after cutting, the increase was five quent seeding of pine can be done successfully. However, times the control for the lightest cut and 24 times the it appears that 50 to 60 percent of the plants may re- control for the heaviest (Patton and McGinnes 1964). sprout at the base. Studies of acorn production in relation to thinning On general pine-hardwood sites, aerial and mist blower are less numerous. A report by Harlow and Eikum (1965) applications are widely used to achieve silvicultural showed that heavy thinning (50 to 90 percent of stand weeding. Helicopter or fixed-wing aircraft application removed) in a stand of turkey oak increased the yield can be expected to kill 70 percent of the overstory (ex- per tree but the control, which had 150 to 160 mature clusive of pines), and mist blower applications 65 to 70 trees per acre, produced five times more acorns per acre. percent of the understory vegetation. Where a good burn Where mast-producing trees are to be removed selective- is accomplished in conjunction with the herbicide treat- ly, care should be taken to retain the individual heavy ment, understory reduction will be much more complete. producers This kill will occur over a period of 2 to 3 years. Total The recent trend toward clearcutting, which is one kill is not obtained because of skips in application, ap- form of hardwood control, has many implications in parent inherent resistance of some species and individ- deer management. Ripley and Campbell (1960) reported uals within a species and occasional unfavorable soil on two tracts of typical mountain hardwoods in western moisture conditions. Understory vegetation can be ex- North Carolina which were clear and selectively cut in pected to increase considerably after the second year, 1949. Two years after cutting there were approximately so that it may become much more dense than originally. twice as many seedlings and sprouts on the clearcut as Grass and herbaceous growth may likewise increase. on the selectively cut area. Browse utilization appeared A 6-year evaluation of herbicide treatments to improve to be lower on the clearcut area. Ten years after treat- deer browse in northern Idaho (Lyon and Mueggler 1968) ment the clearcut area had three times as many commcr- cially valuable seedlings and sprouts as did the selcctive- Chamberlain, E. B. Jr., and Goodrich, T. K. ly cut area. Browsing was moderately heavy in the 1962. Acrial and mist-blower application of herbicides clearcut area and severe in the selective cut. More in southern forests as relatcd to wilcilife man- browse was produced than could be utilized in the :~gen~ent.Twenty-Seventh N. Amer. Wildlife clearcut area, permitting satisfactory rc~gctleration. and Natur. Resources Conf. 'l'r;ltis. 1962: 384- Continued work on these same areas was reported by 3113. Della-Bianca and Johnson 119651. When thc regenerater1 Collins, .J. 0. stands reached sapling stage, browse was out of reach I!)fjl. Ten year acor~~mast production stucty in 1,ou- of deer and crop tree growth greatly reduced. To ovcr- isiana. La. Wild 1,ifc anti Fish. Comm. 73 pi,. come this, removal of all woody stems except desirable r) a~i,:. R. B., and \Vi~lklcr,C. K. crop trees was tested on part of the area. Dense coppice growth resulted, and deer use was heavy. Browse pro- 1968. Brush vs. c1earc.d range :is decr habitat in duction in the cleaned areas varied from 805 pounds southern Texas. J. Wildlife M;in;ige. 32: 321- per acre on lower slopes to 81 pounds on upper slopes. 329. Untre~iterl compartments, both lower and upper, had Dell;i-I3in1-ica, Id., ~inriJohnson. F. ;LI. 3 pounds of browse per acre. 1965. Effect of an intensive cletitii~~gon (leer-browscx In southern Missouri, production of understory vege- protiuction in the Southern Appnlaci-iiar1s. J. tation on ur-idisturbed areas was less than on logging L5'ildlifc Manage. 29: 7211-733. and TSI areas during a 10-year period (Murphy and (;oodrum, P. D. Ehrenrcich 1965). However, increases were not as great 1959. Acorns in the diet of wildlife. Southeast. Assoc. as expected. This was probably due to the fact that the Game and Fish Comm. 13th Ar-inr~.Conf. Proc. type of timber management in eflect during this time 1959: 54-61. created small openings with minor reductions in basal Grano, C. X. area and crown cover. 1961. I-Iardwood reoccupation of bulldozeti sites. In The many studies which have been cited as having Hardwood sprout development on cleared sites, some bearing on effects of hardwood control have pro- pp. 7-8. USDA Forest Serv. South. Forest Exp. duced a multitude of data. As must be expected in so Sla. Occas. Pag. 186. complex a problem, interpretation is difficult and sum- mation is perilous. Whether hardwood control is gooti IIalis. L. I<., and Alc;rt~iz,R. or bad for dcer depends on intensity of the treatment, 1963. Browse plants yield best in forest openings. J and on size and distribution of trerrtmet-it blocks. Where Wildlife Manage. 32: 185-186. section-sized tracts are completely clcareci, deer habitat Halls, I,. K.. and C'rawforcl. I-I. S., Jr. is lost. If clearcutting is done in 5 to 100 acre blocks, 1960. Deer-forcst habitat relationsl~ipsin north Ar- properly distributed and with necessary follow-up treat- kansas. J. Wildlife Manage. 24: 387-395. ments. browse production can be much improved. At Harlow, R. F. the same time, tl~eimportance of acorns and other mast 1959. An evaluntiorl of white-tailed deer hahitat in and fruit requires that hardwood control bc practiced Florida. Fla. Game anti Fish Cornm. Tech. 13ull. in a manner which will not significantly reduce their 5, 64 pp. production. Thus, hardwood control programs apparently must have the following characteristics to be most bcne- IIarlow, 1%.F., and Eikum, R. I,. ficial to deer: 1965. The effect of stand doisity on the acorn pro- duction of turkey o;iks. Southcast. Assoc. Game (1) Treatment blocks should be about 5 to 100 acres and Fish Comm. 17th Annu. Coilf. Proc. 1963: in size. 126-133. (2) Blocks should be st, distributed as to be within range of all deer in the area. 'farlow, R. F., arid Tyson, E. I,. (3) Follow-up treatments should be rnadc as necessary 1059. A ~~reliminaryreport on the effect of mast to maintain browse production. abuntlance (111 the weight and reproduction of decr in cet~tralFlorida. Southeast. Assoc. Game (4) Production of acorns axid fruit s11o~ildnot he re- al~dFish Comm. 13th Annu. Conf. I'roc. 1959: duced. 62-69. ,Johnson, IZ. L. LITERATURE CITED 1961. I-Iarclwood sprouts dominate bottom-land clear- Beckwith, S.L. ings. 111 H;lrdwoocl sprout developmelit on ll)(i7. Effect of site 13repar;ition on wiltllifo ant1 vc3ge- cleared sites, p. 8. USDA Forest Serv. South. tation in the sandhills of central Florida. South- Fortxst Esp. Sta. Occas. Pap. 186. east. Assoc. Game ant1 Fish Comm. 18th Anriu. Korschger-i, Id. J. Conf. Proc. 15164: 39-48. 1954. A study of the fooct habits of Missouri deer. 1Mo. Cotiserv. Comm. 43 pp. Blair, R. M. 1!)60. Deer forage iricreiised by tliinni~igsin a 1,ouisi- lCrill1, J. N. atla loblolly pine plantation. .I, Wiltllifr Mail- l!)(i4. Decr usc of a con~mercialclearcut area. N. Y. age. 24: 401-405. Fish and Game J. 11: 115-118. Lay, D. W. Reid, V. II., and Goodrum, P. D. 1957. Some nr~tritionproble~ns of deer in the southern 1958. The effect of hardwood removal on wildlife. pinc, type. Southeast. Assoc. Game and Fish Soc. A1nc.r. Forost. Proc. 1957: 141-147. (lomln. 10th AI~~LI.Conf. Proc. I!)T,fj: 53-58. Ril~ic,y,'r. I-I., a~~dCampbell, R, A. 1960. Hrowsing and stand regeneration in clear- and Lay. D. W. selcctivcly-cut h:irdu~oods. Twcnty-Fiff.11 N. 1961. Fruit production of some understory hardwoods. Arner. Wilcilile and Natur. Resources Conf. Southeast. Assoc. Gorne anel Fish Comrn. 15th Trans. lR(5O: 107- 415. AIII-IU.Conf. I'roc. 11461: 30-37. Schuster, J. I,., and I-Ialls, L. K. I,ay, D.W. 1963. Timber ovc,rstory dcter~nincstleer forage in lOfi3. Fruit utilization by tlecr in southcrn L'orc:sts. J. shortleaf-ioblolly pine-harciwood forests. Soc. Wiltilifc Manage. 251: 3'70-375. Amcr. Forest. Proc. 1962: 165-l(57.

I,;iy, D. W. Segelyi~ist,C. A., and Green, W. E. Deer food yields in four Ozark forest typc,s. J. lOlj7. The importance of variety to southcrn tieer. 1968, Southeast. Assoc. Game and Fish Comm. 18th Wilillife M;rnagc. 32: 330-337. Annu. Conf. I'roc. 1964: 57-62. Stileler. W. M. ,Jr.. and Schraucier, P. A. 1967. Protluction ant1 dcer utilization of woocly browse Lotti, T., Klawitter, R, 11..and I,eGr;~nde. W. P. on the Jei'forson National Forest, Virginia. 1960. Pl.escril2c.d burning for understory control in Soutl~etist.Assoc. C;artle alid Fish Cornm. 18111 loblolly pine stantis of the Coastal Plain. USDA Anliu. Conf. Proc. 11164: 21-30. Forest Serv. Southeast. Forest Exp. Stt~..Sta. Pap. 116, 19 pp. Stitelcr. W. M., Jr., ancl Shaw, S. P. l!)(j('. Use of woody browse by whitetail deer in I,yon, J. L., and Mueggler. W. F. heavily fo~,c~sted;rreas of Northeastern United 1968. I-It.rbicide treatment of North Idaho browse States. Thirty-First N. Amerc. Wildlife and evaluated six years later. J. Wiltilife Manage. Natur. Resources Conf. Trans. 1966: 205-212. 32: 539-541. Strode. D. D., and Chamberlain, E. B.. Jr. Wildlife habitat management in Floricia national Murphy, D. A,, anci Ehrtrireich, J. 13. 1951). forests. Southeast. Assoc. Game and Fish Ct)mm. 1965. Effects of timhcr harvest ant1 stanri improve- 13th Annu. Conf. Proc. 1959: 38-50. ment on forage production. J. Wildlife Manage. 29: 734-739. Walker. I,. C. 1956. Controlling untittsirable I~arciwoods. Gn. Forest Patton, D. R., and R;lc(;innt:s, H. S. Rc.s. Counc. Rcp. 3, 24 pp. 1964. Deer browse reltitive lo agc and intensity of Woods, F. W., anti Cassatly, J. T. timber harvest. J. Wildlife Manage. 28: 458-46.1. 1961. Sprouting of sandhills scrub oaks following Pearson, A. M. cutting. In IIartiwood sprout develol~menton 15143. White-tails like acorlls. Ala. Conscrv. 1516):8-9, clec~retisitcxs, pp. 1-6. USDA Forest Serv. South. 12. Forest Exp. Sta. Occzis. Pap. 186. The Effects of Prescribed Burning on Deerfood and Cover Paul A. SRuauder Wildlife Management Rrancli Sij~ithert?iiegion, Forest Scrvice U S Dcpciri~~ientof Agrici.iltiire Atl~ntn,Georgia Howard A. Miller Corisulting Forestcr arid Biologist Marietta, Georgia

l'l-c~sc~il~erl~IICI.~Z~II<~ JOT tl~e'l- 711 SOIC~~ZP~.~~2)i~lt' p~~luct.'Iistoric;illy, the lo~~glcnfr:[,;ist;il l'lait:s of the. jorc?srs is c~ izigi~ly rccoii~?~~c~tiderlpl-oqrci,!~, o/ 7ntrt~- soiiihc~ast \\-ere ti?c l~rovitig gni~iiitls for illis tyi~cof rrycrrrcvxt. Il,sc.tl ljy gc~~i~cbirtl r~~ciric~g(~l-.sjol- ?jc'cir,s I?I:II?:~~CI~C'II~.Stoddiird iinrl otli('r'i iisetl f~rc-in opc>r;ition- in f,/~t' S~iitlt,jil-(l ~(~12OC jl(sI (IS rc~lirc~l~l~~OI-tl~(7l- ;il m;inngc~ncnt as early ;is the tweiiiies. Fi1.1. rilltier ~nr~?lr~gccirzc~nIi?r ~0z~t/1(~l-11 pi)l(' jol-i'sts. 11 i,s (111 ('i.('1~1- coiitrollrrl conditions c;in Ile just ;is v;ili~tibic in tlec~r l<>i~llirrzl~cr rr~ci?~rcgc~rncnt n~c?tliotl. Not or11y c>c.onorrt- ~nnrriigcmcntin southt.rn pirie forrsts. ic.c~ll?j,it pl-oi;iilcs ~rrorc~zi~lritiozis l)i.o?~s(~ slij~~lic's I11 gc,~icrnl, lprescrihed huri;iiig iii II;II-tln.ooiis is tletri- (llili 11ft~'rfl?ll>llf 2 :jl?(li..7, Cjl~?ll?~l~~/~j~l~Ot~li('(~.~ irlOr(> rnc.i-it:il Exccpt for rc~ge~ier;itio~i.fir(, shoulii hc iisetl food t 12(171 cln ztnh~trnerli~r(,11. 111 pine st(c~~tl,s,r('~(a(ri-(.J~ very sl);irii~glyin l?ard\~roods.Strc,arn holie,nls silpl)ortiilg is still r~c.edcd on .seci.so~~crlfirc..s itndcr i~crrioirssiic.. mast-prodiicing hardwoods sl~ould hc scp;irated from ~TOIL?IL~c.ol:er, nnrl spccies coirt1ifion.s. 112 gene]-(11. atlj:~ccnt pine stands by ploweti lin-s, Clr'bi-caks, or 1)rexcribed 111~r?1!ingin i1nrd7uootls is detri11~cr11(~1. hackfires. When mast-!)rodtieing ici:irtl\~-,voocis:il.c n n?a,ior About (L 4-ypnr winter prc'scribe brirn rotnlio~zis component of upland forests, prescribed fircs sliorrlil not the most beneficial lliirn for botiz ~inrlcrstory~.c'g~t(~- hc used until thcy ;ire of young pole stand size I Sir;irisky lion growth clnd ntr~stprotlziclion. Pnlrrtcrble sprcic..~ anrl IIails 19fi7i. Rcpcrctcd bur~iil~g,even tiiough ligllt, of flesk?~Sqtngi orclcr crbii~zrlantly jol1ou;in~g firp. will everitually remove hardwoods from pine-hartiwoorl High intensity fire proclrices lligizer signijicc~nf in- stands. Unpublished fiildirigs from the ririgc and v;illcy creases of protein and pliospltoric c~citiJor (L longer l~rovincc,l3ii1e-iiarclwooiiw~ocl lirnbcr type in Virginia, incli- dlrrafioi~than lolo intcnsifg fires. c;~tercsprouting success with hear oak /QI~(>Yc?Lsillici- Spring burns, range r/lt(rlif?jcoizsidrred, crre 1)effer joli(~]in 1964 using l~rescribeilburn i Shrauder 1004 I. than full or winter b~crns,zuitlz szcnrmer fjz~i-~~ing 1ic;ir oak is :i11 excellcrit pcreiini;il uncii.rstory mast 121.0- prohnbly l~eingeqliall?~ good. Ann~cc~lslrrr~rrtc~r ~?LTYL- riucer in sh;ile soil. Additioi1al rosearc.11. especially in ings. i~owel~er,rcs~rlt in n grn.ss?j httl~ittrt lrjith re- i?~ountainhardwooci types, is ncetlcd boforc widcsprcc~ti cE~cced browse ctntl ~i~nstsicpplics bel,tpr swiipd lo pmxcihed burning is recomrne~icled lor man:igcnient quail nznnagernc>nl ti~criz to deer nzc~nc~gcntenc. purposes Forests of the future art? going to be different than I'~.escribcd hurning is an c,xccllent forest maniigemcnt those we arc used to seeing. Product v;ilues and land tool. It is fire al)plicd in a skililul ii-~anricr,unrler oxact- costs are demanding that forest niansgerncnt plan for ing coiiditions of weatl~cr,to fuels of the I'or.cst. in ;i pretlc- maximum monetary returns. In niost cases this requires tcrmineti area for a spi.cific purpose, lo achieve certain highly marketable specics at greater stand densities. rrsults. A trained, cxp~.riencotIforester or resource mari- The old "hit and miss" higli graded forest which set~mcci can use it effectively wit11 little risk. In atidition to to have groduccd fine game habitat, is now ;i thing of coritrolli~lgundcrstory vegct:ition ;inti improving wiltllif'c, the past. h;ihitat, prescribed burning is \viticly useti for rctlucing This llew soutlililnd erivironment will require more excessive fucl, contrc~lli~igbrowti-spot rlisc~;isc~in lo11g- knowledgeable and intcnsivc manip~rlntiorlor undcrstory le;if pille. ;ind prejj;iring sc~>dl~rtlsant1 1,l;inting sites. habitat, planning within land patterns, nntl a recognitio~i This pa1,c.r concerns pr~~scribcciburn as it nffccts deer that all species of game c;~rinotbe accornn~cidatetiwitlli~i iii southern Cori~sts. every land pattern. The land pattern consists of i li 1,eopold 119.501 melitions priscribed. burl1 lor deer as harclwood and pine managed for products requiring 1;irge Ijcing thc cl1capest technique to iiicrcase decr carrying cliameter trees, for cxi~mplc,sawtimber and vcnc,er, 121 c:ipacity on some ranges. pilie managed for wood fiber, and (31 cultivated cropland "ire in a fcirc~it enviroiimc,~~th:is bccxil ~rsc~lwith and grassland pasture. The hatlittit quality of tliis en- rnnrkcd success. In 1940, I3utl ,J(.nkins of tlic. Mic*kig:iii vironment will depentl largc,ly on: tkc iirc;i of forc,stcd C'orisc,rvation Departmc~i! bcgaii ti scric.s of stritiics to land undcr lo~igrotation, t11;it under short rol;itit>n.;igri- tlctcrmine tile vnlucis of fir? in cor-it~-ollilig111;iiii succcs- cu1tur;il lands-. and how far thcy arc allart. It is fortu- sioi~Cor grouse anti tlccr. Iiix found tl~atto nnai~?t;ii~i nate, in our opinion, that considerable acreage of forcst tii,sirable openings, they sl~ouldbe hrlrnc.tI ;it :3- to T,-yc\:ir lands will be nian:igecl in even-aged groups. This pc>rmits interv;ils. The nonopen ~jc~rtio~isrc>s~~ondt~i .ivit!i in- treatment of uriderstory witliout damage to standing creased browse, legumes, anti fr~ritcrops of slir~rl>s. timber--at all but sapling sizes. Firc is triore prnctic;ihIe R;isic;illy, there arc two pcrioris ill tllc life of :I pitie under even-aged management. stand when fire may be s:ii'c~ly usi~i\vitliout t1;ingc.r lo Firc as a tool for understory control has hccn iiscti for )iirics- prior to regeii'r;ltion to scxcurc Fnvorablc! coritii- ycx;rrs by managers where game birds were the prime tions Sor gcrrnination, and after the trces :ire tall c'iiougli sue11 3s old JIOIISC sites, Iioneysuck1e thickets, anti otlio lic,at. Wlict11c.r iir iiot ;irly 11( tic~i'it:;iic:criic' to tloer from Sruit ;11id hcriy p;itclies ~~iovicieslinmec1i:itc looti si~l)l)li<~ 111~t-(~geii(~r:~tiot? b~trti fi!>pf,ntls :I lot on ~v1i;it vcg<:t;itioii for doer. Whc~rerunlier oak iQicczrcits pli~t~ilr~iOCCLIY. \:-as lhcirc. prt~vioiirlyziiiei ~vllr~ilic~ror ~ri,t otlii~rsitc pri,ji- it I)c;irs vigorr)risly Soi. se\.eral years fr~llo~riligfir(.. 1"ii ;i~.;itic)ns,strc11 :IS clic~pping :iiitl lii~rbicitles,\vcrcy irsc~l. gre;~'tlyiricre;isi.s tlic l)rc~teinanti p1losj)liorcius ct~iitei-i of browse. 'rhcsc n~iiii~r;ilsarc rurc,ly ;icIc~cjiiateoli so~illi c.r~itlecr r;ltlgils, 111 fnvo~.of 3- to 5--yearhuriiing int<~rv;il: the tn:ixil~itl~ntic~sii:ibl(~ brit.ivsc con(1iiit11is gc~ii<~l.iill, pcxilc, ;illti jnst hegin to tlcclinc~,at [lie cl(ise of tlie intcbl val. 'I'l~cscgtiitics. of course, nl~~sll?c ~ipliiierlwith kiiowl 60 years M:ituriiig s!:iritls i~rc~vitlrmajor ticcsr 1i;ibit;il ccige of site coritiitions. 1,i.t'~lortk at tlicsc conrlitic~iisi ;~inri yic~lcis the rnaxirniirn in bcnc.fits Eron~ 1ircscrii)eci hiirniiig. 1Ierc fir<, serves several purposcxs, 1 1I retiuces ~irid~~sir;iklc~vocitly gro~vth.121 i)rings pnlatahlc spccit,s Nutrition tio;zrn within rcacli of ticcr. 1:31 iini)rovcs t11r t~ulric~nt DcWitt :inti Ilcrby 119551eom1xrred nutric,!it cc~titcn titiaiily ctf the hrciwscs. 14i gencrkiiiy 1)rovitic.s ii-icretisc~:i of reti maple iAcc>r ri167.1~~1~).white o;ik (QIL('T~I~SCL~OCL, l~c~rh;icc~c~~isfoods tindi.1- sc~niicrpeli ovcrstory cotltlitiorrs. flowcrii~g ciogwood i(lor.ncis florida). and riruritilc~;~ at~d1.51 cXncoilrngesi~ritierstory fruit anri h~rryprotluc- ~grccxrrRrier iSrnilrt.r roti~l?difolin)following a low ;in( tion uildcr sparse ovc.rstorics. high inlelisity firc at the Patuxent Research Refuge ii While risenrch is still ncetiotl on resxilts of seasonal 1917. Protein ctil-itents urcre signilicnntly higlier in thm fires under .i,;iri(ius sitenbriers (S~t~iiani.;rnd Swect1c;ii' iSyn~ploc.osi. quail. Wl-iito-tail deer are known to consume a large variet: Wititcr buriiing on ;i 3- to 5-yc.ar iiiter\9al :is ;r manage- of arlditionnl footis such as flcshy fruits, acorns, lravc: ~-nc\nirrir:tlnod rcsults iii ;;xi c~xcclleiitresjtonse of leg~lines, grasses and other herb;lceous pl:ints and fungi i Stitcle keeps browse p1;irits low, and seldom kills plants, injiires and Shrauder 15167 i. 'I'his was supported in more? rccen g;ime or tlcstroys riests. Vcgctativc rcxsponsc, Irowcver, studies hy tlie Southeastern Forest Experinlent St;~tioi v;iric>swith tic~nsityof thr: o.i7crstory. In sclnishariy areas. and citlicrs 11,ay 19fj7i. Proscribed burn. in ;iddition t, ltgiirnes such as partridge pea and beggar-tielis fre- woody hrowse, also iniproves yicxld and quality of I'ruit: ci~tc~itlyfollow fire, whilc in opcri arcx;is, wootiy p1;ints. griisses and hcrhaccous plants ( 1,ay 1956 1. I~;~itl\voodsprouts, and gr;isst,s often comc iri after burn- Fl(41y lung1 which fortun'itely occur without regarc ing ~Ilcvetand 11111)kinsIlf(ii1i. Plowing out selectcti sl~;is to \tand dcnsity, respond well to fire Nitrogen-toleran species fruit prolifically Their ability to nature rapidly nature's ways of setting back plant succession. It is an following stimulation-hours as compared to months for iniporl;int technique in managing our wiltllifc rc3soiircps. other food--offers the possibility of using fire to corn- Considering all data, there seems little ch;irice for pensate for mast failures and other food emel.gerieics. much damage to deer range in pine forcssts by ~,resc~ribeti Several species are outstariding fire followers and at fire. Benefits from fire increase as the ~~ndcrsttrrybegiris the same time are preferred decr food. Tlzcy are: granu- to grow out of rcnch of deer. Large-scale burninq siioillti lated bolete (Suil11c.s yran~~lafus).little red r~rssulaiR~ts- consider tlie pattern or distribution of tkc I)~i~.iisif s~rla ro.seipe.q), swolleii-stalkecl armillaria iflrrrzillaric~ maximum deer, quail, and turliey poi>ul:itions arc. t1esirc.t-i. ~~eritricosui,Tennessee hygrophori~s(Ef?jgrophorzcs few In gcnerai, the pattern that produces the rncist tiiversity ?zcsseerlsis), Clitocybe iClitocybe). arid dcsstroyi~~gangel of understory habitat will be tlie most he~iefici:il. iniraanita vcrna). The ability of common species of In addition, the following considerations for (leer man- mushrooms to withstand shade and respond to fire will agement are important: become i~icrea~~nglyimportant to wildlife man~lgersin the forest environment of the future. 1. Prescribed burning is ari excellent forest mannge- nient tool that economically sets barli plant suc- Cover ccssiorr to a more favoralsle game habitat. In frecjuently burned pine forests, herbaceous planti 2. A cool winter burn is preferred over ri summcr usually form the dominant undeistory cover Converce- burn. Summer burns cirdariger late-nesting quail ly, such plants arc iuppresceci by the ,~ccumulationof and turkey and decrease i~nderstoryfruits ;xntl llttcr when flre is excluded In additio~ito herbaceous forage preferred by deer. An clxccption is a tleiist~ plants, certaln woody specles such as tltl, trurnpct hardwood-choked-pine stand--a series of hot sum- creeper, and yaupon reiprout prolifically .ind provide rner reclamation fires cluring the growing season dense cover for deer Periodic burn~ngin pdtterns \~ould rnay he needed first to elimin;~tchard-to-kill hard- provlde increased nutritious foods on one area while wood species. dense cover for deer can be produced on adjace~itareas 3. A burning interval of about 3 years is ideal for As plants resprout prolifleally, hunter use generally decrfood and cover. This interval permits an ex- declines on dense areas cellent response for legunzes, keeps most browse plants low, seldom kills plants, or i~ijuresgame. Soil blast from vigorous runner oak is produced the second year. Food benefits produced, mostly Soil studies meritioned by Suman and Carter (19541 sprouts, can last frorn 3 to 10 years depending indicate that the chemical characteristics of the rc?latively oil area and type of vegetation. infertile sandy soils of the Coastal Plain region are not materially affected by burning. The Charleston Etesearch 4. Palatable species of fleshy fungi occur ab~~ndantly Center ISEFES J found that a 10-year annual bnrning following a fire. treatment in flat sandy soils increases the organic matter 5 Repeated annual summcr fires producr a gr'lssy content of the top 2 inches of soil. They found no cvi- habitat more 51;lltable for qu<111tllati deer dence of harmful effects to these soils by prclacribed burning The type and frequency of annual winter flre 6. Protein and pliosphorus contents of plants are resulted In no detrtmental effect on the physical prop- increased by burning. Increaseci rlutritionnl bene- erl~esof density, porosity, or perco1,ition r'ite Houcver, fits are apparent for aboiit 2 years. when burn~ng,tlie so11 must be wet or damp trot only to 7. I3~1rningpromotes prolific rcsprouting. thereby prevent fire from penetrating the so11 and kill~ngbelie- providing increased cover for deer. f~clalmicro-organisms and consumng h~~mus,but alio to protcct the layer of duff ,idjacent to the so11 i Dixon 8. Chemical characteristics of sandy soil in Coastal 1965) 'lains are not affectcd by burning. Physic:il prop- erties of sandy soil are not tietrimentally ~iffected Costs by burning if soil is wet or clamp. As mentioned, fire is an itzexpcnsivc: rnetliod of setting if. Evaluate effectiveness of prc~scrihcclbirrn by: ob- plant succession back by provitling not only abundant, serviiig scorch on bole of tree (should be 18 incllcs but more nutritious dct~rfoodsin n browse management or lower), no discoloration of crown foliage a11d program. Silker 11961 I found that when using experi- complete blackening of the r~~~gliand undesirable c~icedpersonnel at wagi' levcls of $1.00 per hour, burning h;irdwood untlerstory. 13;rrk cracks extending into costs may r:lnge from $0.30 to $0.50 per hour for b:ick- tlie cambium at groutid level iirdicate success of fires in r1orm:il pine-h:irdwood fuels ;ind $0.12 to $0.50 hardwood kill. per acre for strip Iieacifire burnitig in light to moticr;~te 10. Atiditiotlal resoarcii okl i~resci-ibctlbrirtl in hard- fuc.1~. Night burning will probably result in decreitsed \vood typeswitlr diffcrctit soils, especially i11 tile fire costs. Mountain province, is neetic\d. Under l,resilnt knowledge, avoid burning in hardwood types for CONCLUSIONS manageme~itof wildlife. Fire iti the wrong place ur1dc.r certain c.onditiotis can 11. Fire is ;in excellent, iiiexpcrisive niethod of pro- be onc of the greatest cncnrics to mailkinti. It is one of viding i~icrcaseddec:rfood atrci covi,r in pine and pine-hardwood types. While additional deerfoocl Lay, D. W. is procluccri i~nc.ert:iiri hard.~z~oodtypes, rtlucii rct- lijli7. In~ljort~inccof variety to xouthcr~ideer. Soutll- sc:irc.h :inti evall~atioiiremain to be done. cast. Assoc. C;arne and I'ish Comni. 18th Ar~riu. Conf. Proc. 1964: 57-62. LITERATURE CITED I,copolti, A. S. ilcvet, D. D.. nricl IIoplzins, M. I,. 1950. Iletr in reluiio~ito plant succession. FiSteenth liifji:. Rcsponsc of wiltllife habitat to tile j~rescriixxl N. Amcr. Wiitllii'c C'onf. Tr;r~is.11150: 571-530. burning program or1 the natioiial forests in South C:irolina. Southeast. Assoc. G;inic anti Sllrs~ltic~r,P. A. E'isli Conim. 2151. Aiiiiu. C,'onS, t'riic, 11167: 1211- 1964. Pri~scril~c~dbi~r~i in the Ritlge ar-icl Valley I'rov- 133. iricc. Virginia. Ui~pui~lisl~ed. lT)c,Witt, J. B.,nnci Derby. J. V., Jr. Silker, T. FI. 15).55. Changes in nulritivc v;iliio of browse 1tl;irits 1 Prescribetl burning to control i~ntlesirnhleharci- following forcsst fires. ,I. W~ltliifcMari;igo. 19: wootls in souilic~rn pine st:ii~tls. Tcu. Forest 65-70. Scrv. Bull. 51, 44 p. Dison, M. J. Stiteler, W. M., :irid Slir;rutier, P. A. 1965. A guide to fire by prt sc.riptioi-i. USIlA Forest 1 Prc,duction ancl tiecr iitiliz~rtionof woocly browse Serv. Soulli, Rog. 32 1211. on the ,Jefferson National Forest, Virginia. Ii;ills, I,. K., and Rijtlcy, 'T.II. ic1:iiti)rsi Soutileast. Assoc. Game arid E'isli Con?m. l8tli 1961. Deer browse plants of soi~tlicriiforcbsts. USDA Arinu. Conf. I'roc. 19(i4: 21-29. Forest Serv. Sout11. aiitl Soiltl-ie;ist. E'i~rc!st Ezp. Straiisky, J. J., :inti IIalls, I>. K. Stas., 78 1113. 1967. Timber and giilno food relations in pinc-hard- L:iy, D. T.V. wood forests of tlie soutl~ern U~iited Sttites. 1956. Effccts ol l)rescrihed bur~iingoil ror:ige and Fonrtecntl~Cotigr. I'roc., Int. Union Forest. Rcs. mast protiilction in souihern pilie forests. J. Organ. Vol. 7: 208-217. Forcst. 54 : 582-584. Sum;iri, R. I.'., arid Carter, R. L. Lay, D. W. 1954. I3~1rnirigand grazing ]lave little effect on chem- 1957. Brourse cluality and tile c~ffcctsoS prcscribc:d ical propcvAics of coastal plain forest soils. burni13g in soutlicrn pine I'orc,sts. ,J. Izorcxst. 55: USIIA Forest Serv. Southeast. Forest Exp. Sta. 342.347. Rcs. Notcs 56, 2 pp. The GQ~~sof the Southern National Forests in While-Tailed Deer Management

Donald D. Strode Divisic,n of Wiliil~ieMati,~~ct~ie~~t Forest Service, U S Dcparttncrit (~iAgrici~ltirrc Wc~il~it?gtot~,D C Wayne J. Cloward Wildlifc~arici Rarige Man,igcmcni Soiithcrri i?egioii, Forest Servicc, U S Dcpartniet~tof Agririiltiire Atlat~t~i,Gcorgia

Tiic. ,i:iiilc-rciilctl dcc?r Iiiis I)c>cn ~~cc.s!uiilisl~t~tlrli~r- va1iic.s of tile v:irioiis rc,.yi,iirctvi, ;iriti not ~loc*c~ss;iri~y i?i$g ti^? 1111~120 ycczrs (is ti~crlzosl i~til~o~.fcc~tt(in({ iilosl tiic c.oil?l>inntio11 of iiscs tiiiit .ivill givi. tlic5 q~'c':itc\st i11inlc2rons big-yciiflc~ s!)c~c~ic~.s111 1\11? SOltl~1(V'ilNotio?lol tio1l;li. rc>tilrzi or tl?c, grc;itc.sI iiiiil oiitl)i~t jorcsts. It is too i~~ot~ci.oilsili i?iir~iy iirc>i~s. Wilrlli/~, i~cci~ilr~l?i?u~tngr~inc~lt (171 f11osc. for-c'sls is J~~IL:ciiii~c>il lo cnc'oii r-n yc .sl~c>cic~,s1~c.sl slt it(>tljor 11 picri ic.itl(i~.cil.c>ci r llrcy7 N;itii~i?;iIforc~st objc.c~tivc- in miltllii'c~t??:iritigc~it1c'nt crlltl lic~l)i!nf,(LS tic,lc~iriiiiedi~y tr jr~i-c.si-ii.itlc S~L,.).P,I~ is to m;iii;rgc~ li;ibit:it so t1l;it hot11 g~in-~c';itiii ii111ig:ir-ilc~ o/ l?criiiitit ricc~i1.s. This S~LI.I.PU:d lo oll~cr7~:ilrili/t> spc)c.ics. :rnri (21 in nccorcl;ince wit11 tvildliCe's ri~crc~iitio~i:iltiriti The white-tailetl deer is tho most important ltnd rilost rc~latc~dpublic uses anti v;ililr~s. 'l'11(~ Soiiti~c~i-11llc~gioir nu1nc:rous big-garne species on all the Nu1ion;il Sc~rcsts in siil~plcmcntingthis policy st;ltos: "TL1l;in;igc ;ill h.;rtion:il in t.he Soiithcrn Region. Its iio!)ul;irity is very high with forest .ivildIife 11;ibit:it to niect the requirc~rnc~nlsof the both the hunting and nonhunting public. It is scarce or proper wildlife spccies, in niin>hc.rs c~oiisistontwith caipn- absent in certain mountain ;ireas of suitable habitat, city." however. Despite certain hahit;il prcferenccs, its ability to aclapt to various h;ihittit contlitions is a11 in~portnnt. GOALS factor in its management atid polsulnrity. I'o~j~tltc~ioii.--'~I~eIzorcst Scrvice \vc~cllcl like to see It is hard to realize that white-tailed deer werci Sound v~i~ilc-t:iilctldcer c~stablisl~etlanti ~r~:i~>agt~lin 1~11 suitable only in remnant popu1;itions ill the southe:ist :is rcxccntly 1i;ihitat. In rirltiition to :ireas of ui~useti~.;iligC. ~IIC~C iiri, as 20 to 50 years ago. Thc success of rccsiablishing the c~x;imj)lcswlit>rc popiilatic~nshave cxccoti~.tlti?(. c:irryi~?g white-tailed decr on the N;itiori;rl forests was not ticcom- calmhilily of t11c. h~ihitat.Ifere tlic li;irvest ~llOllliih(, lib- plislictl without problems, however. 111 ni:itry areas deer c,r;llizc~ti. populations have gone Irom a "Samiiic to a fc;istn level W lri tc.-fnilctl tlcr~I~~~i~iii!t.-lii thc. p;ist, ciii:ility 11:ihit~it and are seriously darnaging not o11ly thcir own habitat. 11~1s]>(ten either a "fctist or a f;imine," tlc.)~c.ntiing pri- brit also other forest resources. It is obvious that 11-1~: in;irily upon tin~bcr c.utting 1,rogr;ims. Uecr 1i;ihitilt number of animals has to he retlucecl to :i rc;isonablc nrusl bcx produccd 011 a sust;iinctl basis hy nia~~il~ul;itinr: level. FIowcver, public rosislance to sound cic,or-huntitig food ;ir~ticover over a period ol time. Forest resources regulations renlains as a continuing barrier to good man- milist he bettcr coordinatctl, c,sl~ecially timber rn;ili;lgc- agemct~till many States. nielit. IZc.sc~ccrclr.---K~~owledgegai~ieti fro111 habitat rcscnrch MULTIPLE USE on tllc white-tailed deer is the hiisis for its m;~tinge~nc~nt The doctrine of rnultiple use is tho cornerstone of Iaiiti 011 tl~cNatioi-i:il forcsts. More rcsc~tircl~is nc,ctlctl to managc,incnt tiecisions on the Nntioi~alforests, National Inati:igc. tic,i,r habitat on ;I sust:ritic~i btisis. Thc>rc:forc, gr;isslands, and land ut.ilization projects. Unfortun~itcly, it is ;I goill of tl1c National I'c~rc,sts to cool~crntt'with ;ill it rncans many things to rtlaily pcol)lc. IIowcver. tile iriti~rc~stedp:irtios anti agencies iir furtlic~ring Iltibitat multiple Use-Sustained Yicltl Act of .Jurio 12, l!)fjO, rcscarch on the wl~itc-tailoti tlccr. states: Q1~cllifyof r~nperic~~~c~t~.--Sho~~lcI\vildliSc~ inanngelntnt Multil~lc~use is the n~;inugcrnc~irtof 2111 lhc~v:irious i)~'ovitic, a cluality c~x]jeric~~icc!wit11 variety, or just as renewable surface resocirccs of the N:ition;il forcsts ~ni~clihrlnting opliortuniiy as ~~ossibic~?I"~irc.st Service so that they are utilizori in the con~bitration tllnt. 111)licy is to "I'rovitle, in cool~c,r:ilioll ~vitll l11e Sl;itcs, will best moct tile needs of tlio Anlcric:rn peol~lc; Illliiting iinii Sisliiiig 31.~21~suitablc~ to tlic~vtiryitlg neccls. making tho nlost jutlicious use of the land lor some interests, anti sl;ills of t11(, public.; tll(~ilitc,iit bcing to or all of tl~csertxsoiirccs or rc'latecl services OVCI- ;ireiis rc~tair-i sclme areas m;~!?ugedfor q~i;ilityof c~spc~ricnce large cttiougll to provide sufficient 1:ititude for ijeri- r;ithc.r th:rn numbers of i~nrlicip;iiits." odic acljustments in use to conl'orrn to changitig r 17 Ire l~rcselit(leer progr:iri? in t111: Soi~tl~c~rt~lapoiis I-Tunting ~2i-c;lon tlic Ila~iitlr3oonc other, without irnpuirrnont of the i~rotluctivityof tlre N;itional Forest ill K~>nluckyis a11 cs;iml)le of clcrality land, with consideration being given to the relative llutlti~ig.A long history oi ;in ovcrl)ol)ulntc!t1 tiecr r;inge on the Pisgah Wlldlife Management Area on the Plsgali ted kill of 34,000 W~delyvarying dollar values have beel National Forest in North Carolina has made people assigned by various States and areas for harvested deer there demand quantity Recent figures range from 9300 to $1,200 IL we uil the minimal value oi $300 per harvested deer, the 196 HABITAT SURVEY harvest of 34,000 deer had a monetary value of $10,200 000 on the Nat~onalfort.\t\ In the Southern Region Two years ago, the Southern Region started a program What are the nonconsumptive values7 Certainly thl of having each forest survey its own wildlife habitat. wildlife photographer, camper, and hiker value thei Part of this program entails selecting wildlife species experience of observing deer in the National forest: to favor in habitat management on established units for The opportunity of seeing deer, or other wildlife, is on1 the next 10 years, based on existing habitat conditions. of the important considerations in family outings an( This does not preclude the management of all spccies automobile trips through the National forests. Wherea~ found on the unit; however, each forest will give priority other big-game species, such as the black bear, havl to those species for which the habitat conditions are best declined as a result of habitat changes and hurnan in suited. Therefore, the forests will emphasize deer habitat fluences, the white-tailed deer population has increase( management only where the habitat conditions warrant. steadily for the past 20 years. And deer will not be favored on those units better suited Since the white-tailed deer is so adaptable, its import to quail or other species of wildlife. In certain units, several species, such as deer, squirrel, and turkey, will ance wlll contlnue to incrca5e on the southern Nationn receive the same priority in habitat management. forests We are all aware of the act~vitiesand pressure from increased public use and demands on the Nationa At present, habitat surveys are completed on the forests They will not have the same adverse effect upoi Kisatchie, Cherokee, and Jefferson National Forests and white-tailed dcer as on other big-game species. There on the National forests in Georgia. Surveys are planned fore, this deer should continue as a major big-game spe for the others. The rate of accomplishment will depend cies in the Southern Region of the Forest Service. upon personnel ceilings and financing. The habitat needs survey does not intend to deempha- size deer in the forests' wildlife programs. However, HABITAT MANAGEMENT these surveys will probably show that deer have long Habitat requirements and preferences for both fooc been dominant in areas where the habitat is better suited and cover for the white-tailed deer are adequately cov to other species. This does not meal? that habitat needs t=red in other sections of this symposium. for deer will not be considered in these areas; rather, As a land management agency, the Forest Service i the needs of other species will have a higher priority. charged with management of wildlife habitat. ~h~~~ two ways to accomplish this responsibility. The first IMPORTANCE OF DEER and bv far the most important, is through- resource coor How important are dcer as a National forest resource? d~natlon,and second, direct habltat improvements In 1967, there were an estimated 289,000 deer on the Direct habitat improvements are more "glamorous, Natlonal forests In the Southern Region, and an estlma- but the heart of the Nat~onalforest deer habitat manage

Wildlife openings on the Jefferson National Forest in Virginia. nient programs is coordination with all other resources, to the possible detriment of other important game and especially timber. Forest Service Handbook 2121.4 nongame species. This situation was partially responsible "Multiple Use Guide-Southern Region" provides the for the establisliment of the "Habitat Needs Inventory broad guidance in coordinating resources. Further guid- Survey" presently under way in the So~ithernRegion. ance is provided by Forest and District supplements to Habitat corlditions do not remain static. Therefore, the Regional Guide and in individual resource plarls. over a period of time, habitat emphasis will change on These supplements are used to further clarify local situ- a given unit of land. Let's look at a unit of land in the ations and problems. southern Appalachian Mountains. Just after timber re- At the present time, direct habitat improvements for generation cuts, habitat management would emphasize deer are limited. Where improvements have beell estab- deer. This same unit for the next 20 lo 50 years will lished, they generally consist of openings, waterholes, be best suited for grouse, arld from 50 to 100 years for food plantings, and release of food-producing species. squirrel and turkey Keep 111 miild there will be some In addition to these direct habitat improvements, the deer In thlr unit throughout the 100-year period, but Southern Region constructs hunter trails, parking areas, durirlg this rotation there are periods when the habitat and hunter camps. Although these activities do not im- is best suited for the management of other species prove the deer habitat, they are Important in facilitating The future of the white-tailed deer In the soutllern the harvest and help provlde a more enjoyable hunter National forests holds great promise. It is limited only experience. by the potential of the habitat and the ability of the The great lmportarlce of white-talled deer in the South Forest Service and the respective State conservatio~ldc- has encouraged a tendency to overemphas~zedeer habltat partments to wisely manage this species and its range. "Be Goals sf State Conservation Agemies in Deerherd Management

Frank P. Nelson Solith Caroltna Wildlife Resources Department Columbia, South Carolina

In this pnyler (in cztferizpf is n~ccricto erttrctcf from would, as suggested by two respondents, result as a b~ the ciirrctors oJ 16 solit lien,stcrn Si(ltc. gcrme c~nrljisl~ product of proper managcmcnt. This writer's coiiclusio acgt~?ic,iesso?itcJ of f 7Lc.i~ f lzo?~glzfs z~'ii11~.?g(trd to ~~ihitt'- is that directors want maximum hunter days eonsister f~tilcddeer c111d to present cr c.ontposifo pictzcre oj with range capability and physical condition of herc tlzeir gcnrrul co~lceptson (I rcgionc~l1)nsis. This spc3- without depletion of herd density. They would accow cics is lzigl~lyii~~portaiit. It rccei7:cs tncljor c~ftenfiori plish this by adjusting from year to year the arcas oper in tl~egctme ~tzn?zngc.?rLenf progrcci?~. Ii s?~ppoi.ts(L the sex to be harvclsted, and the hag limit. Tlicy woul~ sigizijic.a?zf portioit oj the total l~zrntirtg progrnnl, hope to produce some trophy antlered-deer but this i boflz jr~iizthe sfu71dpoiizt of ecoi~oi~zicscinrl hzc~zter not essential, since any deer bagged is coi~sidered ejjorf. Most States espect cLn ii~creczscin deer d(.rc- trophy in some areas. Trophy is a relative term. si:y, tii.sfril)?clion, criid the, ctcc.on~prr~z?jin~probl(>??ts 2. To what extent do you expect deerherds to sugpor involz:etl in lzerd riz(~?lagc?t~enf.So7i1e trepidation is your total hunting program? Will this change i~ tz)idcnt among directors rcgardi?~g(~n i~zcipie~zt trend the future? tou:ard fee llz~ntii~gby incl?ist~.ictllandou:ncrs. The States which interpreted this as meaning to suppl: In attempting to present the general picture of the a figure gave 30 to 50 percent of the total license sale, role of white-tailed deer in overall grime management as representing deer hunters. A different way of an programs being conducted by State cotlservation agencies swering relates to man-days effort in which three State: in the southeast, a questio~inairewas prepared and sent gave 11.9, 15.0, and 25.0 percent of their total huntin5 to the various directors. States included meinhers of the effort being attributable to deer. The latter figure come: Southeastern Association of Game and Fish Coinmis- from a State which probably places more value on thc sioners, and Texas and West Virginia. Emphasis was white-tail deer in its overall hunting program than dc placed on the need for the directors' personal attelltioil any of the otlier States interviewed. rather than their passing the form on to their deer biol- In circumspect, it appears that white-tailed deer con ogist or someone else for conlpletion. 'I'he directors rc?- stitute a sig~iificaiitportioii of the surveyed States' tota spondcd admirably and in all but a few cases, presented resource and subsequent hunting programs. Respondent: their own views, and individual concepts and philoso- also indicated a coiitinued expansion in distribution anc phies. Even in the exceptions, persons of adininistrative densities, riot only in those States which have supportec responsibility furnished the desired information. Tl~e historic deerlierds but also in border States where res thoughts, therefore, presented it1 this paper are more toration efforts are proving to be effective. Some deet philosophical than technical, anti rcflcct personal atti- biologist may be shocked to learn that the white-tailec tudes of the people who must treat the entire wildlife deer is not necessarily the most important species a. resource. rather than a single species such as that with this time, altliough it is expected that deer hunting wil which this symposium is rnost concerned. continue its rather rapid inercase in popularity during Qucstioils were ciesig~-icd to be tliuuglit provoking. the next decade. Simple "Yes" :ind "No" answers werc impossible witliout 3. What are your p1annt.d limits of deer density'! flow also elaboi.ating. It was t11o11ght earlier that the answers will it be eontrt~llod?Arc you conccrrled with cror would lend then~st~lvctsto consolitlatio~i;1tic1 permit for- darnage'! R;rnge deplt.tioil? mation of a tyl~icalsout11e;istern situntio11- -2 stcrcotyped director, if you will. Tliis provccl to bc rather clumsy, There are 110 standards for density. 11 is the genera howe\c>r,ciuc to the wide r,rrige ot condrt~oti\encou~rtcred consensus amoiig the respondents that density will, oj in the region necessity, be variable, due to the myriad environment< conditions whicli exist in a giver1 State and wllic'cl cxisl The followitig rcnr:irks :ire tlcsrir~c,cifroni the comments from one State to another. Thi, directors arc, cogrrizanl of thc rcsirectivc directors and also rcxilcct, 11o doubt, of thc ncocl to expc~ntirnorc effort. in management ii- il~eteml~ering by this writcr of tli(. cotrtlitions in his own areas of low carrying capacity as a means of iml)roviiii State. Questions and an~ilysisof ans\v(,rs follow the same distribution, par~ticulnrly,ill conjunetio~iwith areas ol order as werc listcd it1 the qucstioiitr:iirc. high liunter clc~rsity--~-centersof human population. 1. What is your gc,ticrnl aim in cleer nl;lll;~gemc~nt?Is Crop ciarnnge is not n serious problem but occurs ir it to provide maxiniiim lrur~titlg clays'! Or is thc localized arcas. Ka11gc dcplt.tion is a little more serious cmphasis on quality'? generally, and there is expcclcd to be an increasing Mo\t respondents rcxplied tv thcs cltett "maximuni alerti~cssfor this problem as herds develop in dcilsitj deer hc1c15 coniiitei~t wit11 r~i~gc'c,il~.ibil~ty " (Ju'111ty and distributioii. (lontrol rne;isures will involve adjusting liu~~t~ng(to inclutie 1lol)hy Iic.,itii, frceclom frorn oiliei liu~ltcrpressure through special seasons, special area: Iruntt~ri,and bettci tirr'in ,ivclc~gclllu11tc~r-5ticci~is rdtro) ~rlltlcither sex h:rrvesting. One respondent po~ntedout the ambiguity of the term from about the third most important species to the nuni- "range deplet~on"by stating-"Deer r'rngei can become ber one species. depleted as a result of natural plant succcsslon or 'is a Input output ratio is considered to be highly favor- re5ult of population densit~csthat are incompatible w~th able. Of the species which require management in the the quantity and quality of footl enczgy ,iv,rilable to form of restocking, habitat improvemetit and harvest the deer population The reyource man,lger (he he wild- regulations, deer obviously produce a high degree of l~feror forester) shoulcl be wise to rccognlze the fact hunter opportunity expressed in man-days. Most respon- that product~onof a r~lsfaaned $table h'irveit oi deer dents were quick to point out, however, that squirrels li a product of FILT~CLZTZP~~tal>l~ SOU~CL' of energy ' and rabbits support considerably more man-clays of hunt- 4. Will future emphasis be on permit hunting'? ing, at practically no input except harvest managcment. 7. Do you detect a tendency on the part of conimcrcial This question was deliberately worded to evoke an- timber companies to charge a fee for hunting thcir swers that might define permit hunting. At least. some lands'? What is your reaction to this'? of the respondents reacted by describing two kinds of permit hunting--a type of lottery wherein a predeter- To the first part of the question, the northern border mined number of people would he given the opportunity States replied in the negative. All other States replied to participate in a special hunt; or requirement that any in a positive way, ranging from slightly to definitely. licensed hunter must buy an additional permit or stamp The rights and prerogatives of private landowners to hunt the designated species. Some of the others, in are clearly recognized, and supported and defenclcd. addition to the above precepts, meritiuned fees being Respondents did, however, express the feeling that rev- charged by landowners for the privilege of hunting their enues collected should he directed toward a sustaining acres. game management program on these lands. Most of the States have entered cooperative agreements with com- In a general way, most directors prefer to conduct mercial landowners for game management and public their deer programs with a minimum of regulations. This hunting. Hunting is generally offered without use fees. entails merely setting a season and hag limit and not Leascs to conservation departments are generally with- regulating number of hunters or their distribution. Some out payment. Two States have had leases on some free- refinements are added from State-to-State but there is leased lands terminated. an avoidance of permits for any purpose, except permits for the purpose of additional revenue and for areas which Comments from individual States indicated concern require special harvest. Some have utilized lottery per- that the commercial landowners may become so en- mits with satisfactory results. Some have experienced grossed in wildlife management that they might exert disappointment, especially in regard to public relations. detrimental influence on the decision of the State ad- ministrative agency. In summary, the opinion appears to favor an avoidance The average director is not particularly concerned of permits, unless necessary to facilitate special harvest. over a fee charged by a landowner. He recognizes prop- 5. To what extent will out-of-Slate hunters be encour- erty rights and that fees collected could improve game aged? managcment and subsequently hunter opportunity. He admonishes the landowner to accept responsibility for None of the polled States makes any great effort to protection and proper game management procedures. attract nonresident hunters. On the other hand, all He would rather have the lands under cooperative agree- States treat the nonresident on a par with resident hunt- ment, without payment for leasing, and without fee to ers except for increased license costs. Provision is made the public user. He thinks that fee hunting will be ac- generally for trip-licenses. cepted hy a portion of the hunters and that public lands Two respondents expressed a feeling that cvci~tually can support those who cannot or will not pay the use public pressure may force a "favorite son" policy. One fee. He wonders if increasing costs of use fees will he stated that nonresident fees may be raised to discourage detrimental to his efforts to obtain authorization for this type hunter. license increase or will affect his chance of establishing No one objects to the revenue generated by this source. a big game stamp. He thinks that fee hunting on these It is expected that no great change in the status of non- lands is inevitable. resident hunters is forthcoming. While this presentation obviously does not, precisely or comprehensively, treat all the ramifications of the How do you rank deer among your other popular 6. status of the white-tailed deer in the southeast, some of species? Do you think of deer in terms of a species the current thinking of administrators is amplified. Each which, with relatively little management, can pro- State has its own problems, not only with deer, but with duce a high return in recreational opportunity? Can other game species. Some States place more value on any other species offer this comparative return for deer than others. The directors' attitudes and concepts the same input? cannot he taken lightly. After all, these are the minds Again, as in question No. 2, deer rank according to that determine where it is we are going, and influence whatever criteria the respondent decided "rank" meant, the speed we are to maintain in getting there. The Goals of Private Forest Holdings in Deer Management

Raymond D. Moody F~shand Wildl~feDivision International Paper Con?pany Mobile, Alabama

Far7ners and nziscellaneous intli~)idzialsown th? day'.; goal 15 not ncces5arily the same today nor will it greatest portion of forest land in tlzr So~~th.Most be the sarlic tonlorrow This li why it ir difficult tc of thcnr like to hu~~edeer on their lancl. Their ?nun- come to any s~ngleconcluslon concerning the goal5 of ugement goals have bcen vagzte and varied fx~lthe these lai~downersIn deer management projit incentive is causing many to give greater em- Perhaps the private landowner, cornpr15ing the largest phasis to deer. The majority of forest industries sector of landownership in the South, 1s the mo5t difficult welcome sportsrrlerl on their land to lzz~nf deer and to evaluate Their goals could possibly vary as widely other game. This improz)cs public relations But if as there arc numbers of landowners, therefore, I hesitate the industry is to provide improved facilities and to draw a def~niteconcluslon So far as I know, there quality clecr hunting the deer progl-an? must be pre- has never bcen a study of the prlvate landowiier to pored to pay its own way. really determine his goals in deer, or any other svildlife management on his lands. Before discussing "The goals of private forest holdings in deer management," I think that at the very outset we Because of this lack of documented information, I should establish exactly what lands we are talkir-rg about. must resort to personal observations. As most of us know, the southern forests account for 111my field of work, I have had an opportunity to work approximately 40 percent of our Nation's total forest with the private ownership sector for quite a number of acreage. Seventy-three percent of the South's forest years. From this experience, I believe it is safe to say lands is owned privately by farmers and other miscel- that the vast majority of private landowners would like laneous individuals; an additional 18 percent also is to have deer on their land, if for no other reason than the personal enjoyment and sportsmanship gained in owned privately, but by the forest products industry; and the remaining 9 percent is owned by various govern- deer hunting. Frequently, however, you will find that mental agencies. So, when we speak of private owner- rare breed of private landowner who is a truly dedicated ship, we are actually referring to two distinct groups- conservationist. He has practiced good deer management the individual landowner and the forest industry. on his land for many years, and in addition to the pleas- ure of hunting, he is a man who enjoys the sheer beauty For clarity in this discussion, I have labeled that group of the animals roaming his land. In some areas of the composed of farmers and other individuals as "private South, these early conservationists must be given at least ownership," and the forest industry group as "industrial partial credit for building up the deerherds that even- forest land." It is necessary to make this differentiation tually moved on to other nearby lands. because the goals each group of owners sets for its land are not necessarily the same. These goals arc frequently A factor that has become much more important in determined by a number of influencing factors. deer management on private lands in recent years is economics. As the demand for more and better hunting In private ownership, for instance, goals often have has increased with our rising population, many private bcen determined by the 5ize of the tracts, the purpose landowners have found it profitable to manage deer on for which the land was originally purchased, and the their land in addition to farming, cattle and tinibcr grow- land's actual capability to produce trees, agricultural ing. The practice is becoming increasingly popular, and crops, cattle or wlldlife inconie from hunting permits and fees for leasing have On the other hand, the forest industry generally had in some cases surpassed that earned in cattle or farming. a predetermined goal for the land at the time of ~ts This was borne out quite vividly last fall when I had acquisition--to grow as much timber as possible an occasion to read a Sunday edition of the Houston Over the years, however, the goals of each of these Post. On this particular day, there were 34 ads concern- groups are subject to reevaluation and change-usually ing paid hunting. They went something like this: deer because of economics. Large acreages, once considered hunting lease on 340 acres for $1,100.00 . . . deer hunting marginal farmland, for instance, have been returned $125.00 per gun per season or $20.00 a day with 3 day to productive forests. Economics simply dictated that minimum . . . and yet another at $300.00 per gun on the owners could earn a greater profit growing trees on 4,600 acres. This is what I mean why I say economics is the land than they could cotton or corn. Likewise, with fast becoming a factor in improving deer nianagemcnt on the increase in population and the growth of our cities, private lands. we have seen excellent forest lands become more valu- Another example of economics playing a major role in able as sites for shopping centers and manufacturing deer management on private lands that I ran across re- complexes. Still other fertile lands have been cleared cently appeared in "Outdoors U.S.A." In case you are and planted in agricultural crops. not familiar, this is a book published by thc United States Therefore, in discussing the goals of landowners, we Department of Agriculture. One of the articles tells are dealing with a dynamic, changing condition. Yester- of a Texas rancher who capitalized on a deerherd that was becoming a definite detriment to his cattle range. He the program that industry's role in deer management opened his ranch to hunters in 1961 and charged $10.00 thus far has not been so spectacular. But this is certainly a day. That first year produced 225 man-days of hunting not a true picture. One only has to look back to the with a kill of 100 bucks and 162 antlerless deer. Realiz- beginning of this program to realize the tremendous pro- ing he had a bonanza in the making, the rancher really gress that has been made in game management on forest began to practice deer management in earnest along industry lands. Of course, deer are a product of land with range improvement for his cattle. Now, this rancher management. We manage the environment which in charges $20.00 per day with a maximum limit of 20 turn reflects the land's deer-carrying capacity. Some of hunters each day, and the 1965 season provided 250 the accomplishments are well worth reviewing. man-days of hunting with a kill of 147 bucks rind 163 Perhaps one of the most significant developments oc- antlerless deer. The practice of paid hunting is not new curred when a number of companies sought out and in Texas. In fact, for a number of years Texas has been employed highly skilled wildlife mznagement personnel. the leading State in paid hunting. These specialists have been able to advise and guide their These are only two examples where private landown- companies in adopting timber management policies ers have found their goals of deer management changing which are compatible with wildlife habitat. as the economics became more attractive. In the good In the South, International Paper Company has em- old American tradition, the return of a profit has caused ployed seven foresters who hold graduate degrees in many private landowners to readjust their thinking on wildlife management to administer the company's game the merits of deer management on their lands. management program. Our organization consists of the When you look at the private ownership sector as a fish and wildlife director, who heads the program; a whole, it is difficult to say that they have a common specialist who conducts research projects at the com- goal in deer management. As I see it, deer management pany's forest research center; and one man in each of goals on these lands depend largely upon the economics our five woodlands regions who is responsible for activi- or the personal satisfaction derived from such a venture. ties in his particular area. Forest industry lands, however, are another matter. In actuality, many of the industry's forest management It appears a general policy and goal is beginning to take practices were excellent deer management practices also. shape. Of course, the first priority on these lands must Clearcutting is a prime example of a silvicultural prac- be oriented toward growing timber, but other factors tice that aids in improving browse for deer. Contrary have emerged in recent years. Game managcment can to what most people believe, the overmature forest does definitely be considered as one facet of this new policy. not provide ideal game habitat. Clearcutting opens the forest floor to sunlight, thus encouraging the sprouting Prior to the emergence of this new thinking among the of hardwoods and forb production. forest industry, I am afraid we must confess that very The trend today is to manage the southern forests on little, if any, consideration was given to deer manage- a much shorter rotation than has previously been the ment. But an increased outdoor oriented population case. This, too, can have a good effect on deer manage- with more leisure time brought out an entirely new ment. As the blocks of even-aged timber are established, aspect of industry-owned lands. the browse and other food plants will tend to diminish Originally, most of the industry's lands were opened as the stand matures, but nearby younger stands will to the public as an extension of our public relations and continue to provide food and cover. good neighbor policy. But, in reality, there was little On some industry-owned land across the South food improved hunting offered This proved adequate for plots and food strips have been planted. Whether this awhile and relations with our friends and neighbors went practice adds to the carrying capacity of the land is ques- quite well. But we soon came to realize, as the number tlonable, but it does tend to concentrate game which in of visitors to industry lands increased, just opening the turn provides for a higher hunter success. land for public use was not enough . . . they wanted more game to hunt. The trapping of deer is another program that has been carried out by at least one company to improve public It soon became obvious to the industry, if more game deer hunting. In this case, deer have been trapped in was to be provided, then some type of program had to high population areas and moved to areas of low popu- be developed, and perhaps more important, some type lations in an effort to establish larger and more vigorous of control had to be established if game management deerherds. were to be successful. Since I am more familiar with specific programs being Using the old "trial and error" method, we are now conducted by my company, let me take a moment to dis- developing these programs and controls. Since this is a cuss them. new vcnture for the industry, and there were no prior At International Paper, we have done considerable programs or case histories to follow, this method of inl- research in the area of fee hunting. Prior to launching plementation was necessary. We realize possibly some a fee hunting program on company lands, experiments mistakes will be made, but we hope that as we feel our were conducted at our Southlands Experiment Forest way through this period, the errors can be turned into near Bainbridge, Georgia. This research program, building blocks for a bigger and better program that will through questionnaires, surveys, and actual application benefit the most people. of organized controlled public deerhunts, proved to us The industry is now about 10 years into this new con- that the sportsman was willing to pay for hunting priv- cept. It may appear to those people not familiar with ileges. Personally, I am opposed to fee hunttng I thlnk it 1s the landowner must be in a position to exercise control neccssary, however, if we are to exercise any typc of over the land, the deerherd, and the deer hunter . . . and control over hunt~ngpressure on the land so that dcer fee hunting may be one of the solutions to this problem. management practlce5 can be successful We will always Another solution that in the long-run may be much have open land, I think, but more than llkely thrse lands simpler is for the landowner to lease parcels of land to wlll not be highly populated with game Also, I am various hunting clubs. Presently, this is quite common sure we will continue to Irase land to the various States and prices of the lease may vary from 10 cents to $3.00 for wildllfe management programs an acre, depending on the resident deer population and Since our early work at Southlands, we have expanded the demand for hunting. This typc of arrangement has our fee hunting system to some of our other lands in its advantages as well as its disadvantages, and in the Georgia, South Carolina, and Alabama We employ van- future we may find providing certain areas for fee lzunt- ous mcthods of conductii~gthe deer hunts, and fees ing and others for clubs to be the answer. charged may range from $2 00 per gun for a daily permit, At the present time, we have approximately 8 percent to $25 00 per ~nd~vidualfor a season permlt of our lands in the South leased to private hunting clubs. We believe fee hunting is certainly one of the most Another 7 percent is in State game management areas, promising advancements in deer management to date. and only 3 percent is being utilized as public fee hunting A program such as this will enable us to control the areas. hunting pressure and thereby carry out a more effective Thus far, we have found that concessions to deer can game management program. In addition, it will provide be made in timber management practices, but it remains some economic return for the time, effort, and expense to be seen how far these concessions can go. Again, it's the company invests in carrying out an intensive deer a matter of economics and control. As an example that management program. may be of interest to you, at I-P we have a research project underway in cooperation with Auburn University Two other companies in the South with which I am and the Alabama Department of Conservation. familiar have also embarked on a fee hunting system. This project consists of clearcutting 1,000 acres of The Chesapeake Corporation of Virginia charges $2.00 overflow bottom land with the objective of encouraging for a permit to hunt on 190,000 acres of land. Their a massive hardwood regeneration program in the form program is listed in the Directory of Private Hunting Lands in the Virginia Hunter's Guide, which is published of sprouts and seedlings. We anticipate this will provide an abundance of browse for an overstocked and, in some by the Virginia Commission of Game and Inland Fish- cases, starving deerherd. We believe that this study eries. In Alabama, Gulf States Paper Company also has will go a long way in determining the effects of bottom gone to a fee hunting system on their lands with charges land hardwood timber harvest on deer. Of course, we ranging from $1.00 to $10.00 for a season permit. also hope to learn what effect the deer are going to have At this stage in the ~ndustry'sdeveloping deer man- on hardwood reproduction. agement program, I believe the key to success depends In summary, the forest industry looks favorably on on deer hunting becoming economically profitable. Of deer management. It is perhaps the most important game course, the extent to whlch landowners go in dcer man- species on industry land and we desire to maintain a agement may well depend upon State trespass laws. If an healthy herd on all sizable tracts in keeping with the owner is to develop an effective deer management pro- primary objective of the land. For many years the ma- gram on his land, he must have control of the hunting jority of the forest industry has welcomed sportsmen pressure The States slmply cannot afford sufficient law on their lands to hunt deer and other game. This is part enforcement personnel, therefore, the ~nd~vidualland- of the industry's basic philosophy of being a good neigh- owners may be forced to employ game wardens bor and an accepted industrial citizen of the area. But Public pressures and demands have become too great if we are to provide improved facilities and qztality deer for us to continue deer management simply as a public hunting, the program must be prepared to pay its own relations gesture. And, typical of all industry, a program way. is difficult to launch on a wide-scale basis, especially one As I see it, the goals of private forest holdings in deer that will cost considerable money until it has proven management cover a wide range of individual and cor- capable of paying its own way. porate attitudes. The goals are influenced by a number Of course, as the program begins to pay its own way, of factors, ranging from personal desires . . . to public it becomes even more imperative that high quality hunt- demand . . . to economics . . . to the ease of implementing ing be provided. It's simply a case of when a sportsman the program. I do believe, however, we are on the right pays money for hunting privileges he must have some- track and the future of deer management on private thing to hunt. In order to provide such quality hunting, forest holdings indeed looks bright. Prevention and Control of Damage to Trees

D. 6. Denton, E. H. Hodil ' and D. H. Arner Department of Wildl~fe Management Mississippi State University State College, Mississippi

The prevention and control of deer damage to wai cxpcxr~mentcdwlth In 1964 on the Crown Zellerbach cottonwood trees in the Delta area of Mississippi was lands near Rolling Fork, Mississippi The fence was con- tried through the use of the following: firccracker structed with wooden stakes 4 fect long, 2 ~ncheswide, fences, odor and tuste repellents, buffer zones, and 'ind I inch thick These stakes were driven ~ntotht physical barriers. The ?nost promising of these tech- ground to a depth of about 1 foot and spaced at ~ntervals niques tested was a debris harrier constructed by of 20 feet Wire staplcs were driven into the stake5 windrowing tree debris around the area planted to approximately 6 inches from the top One M-80 fire- cottonwood. cracker was attached to each stake, on which was taped a detonator firecracker. A nylon string was strung from A great deal of work concerning the control of damage each firecracker and made just tight enough so that by deer to trees has been carried on In recent years In when a deer walked into the string, the firecracker both the United States and Europe Both chemical and would explode. The firecrackers were waterproofed with mechanical means have been utilized In attempts to household paraffin. The total cost of construction of 1 control deer damage Some of the chem~calrepellents mile of fence was $87.83. with certarn European workers have tested and reported However, this fence proved unsuccessful due to several effective are: factors First, ~t was difficult to waterproof the f~re- I. Vaseline, oil (paraffin), and either pyridone or crackers Second, even though the firecrackers were cresol in bentonite, waterproofed, the strlngs of the boobytrap type deton- 2. Anthropin (odor repellent 1, ators were found unreliable In three out of frve cases 3. Bone tar oil, In laboratory tests, the strings broke before enough pres- 4. Sticky paint which repels by its texture, sure could be applied to detonate the main firecracker 5. Cow dung and lime. Even if a reliable f~recrackercould have been made, Several of these repellents were tcstect by other work- the single strand fence would not have been effective ers, most of them in the United States, and wcre found in for an extended period of time Observations made on these instances to be ineffective. a 12-acre field surrounded by the firecracker fence re- The studies reported In this paper are concerned with vealed that deer continued to enter the held by either the protection of cottonwood IPO~ILI?LSdellozdesi planta- leap~ngthe fence or slid~ngunder it In order to prevent tlons in the Mississippi Delta area from deer damage deer from sliding under or leaplng over the fence, a more Cottonwood, within the past few years, has become a elaborate modified verslon of the fence was constructed very important pulpwood species and is now being grown around a 1 acre portion of cottonwood plantation This by most of the larger paper and timber companies with fence was made from the same type of material as the lands located in the Mississippi Delta region. first one. However, this fence was constructed by driv- Some of the greatest concentrations of white-tail deer ing two rows of stakes, each stake spaced 20 feet apart, (Odocoile?rs virginianus) in Mississippi live in the forest with the second row of stakes spaced 10 feet behind the bordering the Mississippi River. During the early spring first row of stakes. Two strings of firecrackers were months, when the nutritious and palatable cottonwood attached to the front row of stakes nearest the plot, one cuttings begin to leaf out, deer browse cxtenslvcly on string at the top of the stake, and the other string at the the new cottonwood plant~ngs The terminal \hoots are bottom of the stake. The back row of stakes (tllose especially vulnerable, and very frequently the young farthest from the plot) had only one string of firecrackers cottonwood is deformed or killed by excessike browsing at the top. Firecrackers were also strung between the In 1964, cooperative research was initiated with the rows in a zigzag fashion from each stake. This structurc was observed to be successful in repelling three deer. USDA Forest Service, Southern Hardwoods Laboratory However, this success might have been due to the form- at the Southern Forest Experiment Station, Stoneville, idable aspect of the structure, and not the firecrackers, Mississippi, and the Department of Wildlife Management since it was observed that the deer stopped short of the at Mississippi State University. fence and did not attempt to enter the enclosed area. The techniques which were tried or investigated were This structurc was abandoned due to its impracticality. flrecracker fences, taste and odor chemical repellents, buffer zones, and barrier devices The results and eval- uation of this investigation are as follows: CHEMICAL REPELLENTS FIRECRACKER FENCES Seven different taste repellents and one odor repellent wcre t~eldtested on cottonwood cuttings in a small bloclc A f~recrackerfence, which followed the design of a design The repellents tested were fence used in Maryland by Flyger and Thoerig tI9611, Pcnnsalt th~ramS-42 Mr. IIodil is now on thr staff of the Marylar~tlGnmc and Inland Fish Commission. Peiinsalt thirain 75 Pcnnsalt thiram 80 iprayer Total cost per acre was $12 16 Deer clamdgc Pennsalt thiram Animal Repelleiit with this treatment was reduced 13 8 percent Penns,ill OMPA Syitemic Repellent 011 plot 4, strips of burlap were soaked in bone ta. Du Pont Ardian 42-5 oil odor repcllent and attached to stakcs spaccd 12 fee apart surrounding approximately 5 acres of cottonwoot Morton Chemical Co~np,inyZIP cuttings. The burlap was treater! twice at 1 month ii~tcr Bone tar oil odor rclpellcnt iMagic Circle) vals. A 4:l dilution with water was used and treatmen Thrie repellents wcre applled so that nii~etreLitrnelits commenced in early March. The total cost per acrc wa: were arranged in two latin quare tiesigni I.:,~ch cheml- $4.15. Ilecr cia~nagewas reduced 24.4 percent. cal appearcld oncc in each row ,ind oilce in cach column In 1967, the same techniclue was used, but trcatec in order to allow for nonrandom movement of d(bcxr The it'tkes wcre scattered tl~roughoutthe field ai well a< inelhod of application was by compreiiion sprayer ex- on the periphery This study revealed no significant drf cept for the systemic repellent m~hlch u'ii applied by fercnce bctwecn the treated and untreated plot? soil injector Oil plot 5, a '2 inch cotton rope was soaked 111 bonc Of the eight repellents tested in this experiment, two tar 011 octor repellent and attachcd to stakes surrounding ihowcrl promise as a posslble effective deer browse de- the periphery of approximately 5 acres of cottonwoot terrent Those were the systemlc repcllent IOMPA)'ilici cuttings At intervals of 15 tcct, soaked burlap strip' the odor rcpellcnt (Magic Circle bone tar oilr On the uJcrc attached to the rope The burlap strips were treatec basis of the percentage of plants browsed and unbrowscd, twice at a 1-month interval, commencing in early March ther~was a significant difference between the percent- A 4:1 dilution with water was used. The total cost pel age of plants browsed on the untreated plots and the acre was 815.92. Deer damage was reduced 40.9 percent plants browsed on the treated plots However, there In another experiment conductcd in the same areL was no significant difference between repellents during 1967, a two strand fence of heavy tw~newas con The taste repellents werc found to be ineffect~vcin structed around the perimeter of a 4-acre fleld of cotton controlling deer damage due to the rapid growth of the wood seedhngs The two strands were placed 18 and 41 cottonwood seedlings, which wlll grow from 6 to 10 inches above the ground Burlap rags soaked in odor feet in a slngle growlng season. repellent were tled at 3-yard Intervals along the fence On the basls of the results obtained from the sm,ill The rags were treated a total of 3 tlmes at month11 plot study, the bone tar odor repellent and the OMPA lntervals beglnnlng in mld-March Total cost of fence systemlc repellent were selected for further testing in rags, rcpcllents, and labor was $12 50 per acre Rrowsc largc field blocks damage reduction ranged from 12 to 53 percent durlng the 4-month testing perlod (table 1) One blg advantagt Systemic Repellent to this type of fencing is that trees and bushes arounc the edge of the f~eldcan serve as fence posts Cost ot In 1965, Dennis Jordan, a graduate student in wildlife the fence could be reduced by uslng less expensive rag: management at Mississippi State University, completed and by dlluting the repellent to half strength further experimentation with odor and systemic repel- lents. The deer repellents were applied to six experimen- Table l -A comparzson of dew damage to cottonwood cu tal plots located in a cottonwood plantation which had ttl~gszn a rope fence plot treated wtth odor repellel been planted the previous winter. On plot 1, the systemic and a control area repellent OMPA was applied to approximately 2X2 acres of cottonwood cuttings. A soil injector was used to apply Treatment Exam~nation Trees damaged by deer 2 ml. of repellent to each cutting. The injection into the at j dftc - R~p~fi;tr~ - - 1 ------ground was made approximately 2 inches from cach Percent Percent cutting. Only one treatment was made, and it was ap- plied in early March, 1965, when the cuttings first began March 10, 1966 March 24, 1966 11 5 1 to leaf out. Total cost per acre for this treatment was March 15, 1967 April 1, 1967 11 44 $14.15. Deer browse damage was reduced 20.7 percent. April 1, 1967 Aprll 23, 1967 40 52 On plot 2, approxi~nately21,5 acres of cottonwoocl cut- Auril 23. 1967 June 17, 1967 35 86 tings were treated with two applications of OMPA inade at 1 month intervals. The method, time of application, On plot 6 bone tar oil odor repellent was placed in a anti amount of injection were the same as in plot 1. The small circle around each cottonwood seedling on an sccond applicatiorl was made in early April. The cost area of approximately 5 acres. The pressure sprayer per acre was $27.20. Deer damage was reduced by 27.0 was used to apply the 4:1 dilution twice at 1-month in- percent. tervals, commencing in early March. Total cost per acrc was $3.92. Deer damage was reduced 20.4 percent. Odor Repellents Approximately 5 acres of cottonwood cuttings were On plot 3, the bone tar oil odor repcllent was applied used as a check plot so that the intensity of deer brows€ 011 the ground around the periphery of approximately on the treated areas could be compared to a norm. 5 acres of cottonwood cuttings. Two applications were A second type of odor repellent tested in 1967 was made monthly for a total of four applications, comment- animal tankage. Tankage is a residue of animal tissue ing in early March, 1964. A 4:1 mixture in water was (50 percent protein, 5 percent fat, and 8 percent crude applied to the ground with a garden type pressure fiber and unknown material) and is used in some part: of the country as a feed supplement for hogs. Tankage cotto~lwoods browsed extensively by deer and those was placed in small bags on posts spaced approximately leaves from cottonwoods showing relatively little brows- 15 feet apart. Approximately 4 acres of cottonwood ing damage was made, along with analysis of soil samplcs were encircled by posts containing bags of animal tank- taken from around the browsed and unbrowsed cotton- age. woods. The results showed no difference in the chemical Deer damage was noted in 34 percent of the cotton- analysis of the soil between the browsed and unbrowsed wood trees within the treated plot, and 44 percent of the plants. No significant differences were found in the trees in the control plot were damaged by deer. This chemical analysis of the leaves, with the exception of small reduction in deer damage did not justify the ex- mineral content. The mineral content was higher in the pense involved in this technique. leaves of the browsed plants than in those of the un- From these experiments it was concluded that bone browsed plants during both years that these analyses tar oil odor repellent applied to rope fence device was were made. effective and more economical in reducing deer damage In the spring of 1967, a field test was devised to deter- than OMPA or animal tankage. OMPA was costly for mine if deer damage to cottonwood plantations could be the results obtained, and was also discouraging because reduced by aerial application of odor repellent when of its extreme toxicity. supplemented by fertilized feeding areas. A large cotton- wood plantation near Fitler, Mississippi, was the location AERIAL APPLICATION OF REPELLENTS for this experiment. Four 10-acre fields, located :"1 mile apart, were treated by spray plane with odor repellent In the spring of 1966 a new technique was tested. Be- from a height of 20 feet. Four adjacent 10-acre fields cause of the apparent effectiveness of bone tar oil odor served as control plots. A total of three treatments were repellent as a deer browse deterrent, it was decided that made at approximately monthly intervals beginning in further tests with this chemical were warranted. How- mid-April. Two parts by volume of bone tar oil odor ever, due to the rising costs of labor and the difficulties repellent were mixed with six parts water. This mixture involved in hand treatment of large cottonwood planta- was applied at the rate of 2 gallons per acre. Two 1-acre tions, it was thought that airplane spraying of the repel- fertilized buffer zones planted to winter wheat and clover lent might offer a more efficient and economical means were established approximately 150 yards from two of of application. A 27-acre field of cottonwood seedlings the treated fields. One-acre buffer zones of fertilized located on Diamond Point Island near Vicksburg was native vegetation were established approximately 150 selected for the experiment. Thirteen acres were treated yards from the two remaining treated fields. by agricultural spray plane, and the remaining portion of the field was reserved as a control. Applications were The two buffer zones of native vegetations, which con- made from an altitude of 100 feet One part bone tar sisted largely of blackberry (Rzibus spp.), Ladies-ear- oil odor repellent was mixed wlth seven parts of water drops (Brunnichia cirrhosa), trumpet creeper (Campsis by volume and applied at the rate of 8 quarts mlxture radicansi, and greenbrier (Smilax sp. ), were fertilized per acre. The repellent was first applied in late Febru- with 1,000 pounds of basic slag, 200 pounds of nitrate of ary, before the young cottonwoods began to leaf out. soda. and 200 pounds of ammonia nitrate, along with ES Four more applications followed at varying intervals, Min El trace mineral group (per acre basis). depending on the degree of browse damage. The dates One hundred seedlings on each of the treated and con- of application were: February 26, March 19, April 9, trol plots were tagged for deer browse evaluation. In May 16, and May 20, 1966. The total cost of repellent addition, 25 trees on each of the four aerially treated plots and aerial application was $2.40 per acre per application. were tagged and received four hand sprayer applied This method of treatment resulted in an average browse treatments of bone tar oil odor repellent at 3-week inter- reduction of 11 percent for each 3-week lntcrval during vals. the testlng period Survival of the seedlings on the treated A check area consisting of 130 cottonwood trees, lo- plot was 20 percent greater than on the control plot cated $1 mile distant from any buffer zone or treated area, was examined periodically for deer damage. BUFFER ZONES Periodic examination of the planted buffer zones Linder et al. in Germany ( 1956) believe that browsing showed that deer utilization of these two wheat and and bark peeling by red and roe deer are largely due clover plots was heavy during late winter and early to their physiological need for trace minerals and nutri- spring. The two buffer zones of fertilized native vcgeta- ents which they are unable to obtain from the native tion were used by deer as soon as the plants began to foods growing in the acid forest soils which arc low leaf out in early spring. Heavy deer utilization of these in potash and phosphorus. These men have tried winter two plots continued into the spring months. feeding with hay and fodder fortified with trace min- Deer damage to the cottonwood seedlings involved erals. They claim that peeling and browsing damage in the experiment was evaluated on April 1, 1967. This was reduced within tolerable limits in an area with 8.5 evaluation was conducted before any repellent was ap- deer per 100 hectares. plied. It was found that deer damage to seedlings planted The browsing of the terminal shoot of forcst crop trees in fields located near buffer zones was only 2 percent was reported by Hauer (1953) in Hungary as a sign of less than to those located '5 mile distant from any buffer green fodder shortage. He suggested the raising of frost- zone. This small reduction in deer damage to the you~lg hardy greens on game pastures ;IS one possible solution cottonwoods did not justify the expense involved in to reduce darnage. Chernical analysis of the leaves from establishing these supplemental feeding areas. Evaluation of deer damage on the treated and control trees showed that 6 percent of the cottonwoods inside arcai at 3-week interv~lsthrough th.2 sprlng and early the fenced plot were damaged by deer, while 70 percenl summer months showed that decr damage was reduced of the cottonwood seedlings on an adjacent control plot an average of 18 percent on the four ,ier~allytrcated wtre damaged by browsing deer. The chief advantage: plots in this type of fencing appear to be its comparatively low IIand treatment with odor repellent of 125 selected cost and high degree of protection. Much of the material trec5 on the aerially treated plots afforded no addit~onal such as the chicken wire, can be used for several years protection aga~nstbrows~ng deer Height measurements This would greatly reduce costs over a period of several on a total of 800 cottonwood seedhngs located on the years. treated and control plots showed that there wai no appre- c~abledifference In height.- .,growth Thew measurements Debris Barriers were made near the end of the first growing season. J. S. McKnight of the Southern Forest Experiment In general, the results of this experiment indicate that Station at Stoneville, Mississippi, L. C. White of Chicagc I-acre supplemental food plots are of little value in re- Mill & Lumber Company, and W. W. Dannenburg of ducing deer damage on cottonwood plantations. When the U. S. Gypsum Company, experimented with a barrier the food plots were tested in combination with the aeri- constructed of the accumulated debris resulting from ally applied odor repellent, an average browse reduction clearing operations in the area. The debris was pushed of 18 percent occurred for each 3-week period during the into windrows, completely encircled the area with the spring and early summer months. Total cost for treating exception of an opening large enough to permit tht I-acre of cottonwood seedlings with an aerial application entrance of tractors and cultivatirig equipment. The of Magic Circle odor repellent was $4.05 per application windrows were at least 20 feet wide at the base, and 10 ($2.80 for repellent and $1.25 for spray pl;ine application). feet high. The entrance to the field was covered with a hanging screen of burlap. The areas protected by bar- PHYSICAL BARRIERS rler fences, which were constructed with felled timber Wire Guards and treetops p~ledflush with the ground, were not pene- Wire exclosures constructed around individual trees trated by deer Some deer were able to enter where a were also tested as a possible technique for protecting great deal of earth was mounded up w~ththe felled cottonwood seedlings from browsing deer. Five foot timber debris. The technique where the debris was piled tall wire exclosures were constructed from 2-inch mesh flush with the ground shows excellent promise, and addi- chicken wire. These exclosures were cylinder shaped tional field studies are needed with this barrier type with diameters varying from 12 to 18 inches. The wire fence. exclosures were placed around individual trees and staked to the ground. This control technique was 100 OTHER TECHNIQUES percent effective, but the high cost ($80.00 per acre) Foresters for the R. F. Learned & Sons Company of makes it impractical for use on large cottonwood planta- Natchez, Mississippi, utilized cotton~voodswitches grown tions. in a nursery for 1 year for planting stock. These cotton- wood switches averaged 12 feet in height and were at Fence least :!/? inch in diameter. A tractor with a posthole dig- One of the most effective types of control tested was a ger was used to dig holes about 12 inches in diameter temporary fence. The fence was constructed of wooden and 4 feet deep. A posthole digger can dig about 500 stakes, 2-inch mesh chicken wire, binder twine, staples, holes per day. The cost involved in digging the holes nails, and bone tar oil. Stakes 6 feet long, 1 inch wide, and planting averaged $51.53 per acre. This technique and 1 inch thick were driven into the ground to a depth was very effective in reducing deer damage, but the of about 1 foot at 10-foot intervals around a 4-acre cotton- cost is excessive. wood plantation. Five foot tall 2-inch mesh chicken wire was stretched by hand as tightly as possible and SUMMARY AND CONCLUSIONS stapled to each post with a high compression staple gun. Alternate 6-foot stakes were braced on opposite sides The results of f~eldtesting a number of repellents and with binder twine, and 155-foot stakes driven 12 inches devlces for controlling deer damage to cottonwood planta- into the ground. A 2-foot long, 1-inch wide, and 1-inch tions In the M~ssissippiDelta arca revealed the follow~ng thick stake was nailed to the top of each 6-foot post. (1) Firecrackers were difficult to waterproof and were These stakes pointed toward the outside of the fence and not dependable 12) Taste repellents proved economically ~lpwardat a 45'' angle. These stalres served as a11 out- ~neffcctivedue largely to the rapid growth of the cotton rigger type device and supported three strands of binder wood (3) Bone tar 011 odor repellent applied on hurla~ twine spaced 6 inches apart. Four feet from the fence strips, wh~chwere hung from a rope fence, proved to bc on the side away frorn the field, 5-foot stakes were driven the most effect~vemethod of the chemlcal repellents 6 inches into the ground at 50-foot intervals. Binder (4) Bone tar oil applied by airplane was ~neffective (5 twine soaked in bone tar oiI repellent was stretched tight The e\tablishn~entof b~lfferLone5 of highly icrl~ll~ec and stapled to the top of each of the 5-foot stakes. The native vegetat~on,as well a5 clover plots, proved ineftec total cost of materials and construction of this fence was tivc in controlling deer damage (6) The most economi 6 cents per foot. Two man-hours of labor was required cally cffcct~vetechn~que teited was the temporary typc for maintenance of the 1.800-foot fence during the 4 fence (7) The witidrowing of debr~sappears to hdvc month testing period. Data based on a sample of 1,000 s~gnlfi~antlyredut.ect deer damage in the Delta area The variance of the reyults obtained with the sarne LITERATURE CITED tcchn~quefrom 1 year to another year, or from one area Fiyger, V., and Thoerig, T. to another area, very probably was the result 01 ditfcr- 1961. Preliminary report on a new principle for pre- ences In deer population Some ycars docs would be vention of crop damage by deer. Southeast. harvested, and a significant reduction in the deer popu- Assoc. Game and Fish Comin. 15th Ant-iu. ConC. latlon would result It is the belief of the writcr~that Proc. 1961: 119-122. 110 chemical repellent will provide effective protection from deer damage to such palatable \peclc.s as cottonwood I-Iauer, L. when the deer pol?ulatlon exceeds the carrying capacity 1953. Game fecdjng with ensiled leaf fodder. Erdt(sz. of the range As of thic, date, th- windrow technique Tud. Int. Evk. 20: 2-12. and the temporary fence appear to be the mo5t econorn- IJintincr, A., Brandl, M., and Wyler, E. ically effcctlve technique to control deer ci,~m,ige to cot- 1956. [New methods to prevent gaine damage. J Allg. tonwoods In the Dclta area Forstztg. 67: 233-237. Measuring Habitat Productivity Richard F. Harlow Wildl~feBiologist Southeastern Forest Experiment Station Forest Scrv~ce,U S. Department of Agriculture Blacksburg, Virginia James B. Whelan Assistant Unit Leader Virginia Cooperative Wildlife Research Unit Blacksburg, V~rginia

This pccpcr rlescribes ~~ossiblcccppr.ouc!zc?s to esti- Following is a discussion of scvcral techniques fr mating habitat productivity in the South based on evaluating forage quality: qualitative and qxuntilative measurements. Meth- Animal production.-Animal growth and reproductio

ods are sugqcsted for- determining quality of forages permit an estimate of the gross nutritive quality ( using modifications of currenl forage evuluation natural forages. However, as pointed out by Reid ( 1962 techniques for estimating ntitritive value. Also dis- animal production provides only a subjective measur cussed are methods and difficulties of measuring of forage quality. But in conjunction with forage prc forage yields where all facets of the ur~derstoryhabi- duction and utilization measurements it has been suc tat are considered potentictl deer food. cessfully used by researchers working with domesti Numerous methods have been devised to evaluate range stock (Frischknecht and Harris 1968) and wit white-tailed deer habitat, but no particular technique deer (Gill 1956; Adams 1960). appears adequate to estimate habitat productivity in the Chernical composition.-A method of forage evaluatio South. Highly diverse habitats, ranging from thc Coastal which has received considerable attention is the detet Plain to the southern Appalachians, exhibit vastly dif- mination of chemical (nutrient) constituents of forag ferent levels of productivity, depending on such factors and their relation to nutritive value. The tradition; as weather, soils, and plant species combinations. And laboratory technique for determining the chemical cor there is only a limited understanding of nutritional re- stituents of forages is the Weende system of proximat quirements and food preferences of deer in the various analysis, described by Maynard and Loosli ( 1962). Tw physiographic regions. Also, the highly variable form, of the fractions obtained by this procedure, nitroger distribution, and seasonal availability of food items with- free-extract (NFE) and crude fiber (CF), are subjec in habitats present complex sampling problems. to considerable error. Although the NFE is relative1 Objectives in measuring habitat include relating qual- digestible, it contains highly undigestible component ity and quantity of forage to the productivity of a habitat. like lignin. The CF is considered to be relatively undj Measurements of forage quantity and quality are neces- gestible; however, it includes cellulose which rume sary prerequisites to determination of habitat produc- microbes digest readily. tivity. Forages often vary widely in their chemical compc This paper describes habitat analysis techniques (both sition (I-Iagen 1953; Harlow and Jones 1965; Wilson 1969 qualitative and quantitative) used in the past and sug- These variations, which occur even between plants a gests some realistic approaches that may be used in the the same species (Short et al. 1966), result from man future. factors, including season, growth-stage, soil type, physi ographic and climatic differences, soil moisture, and lan' QUALITATIVE MEASUREMENTS management practices (Lay 1957). Evaluation of Forage Quality In South Carolina, areas which had the highest value The limiting factor affecting nutritive value of forages for crude protein and phosphorus in certain brows is normally the content of the digestible or apparently plants also had the highest values for average bod available energy. Forage consumed by animals can mcet weight of deer (Thorsland 1967). Conversely, two area their growth requirements only if forage quality provides which had the lowest values for these two plant nutrient the necessary nutrients in adequate amounts; therefore, had lower average body weights for deer. a knowledge of forage quality is desirable. Factors deter- Using data obtained from digestion trials with domec mining nutritive value of forages ~~suallyinclude energy, tic animals, Schneider et al. (1952) developed regressio protein, phosphorus, and Vitamin A. Animal require- equations for estimating total digestible nutrients a ments for these and other nutrients may change when digestibility of any specific nutrient from chemical con meeting the demands for growth, fattening, breeding, position of forages. Baumgardt et al. (1962) mentione or general maintenance. that the correlations associated with those regression A knowledge of the nutritional requirements of deer have not been satisfactory because the wide variation is basic to evaluating forage quality in terms of meeting in amounts and proportions of several nutrients in fol the animal's daily requirements. Information on the ages affect their digestibility and in turn their nutritiv daily nutritional requirements of deer is limited (French value. Other studies (Hart et al. 1932; Hellmers 194 et al. 1955; McEwen et al. 1957; Murphy and Coates 1966; Swift 1948; Weir and Torell 1959; Short et al. 1966) ir Silver 1968; Wetzel 1968; Cowan, personal communiea- dicate that considerable variation in chemical compos tion.' tion occurs between plant species growing on the sam -.. .. -- ' Dr. irnal Nulrition, Pcnn. State Univ., range. Some of these investigators have suggested thk University Park, Penn. 1969. deer select the most nutritious forages; there appeal to be ample evidence supporting this hypothesis. Thus, more than concent.ratiotis of cellulose or hemicellulose in it seems doubtful whether an investigator could select the cell walls. A rcgrcssion equation to predict the range forage having approximately the same nutritive digestibility of organic matter was developed using lig- composition as that which a deer might sclcct. nin, neutral detergent residue, and uronic acid as inde- Edlefsen et al. (1960) comparcd the nutrient cornpo- pendent variables. sition of hand-plucked range plants to that of similar In uitr-o digc~stibilit?~of forcrgt.--'rkle energy v;ilue of plants obtained from esophageal fistulae on . Al- a forage has been shown to be closely related to the t.hough significant differences were found between digestibility of its dry matter or orgarlic matter (Baum- samples, the relative magnitude was not great. gardt et nl. 1962 ). In addition, Baumgardt et c~l.i 1962) It might be possible to select manually range forages mentioned that although the digestible dry matter (high- which closely resemble, in chemical composition, those ly correlated with digestible energy i is normally the eaten by deer. By following a tame deer fitted with a limiting nutritive factor, it is also important to know harness and leash, one could observe its feeding habits the digestibility of the forage protein. Fortunately, these and select similar forages from the same location. IIealy researchers have shown that forage crude protein is (1967 J conducted a forage preference study using deer highly correlated with digestible crude protein. For on leashes; however, forages were not chemically an- routine analyses the prediction of digestibility of forage alyzed. Van Dyne i 1968) made such a comparison in dry matter, using laboratory techniques, is a valuable developing a technique for predicting relative chemical and time-saving procedure for evaluating forage quality. composition of dietary botanical components obtained Johnson and Dehority ( 1968 ) compared several chem- from esophageal fistulae of grazing livestock. Validity ical and in vitro techniques used to predict digestibility of the predicted values was tested by comparing them and intake of forages. The in vitro and chemical data to values obtained for hand-clipped samples from the were compared to in vivo dry matter and energy diges- same range. The predicted chemical composition of tibility, relative intake, and nutritive value index grazed plants was found to be reasonably close to that (Crampton et nl. 1960 1. These workers found that the of hand-clipped plants. This approach, using nonlirlear two-stage digestion procedure of Tilley and Terry ( 1963 ) programming and matrix methods of analysis, also pro- was the best method for estimating dry matter digestibil- vides a reasonable estimate of the digestibility of cellu- ity (DMD ), which agrees with comparisons conducted by lose. It offers a means whereby a great amount of infor- Oh et al. ( 1966). Van Soest et nl. ( 1966) suggested that mation on botanical and chemical composition of the diet the two-stage method could be improved by replacing can be obtained with a reasonable degree of accuracy the second stage (acid pepsin digestion) with a deter- in a relatively short time. mination for cell-wall constituents using the neutral A promlslng techn~quewas developed by Van Soest detergent procedure, which shortens the unmodified (1963) and Van Soest and Wme (1967) to slmpllfy feed two-stage method by nearly 2 days. cvaluatlons The ratlollale for the method 1s that cell Grlmes 11968) was the flrst to compare the relat~ve contents of plants are dlgc,sted almost completely 198 digest~vecapab~llt~es of rumen-flstulated deer and sheep percent) by rummants but plant cell walls are only and an in t)iuo nylon bag microdigestion technique de- slightly digested. This method involves treating (digest- veloped by Lusk et al. (1962 1. He concluded that sheep ing) a feed sample with neutral detergents and yields could be used for determining quality of forages utilized two fractions-the cell contents composed of readily di- by deer. Cowan et al. ( 1969) discussed the applicability gestible compounds, and the cell-wall components con- of the techniques used by Grimes. sisting of fiber insoluble in neutral detergent, called Rumen nr~alyses.-Short ( 1963 ) compared VFA pro- neutral detergent fiber (NDF). However, a portion of duction in white-tailed deer and a fistulated steer. He the cell-wall constituents, represented by the hemieellu- found that mean co~lcentrationsof VFA in rumen liquor loses and fiber-bound protein, is soluble in acid deter- samples were similar for deer and cattle when both gent; the remaining insoluble residue is called acid deter- were fed a concentrate ration. When both were fed aspen gent fiber (ADF). The ADF is composed primarily of and white cedar, deer maintained a higher level of VFA lignin and cellulose ilignocellulose). A value for total production. Ullrey et al. ( 1964, 1967, 1968 1 found VFA digestibility of forage based on the Van Soest technique production in deer fed white cedar exceeded that for probably provides a better estimate of true forage quality deer fed aspen, jack pine, and balsam fir. than the proximate analysis value. Shore and Harretl Rumen contents collected during the summer from (1969) compared the standard proximate analysis and Sltka deer were analyzed by Kle~n(1962) to prov~dean the detergent solution analysis of two southern browse Index to the nutrit~vequal~ty of forage on two Alaskan species. Significant differences betweell the methods Islands Poslt~vecorrelat~ons were mdlcated between were evident, as indicated by the different values ob- percentage nltrogen and percentage volume of mlero- tained for various components in current twigs and old organisms (r =-I t0.67) and percentage nitrogen and twigs. light trausmittancy of the clear fraction (r - -{0.75). Another new technique of chemical evaluation of for- Comparisons of rumen samples from the two islands ages was developed by Gaillard (1968) and is based on revealed that differences were significant for most chem- composition of cell-wall constituents of plants. Labora- ical analyses. Klein's study also suggested that fiber tory digestibility of roughage samples was compared to content of fecal samples might be used as an indicator known in vivo digestibility of the roughages. He con- of forage quality. cluded that lignin and uronie acid concentrations from Kirkpatrick et al. (1969) studied seasonal changes in cell walls influenced the digestibility of organic matter proximate composition of rumen as related to forages con5umed by white-talled deer In the southeast They In vitro fermentation trials have shown promise f found that mean crude proteln levels of rumen contents evaluating forage intake (Crampton 1957; Crampton var~edfrom 26 perccrit 111 spring and summer to 14 per- al. 1960; Donefer et al. 1960; Barnes 1966). The hyp cent in the fall thesis is that the speed at which a particular forage Another techmque whlch would provide a good index digested in short periods (6 to 12 hours J determines ar to forage qual~tymeasures the relationsh~pbetween the rnal intake of that forage. Confirmation of this the01 nitrogen (crude protein) in forages and the extent of its for determining intake would be especially beneficial conversion into rurr~enmicrobial nitrogen. Weller et al. wildlife studies. (1958, 1962) studied digestion in sheep and found that Possible approaches to determining forage quality.. the largest loss of plant nitrogen within the rumen was Figures 1, 2, and 3 outline possible approaches to dete attributed to its conversion into microbial nitrogen. mining forage quality. These are only suggestions bast Unfortunately, the relatively long and detailed analyses on modifications of current techniques, some of whic required preclude use of this procedure as a routine have already been discussed. Caution is needed in inte analysis for determining forage quality. preting data; the methods should be used with reserv Foragc intake.-Foragc intake is of maximum im- tion. portance when evaluating qualitative habitat produe- tivity. Therefore, the chemical and botanical composi- In the first approach, figure 1, an index to nutritix tion of the diet must be sampled carefully to prevent value is derived from an expression of dry matter intab error in evaluations. Sampling may be done with both and dry matter digestibility in relation to metabolic sii animals and plants. of the animal. This approach to estimating the nutriti~ value of forages was developed by Crampton et a Estimating amount of forage consumed includes the (1960). The nutritive value index iNVI) indicates tk ratio technique which involves the measurement of effective feed value of forages by combining a measul an undigestible indicator like lignin in the forage and of the quantity of forage dry matter voluntarily cox the feces (Cook 19561, and the fecal nitrogen index sumed per day and the percentage of digestible energ method (Lancaster 1949; Arnold and Dudzinski 1967 ) . in the forage into its calculation. It is necessary to relal Merits of these methods are discussed by Van Dyne and the quantity of forage consumed daily to the daily COI Meyer ( 1964), who developed a promising new method sumption of a standard or reference forage; thus obtair for estimating forage intake of grazing livestock by using ing a measure of relative intake (RI). This standar microdigestion techniques. Van Dyne's system can only value is derived from feeding a standard forage and dc be used, however, if adequate facilities are available for termining the average dry matter consumption per un maintaining relatively large numbers of deer fitted with of metabolic size (body weight in kilograms raised to tk esophageal and rumen fistulae.

(1) (2) r r.4ME DEER MANUAL SELECTION 1-ON LEASH - (3) (4) I SPECIES COMPOSITION OF PREPARATION OF SAMPLE + DIET AND ',( BY WEIGHT FOR DMD DETERMINATION I -OK- OF COlMPONENTS (SPARKS & (TILLEY AND TERRY I MALECHEK 1968) 1963, VAN SOEST 1966) 1 FESOPHAGEAL COLLECTION OF FISTULATED -FORAGE MIXTURE / STANDARD INOCULA FROM 'DEER ON LEAS13 FROM FISTULA FISTULATED DEER /I-' - - _ 11 -OR- (5) I /D~YORAL ADMINISTRATION IZUMEN INOCULA FROM A DEER ,IN VITRO L4 OF CrLO$ KILLED ON STUDY AREA 48-HR DMD ------

(6) ESTIMATION OF TOTAL DAILY FECAL DETERMINATION OF DM DM1 DM OUTPUT IN FECES X 10 OUTPUT FROM TAME OR FISTULATED- IN FECAL OUTPUT .----rc (INTAKE) = ------100- ',4 DIGESTIBILITY DEER BY GRAB SAMPLES (REID 1962) - -- OF DM (RUMEN)

(RI) x (DMD) NVI (NUTRITIVEVALUE INDEX; LUCAS 1962)"

.. -. .. .- * Modification of Nutritive Value Index (Cra~nptonef al. ISGO), assuming a high correlation b~~ec~nDMD anddigestible cnerg, DMIIday (sample forage) Lcgrnd DMD Dry Mattel Dlgesl~bllity,DM1 = Dry Matter Intakc, DM = Dry Matter, RI - - 1( (~tk,.~')(DMIIday of std. forage1- .75) kg Figure 1.-Estimation of nutritive value for a mixed-forage diet of deer using percentage fecal DM and daily output of feces to predict forage intake. 3'1 power) In deternillnng the standard or reference a runien fist~llatecldeer calibrated to the study area could forage to be used for whlte-talled deer which-ln thelr serve as the source of inoculant. This would standardize natural habitats-exist on a mixed-forage diet, it would the inocula used for all in ~ritro trials and would provide be necessary to select a forage or forage mixture which a somnd basis for comparing dry matter digestibility of provides the quantities of nutrients for meeting nutri- forages collected from several areas. Another possibility tional requirements when fed c~dlibitz~m. for standardizing inocula for in vilro trials would be to After obtaining a value for RI of a forage or ~nixture use rumcn fluid from a fistulated deer, or possibly a of foragcs, a value for its digestible energy is deter- fistulated sheep, which was fed a standard ration. Diir- mined; multiplyilig these two values gives tlle NVI. ing each iia vitro trial, a sample of the st:incl;rrd r:~tior~ Since Raumgardt et (11. ( 1962 ) have shown that digesti- also should be included for microdigestion to perrliit ad- bility of forage dry matter is highly correlated with di- justme~lt icorrection) of digestibility values of range gestible energy, dry matter digestibility could be srtb- foragcs in relation to possible changes in digestibility stitutect for digestible energy in calculati~ig the NVI. of the standard ration. The theory, methods of calculation, and application of An estimate of daily dry matter intake can be obtained the NVI lor forages is discussed by Crampton and I-larris from the ratio of daily dry matter output in feces to ( 1969). indigestibility of rumen dry matter, as shown in step 6. As ~llu\tratedin steps 1 alld 2, the bota~iicaland chem- A modified nutritive value index INVI) may be c;rlcu- ical composition oi torages collected by c>soph,~gcalflstu- latcd by cnultiplying dry matter intake (relative intake) lac and by manual selection should be compared If ;: dry matter digestibility (step 7). significant differences occur, this will necessitate using This procedure might be useful in estimating nutritive the more accurate method. An estimate of dry matter in value of range forages which greatly influence habitat total daily fecal output will be obtained using the chro- productivity. mic oxide ratio method combined with fecal grab sam- The method used in the second approach for esti- ples. mating nutritive value of forages (figure 2) provides an In steps 4 and 5, a representative sample of the diet, estimate of dry matter intake. This method is based on obtalned in step 3, is prcpared for zn uztro dry matter apparent digestibility of rumen dry niatter to calculate dlgestlbll~tydetermination by grlnding a nutritive value index as in the first approach. Steps In llcu of uslng rumen inocula from dcc11 lclllcd on 1 t,hrough 3 of figure 2 are the same as in figure 1, the study area for dcter~nlllingdry niatter rl~~:~~\t~bll~ty,except that it might be possible for the ii~vestigator to

(1) (2) (3) (4) TAME DEER MANUAL SELECTION SPECIES COMPOSITION PREPARATION OF SAMPLE ON LEASH OF FORAGE MIXTURE +OF 1)IE:T AND (, BY FOR LIGNIN DETERMINATION ------WEIGIIT OF COMPONENTS (VAN SOEST 1967 OR (SPARKS & MALECI-IEK 1968, CZERKOWSKI 1965) .c (5) RUMEN SAMPLES FROM ADULT DEER KILLED ON STUDY AREA

(6) DETERMINATION OF % LIGNIN IN RUMEN DM -- (7) CALCULATION OF APPARENT DIGESTION OF DM IN RUMENS ( RUMEN DMD) (ULYATT ET AL 1967) - - - -

(9) ESTIMATION OF MAXIMAI, VOLUNTAKY I/ LIGNIN IN FEED DM 4 " NVI INTAKE (I) USING APPARENT I (CALCU~~yrxDAS-e---- DIGESTIBILITY OF' RUMEN DM -OK-- '< LIGNIN IN DM SAMPLING POINT IN FIGURE 1) ULYATT IST Al, 1967 I 003I (8) IIEVELOI'MENT OF REGRESSION ($3) EQUATION (I a(/, RUMEN DlVID) r h) NVI + USING KC'MEN FISTULATED DEER FED LONG 13OUGlIACES

" I Dry 111,itlrr ~ntak(. Lcgcxnd IIMD Diy Maltrr D~qc\tih~llty,DM1 Dry Mattcr Inlakc, DM Dly Matter

Flgur~2 -JS\t~tnat~on ~j IL?L~IZ~LZ)F?VL~ILP for a mzxed-foiage dzet of deer 1~5111~~c~ppc~~cvtt dlge5tion of t~~tntnalDM to predzct forage zntake. obtain forage san~plesmanually by observing a tame deer determined during in 11ivo digestion trials (Troelsen a: on a leash. Rigsby 1964 1. The equation for the regression of volu tary DM1 on PSI, as determined in vivo, can be used Rumcn samples are obtained from several adult deer estimate voluntary DM1 from PSI obtained on forag on the study area and percentage lignin of the rurnen subjected to artificial mastication. A nutritive val dry matter is determined (steps 5 and 61. Ct)llection time for the rumen samples must be standardized. Late in index can be calculated as described previously, usii estimated DMI, in vivo or in uitro DMD, and the niet the afternoon seems to be inoit desirable bolic size of experimental deer. In figure 3, steps In step 7, apparent digestion of rumen dry matter is through 3 are the same as in the f~rsttwo appro~lclic determined by the ratio of percentage lignin in dry mat- In 5tep 4, rumen-fistulated deer are used to obtain ter of the forage sample to pcxrcentage lignin in rumen measure of voluntary intake of long roughages. Al: dry matter. one portion of the forage sample obtained in steu 3 I11 step 8, niaximum voluntary ~ntakemay be estimated stained and introduced into the rumen through the fistu from the regression of voluntary dry matter ~ntakeon for a 12-hour digestion period. This is illustrated in ste percent,~ge rumen dry matter digcstib~lity Thi5 esti- 4, 5, and 6. mate inay be calculated from a11 equat~ondeveloped spe- The unstained portion of the forage sample is subjectt clfically for decr by feeding long roughages, such as hay, to artificial mastication with an apparatus developed t to rutnen-tistulated deer IIowever, an equation devel- Troelsen and Bigsby (1964). Using another unstairlr oped for sheep fcd long roughage\ may be used forage sample, dry matter digestibility is determined t In step 9, using the estimated dry rnatter intake and either the in vizjo nylon bag technique or by the in ?lit apparent digest~bilityof rumen dry matter, NVI may method of Tilley and Terry ( 1963 J. The rumen-fist be calculated as in f~gureI lated deer serve as a source of inoculant for in viiro c gestibility trials. A sample of a standard ration fed The third approach (figure 3) for estimating nutritive the fistulated deer should he included in each in uit value of forages is based on the assuniption that rate of digestibility trial to permit adjustment of digestibili passage of digesta from the rumen is regulated by the values, as explained in the discussion of figure 1. size of particles present and voluntary intake is related to the rate of passage. This approach employs an arti- Following digestion, a sample of rumen contents ficial mastication technique that reduces forage to a obtained, dried, and a PSI computed (step 7). certain particle size. Particle size determined in this In step 8, an estimate of voluntary intake may be o manner was found to be highly correlated to a particle tained by calculating the regression of known voluntai size index (PSI) and voluntary dry matter intake (DMI) intake on PSI.

-, TAME DEER ON LEASH MANUAL SELECTION OF FORAGE MIXTURE SPECIES COMPOSITION OF DIET AND -- - - ,BY WEIGHT OF COR/II'ONE:NTS (SPARK: MALECI-IEK 1068) ------(4) /' (5) (6) TWO OR MORE RUMEN FEED DEER LONG PREPARATION-OF STAIN INTRODUCE STAINED EVACUATIC FISTULATED DEER ROUGHAGES AND SAMPLE FOR -SAMPLE-+FORAGE MIXTIJRE -OF RUMEN --P - - MEASURE VOLUNTARY RUMEN INTO RtJMEN CONTENTS INTAKE DIGESTION AFTER 12-H ------I DIGESTION IN VIVO OR UNSTAINED IN VITRO DMD SAMPLE - - - --I-- (7) / ARTIFICIAL MASTICATION DRY SAMPLE AND MEASURE SIZE 4 IIEI'LICATIONS OF 10-gm OF STAINED PARTICLES SA!VLPLES (THOELSEN AND (CALCIILATE PARTICI,E SIZE: BIGSBY 1964) INDEX TROELSEN & BIGSBY 1964)" - -- - - + (7) (a)------DETERMINE PSI ON REGRESSION T-OF VOLUNTARY UNSTAINED PARTICLES INTAKE ON PARTICLE SIZE INDEX (PSI) (9) 1 (8) -"Y a(PS1) i b NVI w ESTIMATE VOLUNTARY 1 IC~C~UL~N~- INTAKE USING (8) FIGURE I)

'~PSI: Rlcan suin of perecntage stained or unstained dried samples which p;~ssedthrough a scries of ~icves.divided by 100 and the coc cient of variation.

*" Y 1 Dry matter intake.

Legend: DMD - Dry Matter Digestibility; DM1 - Dry Matter Intake; IIM - Dry Mattcr.

Figure 3.-Estimation of nutritive valtce for (2 mixed-jorage diet of deer using an artificial n~ostication technique to predict forage intake. A PSI is determined on the portion of unstained Horton et al. (1964) describes how the nomograph can sample, and an estimate of voluntary intake is calculated be used to plan and execute the major job of vegetation from the regression equation developed in step 8. surveying in one stage. The model they used is based In step 9, NVI is calculated (as in figure 1) using the on line intercepts to measure cover density, but it can value for estimated voluntary intake and dry matter also be used to predetermine the number of samples digestibility. needed to measure yield. It is based on selecting several Any one of these three approaches may help to deter- plots to sample the extremes of vegetative conditions mine what percentage of the total available forage in a occurring within a type. habitat can be considered as usable and useful. The difficulty of measuring highly variable plant pop- ulations is demonstrated by Lyon (1968), who compared QUANTITATIVE MEASUREMENTS 19 different variations of quadrant and plotless sampling techniques in a known population density of bitter-brush. Sampling Considerations He found that "(1) Many methods would not produce a If forage surveys are to be useful in the South, all correct answer with any sample size, (2) all methods facets of the understory available as deer food should required unreasonably large samples to attain acceptable be measured seasonally by weight or other descriptive precision, (3) several methods required more effort than parameter common to all foods. More than one meas- counting all plants on 1 to 2 acres." urement technique may be necessary to survey all foods adequately. In the past we have used techniques devel- Sampling Procedures oped in the North, where fewer species of plants are important in deep snow zones. Woody plants, forbs, and grasses.-The following five steps will result in useful habitat productivity surveys: Plot size.-Guthrie ( 1964 1 found that in measuring woody browse, a 1-milaere plot was most efficient when 1. Start surveys randomly. comparing plots 1, 2, 4, 9, 25, and 29 milacres in size; 2. Consider using different sized sample plots, such yet in double sampling, the optimum was 4 milacres. as concentric circles or transects of variable length Conkle (1963) reported that optimum plot size was ob- or width. tained by selecting sizes that were most effective in 3. Obtain an adequate sized sample. reducing the coefficient of variation. Mesavage and Gro- 4. Equate food items by weight. senbaugh (1956) found that precision of estimates of 5. Conduct seasonal surveys. equal intensity increased as plots were made smaller and more numerous. A 100-percent clipping method, described by Campbell Whelan (1962), in determining minimum size for and Cassady ( 1955), provides the most accurate measure woody understory sampling units, found that 1/100-acre of forage yield, but is sometimes impractical because of units were adequate. For sampling grasses and forbs the time it consumes. The double-sampling technique ( on southern cattle ranges, Campbell and Cassady ( 1955 Wiim et al. 1944), which weighs and estimates weight recommended a 3.1-foot-square plot because of its con- of vegetation on a number of plots, as modified by appli- venient size and ease of conversion to pounds per acre. cation of the Dry Weight Prediction Method (Blair 1958) should avoid the undesirable features of the 100-percent Plot shape.-Ursic and McClurkin ( 1959 i found rec- clipping method. Another modification of the 100-per- tangular plots gave better representation of local varia- cent clipping method is the Ranked Set Sampling Meth- tion in the vegetation than square or circular plots of od, first described by McIntyre ( 1952). This procedure equal size. Distribution of species largely determines establishes sets of three closely-grouped quadrants which which shape plot is best. Johnson and Nixon (1952) are visually ranked within sets as to highest, inter- found that cruising time is less on a compact rectangular mediate, or lowest in forage weight. Only one quadrant plot than on a long, narrow one. Bormann ( 1953) states of each set is clipped and weighed. that "Variance decreases as plots are increased in length prov~dedthe longest axes of the rectangular plots cross For purposes of comparison, it is necessary to stand- observed contours and vegetational or so11 bandings " ardize terms describing what plant parts deer consume Number of plots.-The number of plots needed to and to what height they feed. General terms, such as "browse," should be discarded and replaced by more obtain a reliable sample from any area is not primarily specific terms, such as evergreen leaves, woody twigs, a fu~lctionof the size of the area, but rather of the varia- etc. tion from place to place within it. The number of sample plots for any degree of accuracy can be determined by Fruzt.-The value of acorns as food for deer has been statistical methods once the normal distribution around documented by Goodrum (19591, Harlow and Tyson the mean has been established (Bruce and Schumacher ( 1959 1, and Duvendeck ( 1962 1. The contribution of 1935; Snedecor 1946 1. Mesavage and Grosenbaugh understory fruits other than acorns, in east Texas, has (19561, using a systematic survey, suggested that the been reported by Lay ( 1961). number of sample plots should be calculated as though Methods of determining annual acorn mast and other the plots were located at random. Grosenbaugh ( 1952 ) fruit abundances have been described by several workers prepared two tables that are helpful in deciding the (Downs and McQuilkin 1944; Uhlig and Wilson 1952; number of samples to take. In one table it is necessary Edwards and Evans 1955; Gysel 1957; Crawford 1958; to have an estimate of thc population's coefficient of Sharp 1958; Lay 1961; Thompson 1962 i . variation; the other table provides a rough estimate of Difficulties encountered in measuring fruit abundance the coefficient of variation for different conclitions. and establishing its importance to deer include: (1) fruits may be eaten by all wildlife, not just deer; (2) In the scrub oak and scrub palmetto areas of the Coa their distribution is neither random nor uniform; ( 3 ) a1 Plain reglons, total frult counts per scrub car1 be ct fruiting is cyclic as well as species specific and differs vcnlently taken at randomly-located plots Adequ: greatly between trees of the same species; ( 4 1 fruiting s~zecounts are comparatively easy to obtaln (Narll success is influenced by such factors as climate, heredity, and Tyson 1959). soil, and stand conditions; (5) traps are necessarily Fallen 1euves.-The irnportance of evergreen foliti small for convenience in handling. as food for deer in the South during the winter 1- Burns et ill. ( 1954 ) placed four randomly located traps been documented. But the possible irnportance of fall per tree crown in one area and one trap per crown under deciduous leaves has not, until recently, been given mu trees of the same species in another area. The standard attention Studies relating to the importance of this fo error of the mean ranged from 40 to 50 percent when to deer have been reported by Watts ( 1964 ) and Die sampled with four traps and 14 to 60 percent when t 1965). sampled with one trap. Gysel (19561found little differ- The uniformity of hardwood leaf production over mc ence in number of acorns between traps when more than of the southern Appalachians lends itself to easy sarr one trap was placed under the same tree. Gyscl (1956) ling. Olson ( 1967) ratidomly placed 10 plastic garba and Thompson 11!)62i found no significant difference in cans (opening 1.89 square feet) on each 0.25-acre p: iiumber of acorns in traps arid adjacent ground plots to sample all litter. Leaf fall probably could be measur when counts were made weekly. with sufficient reliability with acorn mast traps or in Thompson t 1962) compared circular and square plots forage survey. 0.0001-, 0.0040-, 0.0010-, and 0.0023-acre in size and Mz~sl~roon~s.--Theephemeral nature and wide dispc found that none followed the conventional species-area sioii of fleshy mushrooms make it difficult to obta curve. He did find that a polyethylene film cone (3.00 reliable data on this important deer food. Prelimina mill was less influenced by wind, had better drainage, data required prior to conducting an adequate surv was not easily damaged by falling sticks, and had the must include: when they occur, their growth patter greatest efficiency in retaining acorns. how long they remain available, and what affects the Efficient but inexpensive and disposable oak seed traps distnbutlon. have been described by Klawitter and Stubbs (1961) and In telemetric deer studies conducted at the Universi Thompsoil and McGinnes (19631. Fifty-five gallon, open- of Georgia, workers demonstrated the ability of de top barrels make good permanent seed traps (Crawford to detect the presence of fungi obscured from hum, and Leonard 1965; Segelquist and Green 1968). Bushel sight. This indicates litter removal may be neeessa baskets have been used and found satisfactory by Minck- to sample mushrooms when using any sampling methc Ier and McDermott (1960). Merz and Brakkage (1964), Perennial mushrooms (fruiting bodies which displ, and Liscinsky (1966). Openings for small traps ranged annual growth rings i become abundant in cut-over are from 1.5 square feet for bushel baskets to 2.6 square feet and are found primarily on dead wood; they may be ce for steel drums. Downs and McQuilkin (1944) developed sused by using regular field plots. a wooden-wire trap with an opening of 10.89 square feet. Sumpling by photographg.-Brown and Worley ( 196 Their design has been a popular one. described the use of wide-angle to mcasu The necessity for using a large number of small traps crown canopy and to determine aerial coverage, slo to obtain statistical reliability was demonstrated by and aspect, tree heights and distances, basal area stoc Thompson (1962). Based on 18 traps 0.0001 acre in size ing, identification of plant species, and to count trees. located in each of six study areas, he found that the Photography is a quick method of measuring featur standard error of the mean for number of acorns per difficult to ascertain by other field methods. Many 2 mast trap indicated he would need from 53 to 1,228 traps pects can be photographed on a single entry for a poi to obtain the required sampling intensity. location. The worker should, however, recognize ti The following rccomrnendat~ons for measurirlg tree limitations in using photography-its inability to me: fru~tproduction are bawd on a rev~ewof current l~tera- ure the inconspicuous or seasoiially available food iter ture: such as fruit, mushrooms, and fallen leaves. 1. To establish fruit productioll potenlial- The irifluence of canopy cover on understory densi a Determine lserccntage of canopy cover by has been illustrated by Shaw and Ripley (1965), Sch~ forest type for entire area ter ( 1967 ) , and Young et al. ( 1967 ) . Jameson ( 196' b. Determine extent of canopy cover in mast- presents a mathematical equation which fits overstor producing trees undcrstory data better than previously-used equatioi c Coilvert extent ot canopy cover of mast- producing trees to square feet, and d. Deterlnine the square-foot covcrnge of mast- Hcforc valid estimates of habitat productivity are pc producing trees by species. sible, it will be necessary to ascertain the quality a1 2. To deterlninc fruit production quantity of available rood on a seasonal basis and rela a Randomly select sampling points, and these factors to the nutritional requirements of whit b Sample a i~~fficlentnumber of trees cach year to obtain square-foot production of fru~tfor it11 R. L. &2archinton, School each species. 'corgia, Athens, Ga. March 19f LITERATURE CITED ation Workshop. USDA Misc. Pub. 1147. (In press ) Adams, W. H., Jr. 1960. Population ecology of white-tailed deer it1 north- Crampton, E. W. eastern Alabama. Ecology 41: 785-790. 1957. Interrelations between digestible nutrient and energy content, voluntary dry matter intake, Arnold, G. W., and Dudzinski, M. L and the overall feeding value of forages. J. 1967. Comparison of faecal nitrogen regression and Anim. Sci. 16: 546-552. in vitro estimate of diet digestibility for esti- mating the consumption of herbage by grazing Crampton, E. W., Donefer, E., and Lloyd, L. E. animals. Brit. J. Agr. Sci. 68: 213-219. 1960. A nutritive value index for forages. J. Anim. Barnes, R. F. Sci. 19: 538-544. 1966. The development and applicatio~lof in vitro Crampton, E. W., and Harris, L. E. rumen fermentation techniques. Tenth Int. 1969. Applied animal nutrition. Ed. 2, 753 pp. Sari Grassland Congr. Proc., pp. 434-438. Francisco: W. H. Freeman and Co. Baumgardt, B. R., Cason, J. L., and Taylor, M. W. Crawford, H. S. 1962. Evaluation of forages in the laboratory. I. Com- 1958. Sampling the production of fruits and seeds of parative accuracy of several mcthods. J. Da~ry woody and herbaceous plants. In Techniques Sci. 45: 59-61. and methods of measuring understory vegcta- tion, pp. 116-122. USDA Forest Serv. South. Baumgardt, B R , Taylor, M W , and Cason, J L and Southeast. Forest Exp. Stas. 1962 Evaluation of forages In the laboratory 11 Slmpllfled artlflclal rumen procedure for ob- Crawford, H. S., and Leonard, R. G. tainlng repeatable estimates of forage nutritive 1965. The Sylamore deer study. Southeast. Assoc. value J Dairy Sci 45 62-68 Game and Fish Comm. 17th Annu. Conf. Proc. 1963: 9-13. Blair, R. M. 1958. Weight techniques for sampling browse produc- Czerkowski, J. tion on deer ranges. In Techniques and mcthods 1965. Estimation of lignin. Nutrition Soc. Proc. 24: of measuring understory vegetation, pp 26-31. 17. USDA Forest Serv South. and Southeast Forest Exp Stas. Dietz, D. R. 1965. Deer nutrition research in range management. Bormann, F H Thirtieth N. Amer. Wildlife and Natur. Re- 1953 The stat~st~caleff~ciency of sample plot size sources Conf. Trans. 1965: 274-285. and shape in forest ecology Ecology 34 474- 487. Donefcr, E. E., Crampton, E. W., and Lloyd, L. E. 1960. Prediction of the nutritive value index of a Brown, H E , and Worley, D P. forage from in vitro rumen fermentation data. 1965 Some applicat~ons of the canopy camera In J. Anim. Sci. 19: 545-582. forestry J Forest 63 674-680 Downs, A. A,. and McQuilkin, W. E. Bruce, C, and Schumacher, F X. 1944. Seed production of Southern Appalachian oaks. 1935 Forest mensuration 360 pp N Y McGraw- J. Forest. 42: 913-920. Hill Book Co. Duvendeck, J. P. Burns, P Y , Christlsen, D M , and Nichols, J M 1962. The value of acorns in the diet of Michigan 1954 Acorn productlon In the Mlssouri Ozarks Mo deer. J. Wildlife Manage. 26: 371-379. Agr Exp Sta Bull 611, 8 pp Edlefseil, J. L., Cook, C. W., and Blake, J. T. Campbell, R S , and Cassady, J T. 1960. Nutrient content of the diet as determined by I 1955 Forage welght inve~ltorlesoil southern forest hand plucked and esophageal fistula samples. ranges USDA Forest Serv South Forest Exp J. Anim. Sci. 19: 560-567. Sta Occas Pap 139, 18 pp 1 Edwards, M., and Evans, T. 1 Conltle, M T 1955. Mast stimulation and prediction-oak mast pro- 1 1963 The dctcrminatlon of experimental plot size duction. N. C. Wildlife Resources Comm., Work and shape in loblolly and slash plnes N C Plans 1955-1956. Raleigh. i State Coll Sch Forest Tech Rep 17, 51 pp I French, C. E., McEwen, Id. C., Magruder, N. D., and others. 1 Cook, C. W. 1955. Nutritional requirements of white-tailed deer 1 1956. Range livestock nutrition and its importance in for growth and antler development. Penn. Agr. the intermountain region. Utah State Agr. Coll., Exp. Sta. Bull. 600, 50 pp. 17th Annu. Fac. Res. Lect., 28 pp. Frischknecht, N. C., and Harris, L. E. / Cowan, R. L., Jordan, J. S., Grimes, J. L., and Gill, J. D. 1968. Grazing intensities and systems on crested Conlparative nutrit~vevalues of forage spec~es wheatgrass in central Utah: Response of vege- for deer In Range and Wildhfe Hab~tatEvalu- tation and cattle. USDA Tech. Bull. 1388, 47 pp. Gaillard, B. D. E. value of several winter deer foods. J. Wildlif 1968. Calculation of the digestibility and the starch Manage. 4: 315-325. equivalent value of herbage from chemical an- Horton, J. S., Robinson, T. W., and McDonald, H. R. alysis. New Zeal. Soc. Anim. Prod. 28: 49-52. 1964. Guide for surveying phreatophyte vegetatior Gill, J. D. USDA Agr. Handbook 266, 37 pp. 1956. Regional differences in size and productivity Jameson, D. A. of decr in West Virginia. J. Wildlife Manage. 1967. The relationship of tree overstory and herb: 20: 286. ceous understory vegetation. J. Range Managc Goodrum, P. D. 20: 247-249. 1959. Acorns in the diet of wildlife. Southeast. Assoc. Johnson, F. A,, and Nixon, PI. J. Game and Fish Comm. 13th Annu. Conf. Proc. 1952. The most efficient size and shape of plot t 1959: 54-61. use for cruising in old growth Douglas fir tirr ber. J. Forest. 50: 17-20. Grimes, J. L. 1968. Nutritive evaluatiori of various decr foods. I. Johnson, R. R., and Dehority, B. A. Effect of plant species, plant morphology, and 1968. A comparison of several laboratory technique season. 11. Comparison of sheep and deer as to predict digestibility and intake of forage: experimental animals. M. S. Thesis. Penn. State J. Anim. Sci. 27: 1738-1742. Univ., University Park. 97 pp. Kirkpatrick, R. L., Fontenot, J. P., and Harlow, R.F. Grosenbaugh, Id. R. 1969. Seasonal changes in rumen chemical cornpon 1952. Plotless timber estimates: new, fast, easy. J. ents as related to forages consumed by white Forest. 59: 32-37. tailed deer in the Southeast. N. Amer. Wildlif and Natur. Resources Conf Trans. (In press Guthrie, A. W. 1964. An investigation of sampling methodology of Klawitter, R. A,, and Stubbs, J. winter browse. M. S. Thesis. Va. Polytech. Inst., 1961. A reliable oak seed trap J. Forest. 59: 291-292 Blacksburg. 48 pp. Klein, D. R. Gysel, L. W. 1962. Rumen contents analysis as an index to rang 1956. Measurement of acorn crops. Forest Sci. 2: quality. Twenty-Seventh N. Amer. Wildlif 305-313. and Natur. Resources Conf. Trans. 1962: 150 162. Gysel, L. W. 1957. Acorn production on good, medium, and poor Lancaster, R. J. oak sites in southern Michigan. J. Forest. 55: 1949. The measurement of feed intake by grazin, 570-574. cattle and sheep. I. A method of calculatin, the digestibility of pasture based on the nitrogei Hagen, EX. L. content of feces derived from the pasture. Neb 1953. Nutritive value for decr of some forage plants Zeal. J. Sci. Tech. 31: 3. in the Sierra Nevada. Calif. Fish and Game 39(2): 163-175. Lay, D. W. 1957. Browse quality and the effects of prescribe1 Harlow, R. F., and Jones, F. K. burning in southern pine forests. J. Forest. 55 1965. The white-tailed deer in Florida. Fla. Game 342-347. and Fresh Water Fish Comm. Tech. Bull. 9, 240 Lay, D. W. PP 1961. Fruit production of some understory hardwoods I-Iarlow, R. F., and Tyson, E. L. Southeast. Assoc. Game and Fish Comm. 15tl 1959. A preliminary report on the effect of mast Annu. Conf. Proc. 1961: 30-37. abundance on the weight and reproduction of Liscinsky, S. A deer in central Florida. Southeast. Assoc. Game 1966. Cutting intensity study in a pole stage norther and Fish Comm. 13th Annu. Conf. Proc. 1959: hardwood stand. Penn. P-R Final Rep. Pro" 62-69. W048-R014, 23 pp. Hart, G. H., Guilbert, H. R.. and Goss, H. Lucas, A. L. 1932. Scasonal changes in the chemical coniposition 1962. Determination of forage yield and quality fror of range forage and their relation to nutrition animal responses. In Range research method: of animals. Calif. Agr. Exp. Sta. Bull. 543, pp. 43-54. USDA Misc Pub. 940. 62 PP. Lusk, J. W., Browning, C. B., and Miles, J. T. Hcaly, W. M. 1962. Small sample in viva cellulose digestion prc 1967. Foragc preferences of captive decr while free cedure for forage evaluation. J. Dairy Sci. 45 ranging in the Allegheny National Forest. Ph.D. 69-73. Thesis. Penn. State Univ., University Park. 93 Lyon, J. L. PP. 1968. An evaluation of density sampling methods i IIcllrners, H. a shrub community. J. Range Manage. 21 1940. A study of monthly variations in the nutritive 16-20. McEwen, L. C., French, C. E., Magruder, N.D., and others. woody browse for deer. Soc. Amer. Forest. 1957. Nutrient requirements of the white-tailed deer. Proc. 1965: 229-233. Twenty-Second N. Amer. Wildlife Conf. Trans. Short, H. L. 1957: 119-132. 1963. Rumen fermentations and energy relationships Mclntyre, G. A. A. in white-tailed deer. J. Wildlife Manage. 27: 1952. A method for unbiased selective sampling, using 184-195. ranked sets. Australian J. Agr. Res. 3: 385-390. Short, H. L., Dietz, D. R., and Remmenga, E. E. Maynard, L. A., and Loosli, J. K. 1966. Selected nutrients in mule deer browse plants. 1962. Animal nutrition. 533 pp. N. Y.: McGraw-Hill Ecology 47: 222-229. Book Co. Short, H. L., and Harrell, A. Merz, R. W., and Brakkage, G. K. 1969. Nutrient analysis of two browse species. J. 1964. The management of pin oak in a duck shooting Range Manage. 22: 40-43. area. J. Wildlife Manage. 28: 233-239. Silver, H. Mesarage, C., and Grosenbaugh, L. R. 1968. Decr nutrition studies. In The white-tailed deer 1956. Efficiency of several cruising designs on small of New Hampshire, pp. 182-196. N. H. Fish tracts in north Arkansas. J. Forest. 54: 569-576. and Game Dep. Snedecor, G. W. Minckler, L. S., and McDermott, R. E. 1946. Statistical methods. Ed. 4, 485 pp. Ames: Iowa 1960. Pin oak acorn production as affected by stand density, structure, and flooding. Mo. Agr. Exp. State Coll. Press. Sta. Bull. 750, 24 pp. Sparks, D. R., and Malechek, J. C. 1968. Estimating percentage dry weight in diets using Murphy, D. A,, and Coates, J. A. 1966. Effects of dietary protein on deer. Thirty-First a microscopic technique. J. Range Manage. 21 : 264-265. N. Amer. Wildlife and Natur. Resources Conf. Trans. 1966: 129-139. Swift, R. W. 1948. Deer select most nutritious forages. J. Wildlife Oh, H. K., Baumgardt, B. R., and Scholl, J. M. Manage. 12: 109. 1966. Evaluation of forages in the laboratory. V. Comparison of chemical analyses, solubility Thompson, R. L. tests and in vilro fermentation. J. Dairy Sci. 1962. An investigation of some techniques for meas- 49: 850-855. uring availability of oak mast and deer browse. M. S. Thesis. Va. Polytech Inst. Blacksburg. 65 Olson, D. F., Jr. PP. 1967. Leaf and litter production in relation to stand characteristics for even-aged yellow-poplar Thompson, R. L., and McGinnes, B. S. stands. USDA Forest Serv. Southeast. Forest 1963. A comparison of eight types of mast traps. J. Exp. Sta. Study Plan FS-SE-1102, No. BC-31, Forest. 61: 679-680. 15 rJP. Thorsland, 0. A. Reid, J. T. 1967. Nutritional analyses of selected deer foods in 1962. Animal performance 11. In Pasture and range South Carolina. Southeast. Assoc. Game and research techniques, pp. 43-45. Ithaca, N. Y.: Fish Comm. 20th Annu. Conf. Proc. 1966: 84- Comstock Pub. Assoc., Cornell Univ. Press. 104. Schneider, B. H., Lucas, H. L., Cipolloni, M. A,, and Tilley, J. M. A., and Terry, R. A. Pavlech, H. M. 1963. A two-stage technique for the in vitro digestion 1952. The prediction of digestibility for feeds for of forage crops. J. Brit. Grasslands Soc. 18 : 104. which there are only proximate composition Troelsen, J. E., and Bigsby, F. W. data. J. Anim. Sci. 11: 77-83. 1964. Artificial mastication-a new approach for pre- Schuster, J. L. dicting voluntary forage co~lsumptionby rumi- 1967. The relation of understory vegetation to cutting nants. J. Anim. Sci. 23: 1139-1142. treatment and habitat factors in an cast Texas Uhlig, H. G., and Wilson, H. L. pine-hardwood type. Southwest. Natur. 12: 1952. A method of evaluating an annual mast index. 339-364. J. Wildlife Manage. 16: 338-343. Segelquist, C. A,, and Green, W. E. Ullrey, D. E., Youatt, W. G., Johnson, 13. E., and others. 1968. Deer food yields in four Ozark forest types. J. 1964. Digestibility of cedar and aspen browse for the Wildlife Manage. 32: 330-337. white-tailed deer. J. Wildlife Manage. 28: 791- Sharp, W. M. 797. 1958. Evaluating mast yields in the oaks. Penn. Agr. Ullrey, D. E., Youatt, W. G., Johnson, H. E., and others. Exp. Sta. Bull. 635, 21 pp. 1967. Digestibility of cedar and jack pine browse for Shaw, S. P., and Ripley, T. H. the white-tailed deer. J. Wildlife Manage. 31: 1965. Managing the forest for sustained yield of 448-454. Ullrey, D. E., Youatt, W. G., Johnson, H. E., and others. Watts, R. C. 1068. Digestibility of cedar and balsam fir browse 1964. Forage preferences of captive deer while fr for the white-tailed deer. J. Wildlife Manage. ranging in a mixed oak forest. Penn. Sta 32: 162-171. Univ. Coop. Wildlife Res. Unit. Pap. 112, 16 F Ulyatt, M. J., Blaxter, K. L., and McDonald, I. Weir, W. C., and Torell, D. T. 1967. The relations between the apparent digestibility 1959. Selective grazing by sheep as shown by a cor of roughages in the rumcn and lower gut of parison of the chemical composition of ran, sheep, the volume of fluid in the rumen and and pasture forage obtained by hand clippi] voluntary reed intake. J. Brit. Soc. Anim. Prod. and that collected by esophageal-fistulatl 9: 463-476. sheep. J. Anim. Sci. 18: 641-649. Ursic, S. J., and McClurkin, D. C. Weller, R. A,, Gray, F. V., and Pilgrim, A. F. 1959. Small plots for measuring vegetation composi- 1958. The conversion of plant nitrogen to microbi sition and cover. In Techniques and methods nitrogen in the rumen of sheep. Brit. J. Nut of measuring understory vegetation, pp. 70-78. 12: 421-428. USDA Forest Serv. South. and Southeast. Forest Exp. Stas. WeIIer, R. A,, Pilgrim, A. F., and Gray, F. V. 1962. Digestion of foodstuffs in the rumen of tl Van Dyne, G. M. sheep and the passage of digests throuzh i 1968. Prediction by nonlinear programming of rela- compartments. 111. The progress of ilitrogr tive chemical composition of dietary botanical digestion. Brit. J. Nutr. 16: 83-90. components. J. Range Manage. 21 : 37-60. Wetzel, R. S. Van Dyne, G. M., and Meyer, J. I-I. 1968. The seasonal utilization of calcium and phc 1964. A method for measurement of forage intake of phorus by white-tailed deer. M. S. Thesis. Pen grazing livestock using microdigestion tech- State Univ. University Park. 41 pp. niques. J. Range Manage. 17 : 204-208. Whelan, J. B. Van Soest, P. J. 1962. The correlation between available deer brows 1963. Use of detergents in the analysis of fibrous forest cover type and forest site. M. S. Thes: feeds. 11. A rapid method for the determination Va. Polytech. Inst. Blacksburg. 125 pp. of fiber and lignin. J. Assoc. Offic. Agr. Chem. 46: 829-835. Wilm, H. G., Costello, D. F., and Klipple, G. E. 1944. Estimating forage yield by the double sampli~ Van Soest, P. J., and Wine, R. I-I. method. J. Amer. Soc. Agron. 36: 194-203. 1967. Use of detergents in the analyses of fibrous feeds. IV. Determination of plant cell-wall Wilson, A. D. constituents. J. Assoc. Office. Agr. Chem. 50: 1969. A review of browse in the nutrition of grazir 50-55. animals. J. Range Manage. 22: 23-28. Van Soest, P. J., Wine, R. H., and Moore, L. A. Young, J. A,, Hedrick, D. A,, and Keniston, R. F. 1966. Estimation of the true digestibility of forages 1967. Forest cover and logging. Herbage and brow by the in vitro digestion of cell walls. Tenth production in the mixed coniferous forest Int. Grassland Congr. Proc. 1966: 439-441. northeastern Oregon. J. Forest. 65: 807-81 Evaluating Food Use-New Methods and Techniques

H. S. Crawford Southeastern Forest Experiment Station Forest Service, U. S. Departnient of Agriculture Blacksburg, Virginia

To e7+nluatetrue deer foods zt 2s necessary to de- EVALUATING DEER FOOD : (1) terrr~iitcobject ivelg the plants and plant parts Wcbstc?r's New Collegiate Dictionarg ( 2nd edition I ingc~.slcd.(2) the rzufrient content of ingested ma- defines food as "Nutritive material taken into an organ- te~inl.13) ihe digestible portion of the ingested ma- ism for growth, work, or repair and for maintaining terial, and 141 the ability of digested rrutrients to the vital processes." Certainly the entire volume of vegetation within reach of a deer is not food, as has To ccalltrrte trcnds zn the cond~tzonof deer kubztat, been recognized by biologists for years. To evaluate we must keep a runnzng balance of tlze true food deer food in the environment we must measure: (1) ~.ul?~eoj the lzabztat Much of tlze deer's food comes quantities of plant parts ingested, (2) quality of the in- from pctrfs of plunts thaf are present In thc hahztat gested material-its chemical composition and caloric foi rhmi per1od5 and sho~ulzttlc evzdeizce of utzlzza- value, 13)digestion of plant nutrients, and (4) ability tzon New lechnzq~rermutt be de~~elopedthat adc- of digested nutrients to meet animal requirements. quately sample plant prodzlctzor~ and utzl~zcct~onof Once the nutritional requirements of deer are well cach plant or plant part In the mzxed dzet of d~er established, food may be equated to potential animal numbers. All necessary steps for determining food have Changes In the supply of food whlch support a deer herd have been evaluated by examlnatlons of the b~o- seldom been followed in past investigations. We have been weak even in objectively determining what plants loglcal charactcr~st~csof the deer and by examination of the habitat Sampllng deer and rclatlng gross changes or plant parts were ingested. in animal physical cond~tionand fecundity to food supply FOODS INGESTED 15 an appro'lch used by many wildllfe management agen- cies Thc w~ldhfeliterature abounds with reports on Dunkeson (19551 questioned existing methods of deter- many an~m,~lmeasurements Anlmal measurement sys- mining deer food ingestion when he alluded to a "woody- temi arc usually ielat~vclysrmplc and inexpensive be- twig bias." IIe observed plant materials eaten by a cauie data are easily collc~ctedtluririg irurilirrq wasuris semitame deer in a large enclosure in the Missouri Ozarks IIowcver anlmal changes lag behlnd hab~tatchanges and found that forbs, mushrooms, and fruits were fre- By the tlme habltat deterioration is reflected by apparent quently consumed. Korschgen 119621 made stomach changes In the deerherd, 11 is usually too late to reverse analyses of several hundred deer. His work showed the the trend in habltat A deterlorated habitat recovers great variety of plants eaten and the minor importance slowly even with complete protection (I-lalls and Craw- of woody twigs. Harlow (1961)in Florida and Lay (19671 ford 19601 Evcn ii animal changes were apparent prlor in Texas have reported similar results. Current studies to hab~tatchanges, the land manager would be handi- in the southeast are showing little use of hardened capped because he lachs deta~led information needed woody twigs and heavy use of a variety of other plant to apply proper h'lbitat modillcatlon measure4 parts.' There is now little basis for retaining a woody Measuring the food ava~lableand food consumed In twig bias in the South. Lay in this symposium reports the liabltat requires 11)knowledge of the plants and on foods of deer in greater detail. plant parts which make up the dccr 5 food supply, and Discounting unusual items, such as old inner tubes, (2) suitable tcchn~quei;to mc~isurelood ava~lab~lltvand the great variety of plant parts ingested by deer in the utth~ation If habitat changes can be detected before South can be separated into general- categories:- they noticeably affect animals. management can offset 1. Dormant woody twigs, hardened-minor impor- impending habitat deterioration. Measuring available tance. deer food and its utilization is more difficult and expen- sive than measuring the resultant changes in the deer. 2. Photosynthesizing woody twigs-green stems of dogwood, blueberry, greenbrier, sassafras, and other If we desire to manage deer rather than have them plants are consumed wliile leafless, probably when stems manage themselves, howcver, we must maintain and improve their habitat. Allowing deer to manipulate are photosynthetically active. lhcir habitat has resulted in widespread habitat deterior- 3. Succulent buds, twigs, and leaves of woody plants ation. -eaten while actively growing, mostly in spring. My assignment for this conference was to report on 4. Evergreen leaves-a major winter food which in- new techniques for evaluating food use. Rather than cludes leaves of woody and herbaceous plants. describe new techniques, 1 will discuss the general meth- 5. Fruit--a major autumn and winter food when ods of evaluating food use. Techniques mentioned are not available, and also consumed in other seasons. - - neccssarily new but are not as yet used to any extent by ' Data on file at Southeastern Forest Experiment Station, the wildlife profession. Blacksburg, Va. 6. Grasses-particularly cool-season grasses which the basis of palatability, we could objectively evalu are eaten when actively growing during fall and spring. potential ingestion of all plants, both those present 7. Seasonal forbs-heavy use of many species, pri- the environment and those that mlght be introduc marily during their period of active growth, hut also Dunkeson (1955) indicated that deer selected plants after growth is completed. smell New work by Longhurst et al. ( 1968) ind~ca 8. Fungi-many species are consumed heavily when that plant selection by deer is first determined by sml available during several seasons. then taste; and after the plant is found acceptable, is selected by visual recognition. Efforts are being m: 9. Dried leaves-little is known about their impor- by Longhurst and his coworkers to isolate plant co tance, but they are often ingested. ponents that contribute to aroma. If thls can be acco Several monographs and symposia have dealt with plished, a new area of research wlll be opened.. the many methods used to determine kinds and amounts It should be possible to develop techniques of obj of plants ingested. Brown (1954) devoted four chapters tively determining palatability and use them in habi to measurements of plant utilization in open grasslands, evaluation. It would then be possible to distingu shrub lands, and dense pastures of humid regions. A plants or plant parts that would be consumed-assum] "Symposium on Forage Evaluation" was reported in certarn animal densities-and base habitat evaluat 1959 in Volume 51 of the Agronomy Journal. A chapter on only these plants or plant parts. was devoted to methods of measuring forage utilization The subjective palatability ratings used in the p in "Basic Problems and Techniques in Range Research" necessarily had to be related to crrtaln environme (National Academy of Sciences-National Research Coun- and could not be considered absolute throughout 1 cil, 19621. Several symposia held by the Forest Service range of the deer Plants or plant parts desirable have dealt in part with measuring ruminant utilization deer In one area may not have been desirable in a slml of plants (USDA Forest Service 1959, 1963 ). Wild- environment nearby. For instance, flowering dogwc life lnvestigational Techniques ( Mosby 1963 ) contains (Cornus florida L.) was eaten in the Arkansas Ozai a chapter dealing primarily with stomach content an- (Halls and Crawford 1960; Crawford and Leonard 19E alysis. A Forest Service Range and Wildlife Habitat but not on nearby areas in Missouri (Korschgen 191 Evaluation Research Workshop, held in May 1968, in- Dunkeson 1955). This was probably caused by a dt cluded several papers on determining forage quality irruption in Arkansas, which was serious enough and animal consumption. (The transactions will be eliminate more desirable deer foods from the plant co printed as USDA iVIiscellaneous Publication 1147. ) Addi- munity. Deer die-offs were noted (Alexander 1954; R. tionally, many separate papers present techniques for Leonard, personal correspondence) during years of acc determining ingestion of plants by deer. Methods dis- shortage. Dogwood eaten in the Arkansas habitats prc cussed in these works vary from a visual estimate of ably would not have been consumed if more desiral plants removed based on plant appearance and direct foods had been available. An objective palatability rati observations with binoculars of feeding animals in the system would have placed dogwood in its proper p wild state to more sophisticated studies of tame lead spective as an alternate food, while a browse surt deer and animals equipped with rumen or esophageal system would have rated it very desirable simply beca~ fistulae. it was browsed. The criterion of a satisfactory method for measuring Standard survey methods can rate desirable foods food ingestion is how well it measures all categories of unimportant either because they are not abundant ingested plants or plant parts-not just their occurrence, because utilization cannot be measured. Some who stu but the amounts consumed. The major shortcomings of deer recommend a low ranking for palatable foods wh past techniques are that they were not quantitative, plants are not abundant because management bas unbiased, or adequately replicated. on "ice cream" species dictates acceptance of fewer de The use of a tame deer fitted with an esophageal However, we should objectively rank these plants fistula appears promising for obtaining sound quantita- palatability alone and, through research, learn to mod tive samples. Studying tame deer without a fistula the environment and increase palatable species. Bas provides information on kinds of food eaten but not on subjective survey ratings, it is conceivable that reco amounts (Wallmo and Neff, in press). Proper replication mended habitat management measures could be inc requires that an adequate number of tame, fistulated rectly directed toward increasing unpalatable food deer be available. Providing facilities, personnel, and the expense of palatable food. feed to maintain a herd of captive deer will be expensive, but if we intend to get sound data we must be ready to Food Quality pay for it. Unfortunately, no data exist on how many Food quality in deer habitats has been determined fr~ deer will be needed to sample the various habitats ade- an arbitrarily assigned component of vegetation, usua quately. It may not be necessary to examine the entire the current season's total growth of certain woody plar sample taken through a fistula. Sparks and Malechek The same practice has been used in sheep and cat (1968) describe a method to determine botanical compo- studies, but the assigned vegetation component v sition by weight based on 100 microscopic examinations usually the season's total growth of certain grasses a of each sample. forbs. Weir et al. (1959) compared the difference Some plant species and plant parts are ingested and chemical content of fistula samples and clipped sampl others are not. Plant material readily ingested or sought They concluded " . . . the selective grazing practic out by the animal is considered palatable. If we knew by animals . . . makes it essential that our research evaluate the nutritive value of forages be based on meth- priate proportion gives the weight of various nutritional ods which represent the lorage actually consumed rather components in the sample. than forage available." Because of the great variety of 7. Determine the weight of nutritional components plants consumed by deer, this conclusion applies even of the vegetation. It is the total weight in step G ad- more in deer habitat evaluation. justed for the weight of similar nutrients from the saliva Proper use of esophageal fistulae should accurately contaminant. determine the components of ingested forage. This Van Dyne and Torell 11964) criticized Cook's procedure technique can also be used to provide samples for nu- because of assumptions (1) that hand-plucked samples trient analysis if certain problems are recognized. Chem- are similar to grazed forage, 12) that saliva composition ical composition of samples collected by esophageal fis- or secretion rate does not change throughout thc samp- tulae are biased hy contamination from saliva, and this ling period, and (3) that fistula samples are completely bias changes with the animal's diet (Rice 1968). Samples saturated with plant or salivary moisture. are also affected by the method of handling-failing to freeze the sample soon after collection, draining Use of a water-tight container in the modified pro- samples, ovendrying, or rinsing with tap water (Bow- cedure ensures that samples are completely saturated. man and Lesperance 1967; Rice, in press). Contamination is assured, but can be quantified. Col- lecting the sample over a short time-perhaps 1'5 hours Cook 11964) corrected for saliva contamination by --and immediately placing it on dry ice would limit determining the nutrient content of saliva from each bias resulting from chemical changes in plant tissues. animal and then saturating a dried sample of ingested forage to determine total moisture held by the dry ma- Calibrating the animal to the range and s2mpling over terial. Field moisture content determined from hand- a short period should limit changes in composition of plucked samples of the forage was subtracted, and nu- the saliva. Saliva composition changes can be checked trient values of the vegetation were differentiated from by analyzing saliva samples taken during and following saliva values. Fistula collections were made in bags vegetation collections. with screen bottoms which let excess saliva drain, cre- The modified technique also assumes that hand- ating the possibility of leaching loss. plucked samples are similar in moisture content to The following untested modification of Cook's pro- grazed forage. This assumption could be checked with cedure is suggested for further study when using fistu- the isotope-dilution technique (Van Dyne and Torell lated deer: 19641, wherein the animal is dosed with an isotope, and moisture contributed by animal saliva can be distin- 1. Calibrate by allowing the animal to feed normally guished from plant moisture. Isotope-dilution was recom- on the study area for a few days before sampling. This mended as a means of measuring salivary contamination should allow a relatively constant chemical composition of fistula samples and its use should be considered. Re- of the saliva and some time for the rumen population cause of restrictions on use of radioactive isotopes, this to adjust. Length of the proper adjustment period is technique could serve as a check for other, more general unknown and should be studied. methods of determining contamination. 2. Take a saliva sample through the fistula immedi- Van Dyne and Heady (19651 studied variability in ately before the animal begins to feed. Freeze the sample measuring dietary constituents from a range when sam- for subsequent chemical analysis to determine weights pled by esophageal fistulae in cattle and sheep. They of various nutritional components. reported that six animals would be adequate to sample 3. Collect vegetation through the fistula in an air- most dietary constituents within 10 percent of the mean tight, water-tight container placed within an insulating at the 95-percent confidence level. Habitat and feeding container of dry ice. Determine weight of wet sample. habits of deer are probably more variable than those Keep the sample cold until it can be freeze-dried. The of cattle or sheep, and more animals may be required. length of time that the sample can be collected is limited Maintaining control over several lead deer requires con- to the time of active feeding and by the size of the col- siderable manpower. Marchinton and Baker (1967) des- lection bag. Sampling should be terminated before the cribe the use of a free-ranging tame deer carrying a animal starts to ruminate because regurgitation con- radio transmitter to aid in locating the animal for obser- taminates the sample. vation. Radio transmitters could be placed on free- 4. Observe the vegetation being collected through ranging fistulated animals if some means of automatic the fistula and hand-pluck vegetation similar to that fistula closure were developed. It might then be possible eaten by the animal. Determine moisture content of for a few men to collect samples from several tame, the hand-plucked vegetation. (Hand-plucking may be fistulated deer. deferred until after the samples are examined in the Short (1967)discusses several precautions to take when laboratory so the composition can be more accurately collecting plant samples for chemical analysis. Plants determined.) may have to be collected at comparable times of the 5. Determine weight of saliva in the wet sample. day, because chemical composition of the protoplasm It is the total weight less the sum of the dry weight of will be different after a day of photosynthesis than after vegetation and the weight of plant moisture determined a night of respiration. During periods of rapid growth, from the plucked sample. chemical properties of plants will differ in a shorter time 6. Determine, by chemical analysis, proportions of than during less active periods. Thus, sampling fre- various nutritional components of the sample collected quency should be related to phenological stage, the im- in step 3. Sample dry weight multiplied by the appro- portance of which has often been overlooked in habitat evaluation Short also stresses the importance of im- range and l~vcstockworkers for years Difficulties mediately stopping respiratory processes ill the sample using the technique are probably not insurmoui~t~ by quick-freezing and drying in a cold, low-$ressure '~ndin the ncxt icw years it should become a u\c system. tool lor deer management research Captivity, h,indl Arguments about which nutriellts are important to by humans, and f~stullzationm,iy alter normdl feed the an~mxland the validity or accur'icy of method\ for paltcrni of dcer Feeding patterns of cxpcrimcnt,il , measurirrg these nutrients are beyond the scope of this in,ils will have to be comp,ired with thoscl of wild dt paper. and in the~nselveswoulti provide substance for several symposia. It is important to nt~tcthat new meth- Animal Response to Digested Nutrients ods are evolvil~gin these areas. Dietz's paper (in press) The response of an animal to the nutrients diges is recommendctl lor recent in for ma ti or^ on food quality from its food is the final evaluation of ;I food. Anii response may be mcasured by several criteria---its abi Digestibility to gain weight, produce offspring, grow antlers, or cc Even though plants with high nutrient values are binations of these or other criteria. Equating digesti ingested, it does not follow that the totd value goes nutrients with animal performance ties food ingest to the animal Plants or plant parts may tiot be cqually food quality, and food digestion to management gc digested. Nagy (in press) points out " . . . that we are which select the important criteria. really fccding billions of micro-organisms whose diges- Determining animal nutrient requirements is proba tive activities in turn provide most of the i?utritiotlal the most important need in deer management at t l~eedsof the ruminant animal " He stres\cs the need time. Nutrient requirements of game were discussed for an adcquate underitand~ngof the rumen micro-or- the recent Forest Service Range and Wildlife flab gani\ms We should be concerilcd w~thfoods that will Evaluation Research Workshop (Halls, in press). maintain a proper rumen population, and not necessarily with all toods ingested by clecr If a plant part is heavily EVALUATING FEEDING PRESSURE IN THE HABlTb eaten, but poorly digested, it cannot be considered a valuable component of the diet unless it supplies an Predicting habitat trends by studying the effect essential dietary element needed only in small quantities deer on vegetation depends on measuring abunda and utilization of plants or plant parts which are Some plants contain substances which reduce tile portant deer foods. Observing signs of browsing activity of rumen organisms (Nagy et al. 1964; Oh et al. woody plants tells little about utilization of the tc 1967, 1968 i. An excess of these substances reduces the food supply, except that if signs of browsing are plenti numbcr of rumen organisms, digestion is slowed, plant in any season except spring the supply of quality fo material accumulates in the stomach, and plant intake likely is too low. This does not solve the problem is reduced (Nagy, in press). An animal may die with determining trends before the habitat has deteriora its stomach full of undigested plant material of a single to a point where the trend cannot be reversed eas species which is digestible when eatei~in a mixed diet. Much of the deer's food comes from plants or pl Digestibility of plants from a given cnvironment must parts present in the habitat for short periods and can be measured using deer which come from the same kind be measured by surveys made once or twice a yc of environment and which have fed oil a variety of Mushrooms sprout and disappear within a short peri plants found in that environment. For instance, Oh et al. Fruits of many species ripen and fall from spring throl ( 1967 I foulld with in uitro trials that digestion of Doug- late autumn. Herbaceous plants grow, mature, and I las-fir was higher in rumen fluid from deer adapted to appear throughout the year. an area containing Douglas-fir than from deer adapted To compound the difficulty, signs of utilization are to an environment without Douglas-fir. Rumen organ- obvious-mushrooms, blades of grass, basal leaves, ; isms are specific for plants from specific habitats. small forbs often are plucked at ground level, with li Digestion of plants also differs by animal species. For sign of use. I observed several deer nuzzling thro~ instance, shecp are better able to digest some plants leaf litter of an Arkansas upland hardwood stand than arc deer, while dcer are superior at digesting others over an hour. Using 7-power binoculars at a dista (Longhurst et nl. 1968; Short 1963, in press). Digestion of 20 to 30 yards, I could not discern what they w may also be influenced by animal age, sex, activity, and eating. Immediately after they left, I looked for si health. of utilization. By crawling on hands and knees ; Several methods of determitling digestibility were dis- searching diligently, I found signs where they : cussed in the receilt Range and Wildlife Habitat Evalu- pulled small blades of grass sprouting under 2 to 3 inc ation Research Workshop (Pearson, in press; Short, in of litter. pressi. Samples for digestibility trails in cattle and sheep Little work has been done to establish techniques t hove been collected from esophagcal fistula samples adequately sample each type of plant and plant (Van Dyne and Torell 1964) to ensure that digestibility which occurs in a mixed diet. Measures to detect is related to the plants and plant parts consumed. vironmerltal trends must be made on frequently-in\ I could find no published reference to the use of toried plots where each plant part is mcasured or coun esophagcal firtulae on dcer The difficulty of the tech- Plant changes due to animal consumption must be 5 nique and the difficulty in taming animals for fistuliza- arated from changes due to seasonal progression. lion have probably held back its development as a tool closures paired with open plots would provide this CI In wildlife management, although it has been used by parison. The proper number, size, and distribution plots or plot pairs needed to sample each vegetative Harlow, R. F. category depends on variability in each category and 1961. Fall and winter foods of Florida whitetailed the accuracy desired. Additionally, consumption must be deer. Quart. J. Fla. Acad. Sci. 24: 19-38. related to plant production because the same degree of utilization considered moderate during good growth years Korschgen, I,. J. becomes excessive during years of poor growth. Time 1962. Foods of Missouri deer, with some management and effort required for adequate sampling will probably implications. J. Wildlife Manage. 26: 164-172. be considerable. Intensive management may have to Lay, D. W. be restricted to areas which sustain heavy hunting pres- 1967. The importance of variety to southern deer. sure. Southeast. Assoc. Game and Fish Comm. 18th The degree of utilization that food plants can with- Annu. Conf. Proc. 1964: 57-62. stand and still maintain their position in the plant com- munity must be determined. When plants decrease after Longhurst, W. M., Oh, H. K., Jones, M. B., and Kcpner, over-utilization, the food value of the plants which re- R. E. place them must be compared to food value of the orig- 1968. A basis for the palatability of deer forage plants. inal population so habitat change can be evaluated. We Thirty-Third N. Amer. Wildlife and Natur. Rc- should strive to reach a state of knowledge that allows sources Conf. Trans. 1968: 181-192. us to keep a running balance of the habitat's true food value. At present, this is more information than man- Marchinton, R. L , and Baker, M. F. agers can use because they cannot readily improve the 1967. Food habits of a deer as observed with the aid habitat or decrease the deer population. However, de- of telemetry. J. Ala. Acad. Sci. 38: 233-234. tailed habitat information will be necessary for intensive management needs of the future. Mosby, H. S. 1963. Wildlife investigational techniques. 419 pp. Ann Arbor: Wildlife Soc.; Edwards Brothers. LITERATURE CITED Nagy, J. G. Alexander, H E. Biological relations of rumen flora and fauna. 1954 Deer problems-new to Arkansas, an old story In Range and Wildlife Habitat Evaluation Work- elsewhere. Ark Game and Fish Comm , Fed. shop. USDA Misc. Pub. 1147. (In press) Aid Pub, 22 pp. Nagy, J. G., Steinhoff, H. W., and Ward, G. M. Bohman, V. R , and Lesperance, A L. 1964. Effects of essential oils of sagebrush on deer 1967 Methodology research for range forage evalu- rumen microbial function. J. Wildlife Manage. ation. J. Anim Sci 26: 820-826. 28: 785-790. Brown, D. 1954, ~~th~d~of surveying and measuring vegetation. National Academy of Sciences-National Research Council. Commonwealth Bur. Pastures and Field Crops 1962. Basic problems and techniques in range re- Bull. 42, 223 pp. search. NAC-NRC Pub. 890, 341 pp. Cook, C. W. Oh, H. K., Jones, M. B., and Longhurst, W. M. 1964. Collecting forage samples representative of in- 1968. Comparison of rumen microbial inhibition re- gested material of grazing animals for nutri- sulting from various essential oils isolated from tional studies. J. Anim. Sci. 23: 265-270. relatively unpalatable plant species. Appl. Mi- crobiol. 16: 39-44. Crawford, H. S., and Leonard, R. G. 1965. The Sylamore deer study. Southeast. Assoc. Oh, H. K., Sakai, T., Jones, M. B., and Longhurst, W. M. Game and Fish Comn~.17th Annu. Conf. Proc. 1967. Effect of various essential oils isolated from 1963: 9-13. Douglas-fir needles upon sheep and deer rumen Dietz, D. R. microbial activity. Appl. Microbiol. 15: 777-784. Definition and components of forage quality. Pearson, H. A. In Range and Wildlife Habitat Evaluation Work- Digestibility trials: in vitro techniques. In shop. USDA Misc. Pub. 1147. (In press) Range and Wildlife Habitat Evaluation Work- Dunkeson, R. L. shop. USDA Misc. Pub. 1147. (In press) 1955. Deer range appraisal for the Missouri Ozarks. Rice, R. W. J. Wildlife Manage. 19: 358-364. Stomach content analysis: A comparison of Halls, L. K. the rumen vs esophageal techniques. In Range Nutrient requirements of livestock and game. and Wildlife Habitat Evaluation Workshop. In Range and Wildlife Habitat Evaluation Work- USDA Misc. Pub. 1147. (In press) shop. USDA Misc. Pub. 1147. (In press) Short, H. L. Halls, L. K., and Crawford, H. S. 1963. Rumen fermentations and energy relationships 1960. Deer-forest habitat relationships in north Ar- in white-tailed deer. J. Wildlife Manage. 27: kansas. J. Wildlife Manage. 24: 387-395. 184-195. Short, H. L. Van Dyne, G. M., and Heady, H. F. 1967. Forage analysis for deer management studies. 1965. Dietary composition of cattle and sheep gra; Southeast. Assoc. Game and Fish Comm. 20th in common on dry annual range. J. Ra Annu. Conf. Proc. 1966: 15-18. Manage. 18: 78-86. Short, H. L. Van Dyne, G. M., and Torell, D. T. Digestibility trials: in vivo techniques. In 1964. Development and use of the esophageal fist1 Range and Wildlife Habitat Evaluation Work- a review. J. Range Manage. 17: 7-19. shop. USDA Misc. Pub. 1147. (In press) Wallmo, 0.C., and Neff, D. J. Sparks, D. R., and Malechek, J. C. Direct observation of tamed wild animals 1968. Estimating percentage dry weight using a rnicro- measure their consumption of natural for; scopic technique. J. Range Manage. 21 : 264-265. In Range and Wildlife Habitat Evaluation Wc USDA Forest Service. shop. USDA Misc. Pub. 1147. (In press) 1959. Techniques and methods of measuring under- Weir, W. C., Meyer, H., and Lofgreen, P. story vegetation. Symp. Proc. 1958, 174 pp. J. G. 1959. Symposium on forage evaluation: VI. The South. and Southeast. Forest Exp. Stas. of esophageal fistula, lignin, and chromo USDA Forest Service. techniques for studying selective grazing , 1963. Range research methods. USDA Misc. Pub. digestibility of range and pasture by sheep , 490, 172 pp. cattle. Agron. J. 51: 235-237. Problems in Censusing the White-Tailed Deer

James H. Jenkins and R. Larry Marchinton' School of Forest Resources University of Georgia Athens, Georgia

Conzparzsons are made between drzves, strzp flush- fairly recent titles in the literature: Lewis and Safley zng, pellet group counts, track counts, conventzonal (1967), "A comparison of some deer census methods in aerzal photography, trap-retrap methods, kzll data, Tennessee," five methods discussed; Dasmann and Taber and znfrared lzne scannzng The latter eqz~zpmentzs (1955), "A comparison of four deer census methods;" now avazlable at conszderable cost but zt has not been and Eberhardt (1960), "Estimation of vital characteris- adequately tested Although not a census technzque, tics of Michigan deerherds," in which at least three radzo telemetry helps to refzne baszc census methods independent methods were used in estimating population Infrared and mzcrowave radzometry may well be levels. fertzle fzelds for bzologzcal research Allen (1962) has pointed out that he feels that the main reason that the fishery biologist has made more The lot of the deer biologist has been hard and frus- progress in putting fisheries management on a scientific trating and will continue to be until more reliable and basis in the past 30 years is because basic censusing is convincing censusing techniques are developed. Sports- easier. A farm pond researcher can drain a pond and men continue to openly disbelieve the biologists' "popu- count and weigh his fish. The deer biologist has no such lation estimates" and although it may be heresy to sug- easy way out as yet. Adequate harvesting of deer is gest, there is often some justification for this. We shall often delayed until long after the carrying capacity has attempt to discuss the advantages, disadvantages, and been exceeded. Lewis and Safley (1967), in discussing validity of eight methods that have been used with a the "Peninsula deerherd" in upper Norris Lake, an area few peeks into the future with the newer technology of modest carrying capacity for deer and cattle, point now becoming available. We shall attempt to show the out that deer by the "best" census estimates must have role of deer movement and home range as a problem reached the incredible population level of around 129 facet of censusing. All States with adequate protection per square mile in 1957 before declining around 70 and a buck law have developed overbrowsing ranges, die- percent. It is obvious that deer are a valuable resource off or stagnation problems, underharvest and in general and it behooves us to devote more effort and expense wasteful and inefficient resource management. In gen- to developing better census methods. eral these conditions stem from inadequate and uncon- vincing census figures. This can be particularly acute CENSUS METHODS in areas of low carrying capacity such as the southern Drive census.-This method was widely used back in Appalachians. One of us particularly remembers going the Civil~anConservation Corps days when manpower on a sportsman "show-me" trip 20 years ago in north was available for the drives and for clearing firebreaks Georgia. The carrying capacity was undoubtedly around around selected areas, often I square mile in reasonably 15 to 20 deer per square mile and the population estimate level country. With experience these drives give good by several methods was around 25 to 30 per square mile. population estimates and may be very accurate. Some- The unbrowsed vegetation in the small fenced enclosures times in enclosures where conditions are unnatural freak (40 by 40 feet) was impressive and could be seen, counts result because the deer simply won't flush, but standing out, over 100 yards away in the bleak October in the field with disciplined drivers backed by skillful woods. Dead Smilax reinforced the conviction of the horsemen, good flushing and counting can be achieved. field men that a doe season was long overdue. Un- The main disadvantage to this method, assuming that fortunately, the most impressive thing about the 200-mile typical areas can be picked, is that the sample size is so tr~pthrough the heart of the deer country to a few of [ tiny due to the labor of setting up areas and carrying us who considered ourselves professional wlldlife man- I out the work. The need for relatively level terrain, agers was that no one actually saw a deer all day long roads, firebreaks, and 30 to 50 people severely limits ; The sportsmen representatives were reasonably under- this useful but exhausting method. In rugged mountain- standing and the relaxed seasons were grudgingly ap- [ ous or dissected terrain often devoted to wildlife, since proved. Many State conservation departments stay in no other uses are apparent, the method is often hopeless. "hot water" simply because they are trying to do a good job of conscientious deer management. Many of us have Strip flushing census.-Erickson ( 1940 has described given up on scientific management of a deerherd until a one-man strip flushing count method similar to the our methods catch up with our ideals. That this whole cruising method used for grouse earlier. One simply field is still in an unhealthy state is attested to by some calculates the average flushing distance, no mean feat,

~------and the number of miles walked. Lines are run at M-mile 'Research on which many of these conclusions are based is intervals on at least 4-square-mile samples by "stalking" supported by the College I+:xperiment Station of the Univer- at daylight and in the evening. sity of Georgia College of Agricult~~reExperirncnt Stations, MeIntire-Stennis Projects, and the Georgia Forest Research The advantages are that one or a few men can obtain Council. a rough estimate of population cheaply and quickly. On the other hand, it depends on "jumping" deer and as- at best, deer trapping is hard work and under m sumes that beds are randomly distributed which they conditions simply impossible. In general, this met are not. The method breaks down completely in the shows little practicability except where trapping is be "laurel thicks" so common in the Appalachians. The done anyway as for restocking. method is best for relatively level open lands and has Kzll figures.-This method requires data on the been used fairly succcssiully in Minnesota and Oklahoma. and numbers of animals killed by hunters and the st] Pellet group counts.-This method has been described ture of the wild population. In general, many stuc by Bennett et (~1.(1940). It is based on the assumption have shown that under heavy legal buck hunting arol that periodic accumulation of deer droppings bear a di- 10 to 12 percent of the population will be taken. Un rect relationship to population density. A defecation either sex hunting, the kill may vary depending u rate of 12.7 pellet groups per deer per day is generally many factors. Lewis and Safley (1967) found that used Th~ssame f~gurehas been used previously by percent of the herd could be removed under heavy hr range managers for sheep A line varying from 6 to 11 lng pressure. They concluded that this method was feet w~deand 1,000 or 2,000 feet long has been used most practical for use on typical management areas with an accumulation pcr~odof I month It has been Tennessee. used with fair efficiency from January through March This method, if it can be called a census method in the Appalachians. It is possible to use triangular- easy to apply since kill and age data are often availa shaped transects and painted lines. Mathematical com- and requires little extra work, but it is not applica putations can be worked out in order that the number to parks or in areas of closed seasons. At best it .i of pellet groups per line can he converted directly to remain a rough approximation and assumes that n deer pcr square mile. hunting mortality is minimal. The method is cheap and interesting. If approached right, refuge managers often elljoy making these counts Infrared line scanning.-There has been considera and a large amount of data can be obtained quickly and interest in the possibilities along these lines for v easily. Unfortunately greenery developing on the forest over 10 years. Biologists have been intrigued by floor and falling leaves lin~itsthe method to late winter. progress being made on forest fire detection using he Dung interfere particularly in the Piedmont and detecting crystals. The line scanners produce ima Coastal Plain. This census technique is almost confined which resemble photographs but are built up a line to late winter and in many ways this is the wrong time a time like television. Infrared film simply will to census deer, so this limits the usefulness of this meth- photographically pick up animal body heat but the 1 od. In hot areas or heavy rainfall areas it has little value. scanners will. Progress on fire detection, which is mi simpler than deer detection, has been summarized Traclc counts.-This method has been used extensively Hirsch et al. (1968). Croon, et al. (1968) discuss I in west Florida. Grid-like roads were driven on the whole subject in relation to big game censusing. 7 morning following an afternoon rain. Home ranges of method was tested on the George Reserve near P the deer were determined to be about 1 square mile. Arbor at midday on January 4, 1967. They knew Counts were compared to drive censuses. It was con- population of deer on this 2 square mile enclosure cluded that the number of deer crossing the roads per be about 101 animals. They also put out some deer linear mile approximated the number of deer per square pens. The imagery resulting from the overflight ir mile (Tyson 1959). Counts can be made by game rangers cated 98 animals. Ilowever there was little evergrl from vehicles, and in flat country with a stable climate, coniferous canopy and where it was present the pen] consistent data gathering is feasible. More recent work deer could not be discerned. For open country or ha in the mountains and Piedmont by at least two game wood areas aftcr leaf fall the advantages are obvic departments indicated wide variation and some groups You simply overfly, observe, and count the anim have given up on this method. It is possible to get a large sample and, under some c This method has the advantages of being cheap and ditions at least, excellent accuracy can be obtain easy to carry out on a large scale. However in some Infrared Thermal Mapping equipment (Blythe and I ecological situations it appears of little value. Harlow rath 1968) is available from Bendix Aerospace Systt and Downing 11968) have shown that due to variability Division. It weighs only 55 pounds and can be used it oftcn takes an impractically exhaustive amount of almost any small aircraft. It covers a range of emiss data to be statistically valid. This is particularly true from 0.7 to 14 microns wavelength. This goes well ab~ at the lower population densities. that visible to the eye or to photographic film but d Complete census by airplu?re.-This method has been include the wavelengths emitted by mammals (aroun used in the West and in deer "yards" and in east Africa to 14 microns). The usual flights are made at 1,000 f for a variety of plains antelope. Counts are made visu- and objects over 3 feet in d~ameterwith one-half deg ally or with aerial camera equipment. This method has centlgradc temperature d~fferencescan be detected ' bcen of no value in the southeast. equipment at thls t~meseems well adapted to deterrr I,incol~zindex.-This estimate is calculated by marking lng rlver water temperature differences and for trapped deer and checking their recovery on managed dctection. Bendix put on a demonstration in Febru~ hunts. Lewis and Safley (196'7) attempted this but hunt- 1969, at Tampa which one of us attended. We have E ing and trapping must be representative of the popula- been shown imagery of cattle and deer. The equipm tion. The amount of effort needed soon becomes imprac- costs around $40,000 and perhaps more for the up tical. Such a method works well for cotton rats where range where deer detection may be best. We discus a high percent of the population can be marked but, this whole subject with McCullough (personal commL cation) and he concluded that the method has all too that these areas although used only for a brief period many disadvantages at this time: (1) equipment is dif- during the year may contribute substantially to the ficult to keep operating; (2) it is expensive; (3) it won't support of the deer population. Therefore, for population punch through conifers; and (4) in open or hardwood estimates to be meaningful, a deer biologist must have country after leaf fall it has few advantages over con- information concerning how large an area is actually ventional aerial photography which is far cheaper and being censused. easily available. The development of transistorized radios for use on From an overall standpoint this method has good po- animals opened new doors for studies of movement- tential in open lands if animals are out at night and not ecology beginning in the late 1950's. Progress, however, in the daytime, and in deciduous areas after leaf fall. did not proceed at the rate which was expected or in It would appear to be of limited value in the southeast proportion to the number of researchers which became with its extensive conifer overstory. It will not record involved. Until very recently the preponderence of through clouds either. technical papers on telemetry in ecological studies con- cerned themselves with the technique and presented RADIO TELEMETRY relatively little new ecological information. This we think can be attributed to several factors. Although radio telemetry is not used to census deer First, the development of good telemetry equipment was populations directly, the data obtained from this tech- slow due to lack of the large amounts of capital needed nique are often necessary to refine basic census methods for basic development and technical research. A related sufficiently to obtain accurate population estimates. problem was the lack of communication between scient- Tester and Heezen (1965), studying the responses of ists and technicians in the widely separated disciplines three radio-equipped deer in relation to a drive census, of communications engineering and ecology. It is very pointed out that censuses of animal populations are usu- difficult for a single individual to have sufficient exper- ally predicted upon a knowledge of the animals' be- tise in both disciplines to develop equipment for his havior. They further stated that the technique of radio- specialized needs and to apply this equipment to ecolog- tracking provides an excellent opportunity for adding ical problems efficiently. This problem is beginning to to our knowledge of animal behavior and for observing dissolve as more technicians are becoming available with the natural responses of individual animals to census experience in meeting the special construction require- methods. Many of the previously discussed census tech- ments of biotelemetry systems. niques necessitate knowledge of such variables as activ- ity cycles, die1 movement patterns and the influence of A second basic problem is that researchers tend to meteorological factors on them. Any census technique, expect the equipment to take the physical work out of for example, which necessitates direct observation, pho- field investigations. This may be true to some extent tography or the other forms of remote sensing presently in the case of automatic systems such as those described available, requires that the animals be in a relatively by Heezen and Tester ( 1967 ) , and Cochran et al. ( 1965 ) . open area for accurate detection. Movement pattern These systems cannot be efficiently used to answer some and activity cycle information which allows the investi- of the questions which we are seeking answers to be- gator to predict the likelihood of deer being in such cause of their high cost, relative immobility, and certain technical problems discussed in the above papers. areas or the percentage of the populations to be exposed at a particular time becomes important. The value of Portable equipment is now available and can be ob- circadian movement patterns for use in calibrating track tained at reasonable costs. To have maximum utilization count census methods is also clear inasmuch as distance of portable equipment, however, requires a great deal traveled per unit of time is directly related to the number of physical effort. It necessitates, at least when studying of tracks made. It is important that wildlife biologists species such as deer which have variable activity cycles, be able to predict the movement and activity parameters working throughout the night as well as the daylight for the "average" deer in the population under study hours. Relatively few people are willing to give the during the unit of time tracks are being accumulated. kind of effort required to utilize the potential that the This is influenced by a great many variables ranging portable systems now available have. A final problem from meteorological conditions to population density. is that researchers expect too much from the equipment I in terms of dependability and performance and as a The location of deer in time and space also has im- portant implications in estimating their number in rela- result quickly become discouraged. A "rule of thumb" tion to their primary substrate, i. e., ecological density. which we have found useful in setting up telemetry Since much of the value of census information is related projects is to establish a goal for equipment performance which meets the minimum performance criteria which to estimation of carrying capacity measured in numbers can be established and still obtain some useful infor- per unit area, it becomes important to know the size of mation. the land resource base which supports the individuals comprising the population. The latter is sometimes Fortunately for those of us who wish to obtain detailed rather complex since deer may use certain portions of information of deer movement-ecology, whether this in- their annual range only on rare occasions, but these sel- formation is for refining census techniques or for other dom used areas may nevertheless be of considerable im- purposes, wildlife telemetry has "come of age." We portance in the maintenance of the animals. An example feel that it is now possible to obtain excellent and de- of this is exhibited by der in agricultural areas that pendable telemetry systems at reasonable costs and travel to and feed on agricultural crops considerably within reasonable lengths of time. We presently have out of their normal home range. It is important to know a number of very satisfactory telemetry systems func- tioning on deer in the southeast and during the past Dasmann, R. F., and Taber, R. D. several years have telemetrically analyzed the move- 1955. A comparison of four deer census met1 ment-ecology of more than 60 deer in various areas of Calif. Fish and Game 41: 225-228. Alabama, Florida, Georgia, and South Carolina (Marchin- ton and Jeter, 1967; Marchinton, 1968; Marshall and Eberhardt, L. Whittington, 1969). These animals have been located 1960. Estimation of vital characteristics of MicE in a wide variety of habitat types and a wide range of deer herds. Mich. Dep. Conserv., Game deer population densities. It is not the purpose of this Rep. 2282, 192 pp. paper to go into a detailed discussion of the movement- Erickson, A. B. ecology of deer. However, ~t should be pointed out that 1940. Notes on a method for censusing white-t, there is an increasing fund of telemetrically obtained deer in the spring and summer. J. Wil information concerning the movement-ecology of deer Manage. 4: 15-18. in a variety of habitats in the southeast and this infor- mation should be of considerable value for censusing deer Harlow, R. F., and Downing, R. L. in the areas where these studies have been made. 1968. Evaluating the deer track census methot Research needs.-All of the census methods leave much the Southeast. Southeast. Assoc. Game to be desired. We are by no means in a hopeless situation Fish Comm. 21st Annu. Conf. Proe. 1967: 3! but newer approaches would be welcome. Zimmerman (1969), in discussing forest fire detection, states, "Al- Heezen, K. L., and Tester, J. R. though microwave radiometry has the potential for op- 1967. Evaluation and radio-tracking by triangulz eration under conditions of complete cloud cover, and with special reference to deer movement. infrared does not, the current state of technology dic- Wildlife Manage. 31: 124-141. tates the choice of infrared in spite of this one major drawback." Both infrared and microwave devices are Hirsch, S. N., Bjornsen, R. L., Madden, F. H., and sensitive to heat where visual systems have no sensi- Wilson, R. A. tivity. Considerable effort has gone into research to 1968. Project Fire Scan fire mapping final re detect man-sized animals for counter insurgency efforts. April 1962 to December 1966. USDA Fc Small radar units open up new possibilities on the Serv. Res. Pap. Int-49, 49 pp. Intermour ground. Scent detecting devices have been developed Forest and Range Exp. Sta., Ogden, Utah. and capacitance sensors are available which can detect Lewis, J. C., and Safley, L. E. deer simply by their presence in a detector zone. We 1967. A comparison of some deer census method will need to carry out further radio telemetry studies in Tennessee. Southeast. Assoc. Game and a variety of habitats and seasons because most census Comm. 20th Annu. Conf. Proc. 1966: 56-62 methods depend in one way or another on the basic nature of the an~malmovements, range, and responses. Marchinton, R. L. Deer census methods have come a long way since the 1968. Telemetrically determined movement and deer drives of the Civilian Conservation Corps days in tivity parameters of white-tailed deer (Odc the thirties, but it would appear that newer technologies Zeus virginianus) in the Southeastern Ur may make more val~dand accurate methods available, States. (Abstr.) Bull. Ecol. Soc. Amer. 49: 6f hopefully in the near future. Marchinton, R. L., and Jeter, L. K. 1967. Telemetric study of deer movement-ecolog: LITERATURE CITED the Southeast. Southeast. Assoc. Game Allen, D. L. Fish Comm. 20th Annu. Conf. Proc. 1966: 1962. Our wildlife legacy. 422 pp. N. Y.: Funk and 206. Wagnalls Co. Marshall, A. D., and Whittington, R. W. Bennett, L. J., English, P. F., and McCain, R. 1969. A telemetric study of deer home ranges 1940. A study of deer populations by use of pellet- behavior of deer during managed hunts. So group counts. J. Wildlife Manage. 4: 398-403. east. Assoc. Game and Fish Comm. 22nd A1 Blythe, R., and Kurath, E. Conf. Proc. 1968: 30-46. 1968. Thermal mapping. Bendix Tech. J. (summer): 66-71. Tester, J. R., and Heezen, K. L. 1965. Deer response to a drive census determine( Cochran, W. W., Warner, D. W., Tester, J. R., and radio tracking. BioScience 15: 100-104. Kuechle, V. B. 1965. Automatic radio-tracking system for monitoring Tyson, E. L. animal movements. BioScience 15: 98-100. 1959. A deer drive vs. track census. Twenty-Fol N. Amer. Wildlife Conf. Trans. 1959: 457- Croon, G. W., McCuIIough, D. R., Olsen, C. E., Jr., and Queal, L. M. Zimmerman, E. W. 1968. Infrared scanning techniques for big game cen- 1969. Forest fire detection. USDA Forest Serv. 25 susing. J. Wildlife Manage. 32: 751-759. Wash., D. C. The Use of Models in Resource Management

Don W. Hayne Institute of Statistics North Carolina State University Raleigh, North Carolina

The terms "mode1.s" and "systems analysis" are on the basis of a more or less intuitive understanding new in wildlife management but the basic concepts of a complex situation, workers in these other fields are old. A model is an idea of how something works. have made real progress in scientific method. We must A systems analysis is accomplished for a complex study what they have done. process by breaking it down into parts or subsys- First, I believe there is a basic difference in the kind tems, the functioning of which can be studied sep- of objective. The newer view sees a model as a direct arately. The functioning of the whole process may, means to an end, which is prediction (and perhaps con- under fortunate circumstances, be understood as trol). The older view, though not really contradictory, the combined functioning of the parts and their was much less specific. It held that if we come to know interactions. Computer simulation of the system enough we will eventually learn to predict and control. mag be useful. In this view, the model is seen as part of the accumula- A possible systems analysis of the deer problem tion of knowledge, and this accumulation can easily of the southeastern States is descrzbed zn general become the main objective of research. features. It is suggested that a computer szmulatzon of the entire system zs probably imposszble zn the The different steps in building, testing and using a present state of knowledge, but that no doubt por- model may be listed as follows (Churchman et al. 1957; tzons can be simulated well enough to expand our Watt 1966, 1968). comprehension and ability to manage ratzonally A. Description 1. Formulating the problem This is a discussion of models and systems analysis 2. Constructing a mathematical model with special reference to deer. It is written for those B. Analysis who may be uncertain as to what a model is or what a systems analysis is supposed to do. 3. Deriving a solution from the model C. Prediction The terms "models" and "systems analysis" have been 4. Testing the model and the solution heard increasingly of late in wildlife management. These D. Optimization terms have been in fairly general use for perhaps 10 5. Establishing controls over the solution years, and were in technical use well before that. My 6. Putting the solution to work: implementation first point is that biologists have used the same ideas for a long time. E. Data acquisition. A model may be defined as "your idea of how things The steps are not always taken in sequence. A certain work." Anyone who has worked in deer management amount of data acquisition is required before a problem "operates from a model," as the phrase goes these days. can be formulated, and obviously some data are required In other words, he has put together in his mind a concept for testing the model. On the other hand, a successful of how the many interrelated factors operate, and has model dictates the kind of data required in the future. directed this analysis toward logical management of the To formulate an ecological problem and describe a resource. If he chooses to use the modern verbiage, model requires selection of the important factors from he has every right to refer to his intuitive appreciation the infinite number which operate in any biological situ- of, say, deer population dynamics, as his model To ation. Then some specification of how these operate is write it out as a mathematical formula does not make required, preferably in a quantitative form. Thus simpli- it any more or less of a model. It does make it easier for fication and abstraction is required at the start. Model others to understand. description may be deterministic, meaning that it will 1 Historically, ecologists have long struggled wlth the always have the same outcome if it starts at the same / quantitative aspects of complex situations and the idea point, or it may be stochastic, meaning that operation 1 that the world of nature is many-factored and highly of chance is allowed for. It is quite fashionable to claim j complicated is built Into their trainlng It is second that one will first develop the deterministic model and I nature for them to at least mentally abstract the im- then add the stochastic element. The latter step rarely j portant factors and to attempt to understand their mean- seems to follow. Further, it appears probably that the Equally, they learn early that nature is variable differences involved may be relatively minor if fairly 1 $:as been a little ddficult for biologists and statist~cians large populations are involved. I to understand what all the fuss is about in engineering After each process and relationship has been stated and bustness adm~nistration,from which most of the in a quantitative or mathematical way, then a solution recent emphasis on model building and systems analysis must be obtained for the whole model before we can have come But ~t is easy to become too complacent use the model, or even test it to see if it is realistic. about this. While ecologists have been trying to progress Since all the pieces of the whole are stated in mathema- tics, it would seem that a mathematical or analytic Population models, while only part of a resource n solution would be the answer. Occasionally, this may agement problem, are often of greatest interest to bi be possible, but often the assumptions required to allow gists. A population model is primarily concerned T a mathematical solution for the whole model are so re- the balance between mortality and reproduction, tal strictive and unrealistic that the resulting analytic solu- account of the factors which affect these two proces tion is not trustworthy biologically. Even our intuitive Watt (1968) has listed four categories, according to dej understanding of ecology involves such complexities as of complication of the model. to exceed the capabilities of mathematics. 1. Models which explain change on the basis of One difficulty is that biologists are rarely even fair relation between the number of reproducing mathematicians. One solution to this problem may be mals and the number of offspring. This kinc for them to learn more and more mathematics. This is model requires less information than the o. a good answer for those who can push deeper and deeper and may be used where there is less informal into mathematics, and do original work to solve biologi- but the capabilities are limited. cal problems. But this is rarely a practical solution for an applied problem. 2. Models which use information on the age struc, of the population relating present number: A second method of seeking an analysis of a model those of past time. More information is used is by simulation using cqmputers. Usually this exploits the model is still limited. The population ml the tremendous bookkeeping capability of computers used by Davis (1967) in his application of dyna which can be instructed to look at the status of a large programming to deer management, comes ur number of factors and according to instruction to calcu- this category. late the effect of each of these factors on the variable being followed, say the population and then report the 3. Models that consider many factors intrinsic to resulting change in the population over a short interval. population but assume extrinsic factors to ren This same process repeated many times allows many fac- constant. Watt refers to these as "complic; tors to be included for their effect on the biological steady-state models." Intrinsic factors are, h process and does not require that some known mathema- ever, allowed to include the character of the : tical function be found which describes the whole pro- vesting agency, the exploitation rate, and nat. gress of the population in time. As an oversimplified mortality. example, a computer would be ideally suited to keep 4. These models are more complex and thus n account of your personal money resources, your bank flexible. Both intrinsic and extrinsic factors I account, and your household cash, adding and subtracting be included, with as many environmental fac as you earn and spend, and capable of calling up a being included as may be desired. These are balance at any moment. Similarly, it is able to consider steady-state models. They tend to have high the progress of a biological process and providing it is mands for information, being in principle properly instructed, to add or subtract over many time limited as to complexity. units. A system, to quote Watt (1966) is "an interlocl There has heen a tendency to feel that this capability complex of processes characterized by many recipr of the computer will free biologists from the need to cause-effect pathways." A systems analysis is an atte know advanced mathematics, and provide a simple man- to understand a system well enough to manipulat ageable tool which can do just as well. Experts in the as a whole. A systems analysis starts with the const field tell us that this is not true. They warn that study tion of a large and complex model to describe the sysl of the operational characteristics of computers has in The rest of this paper discusses systems analysi itself become an advanced and exacting branch of math- the form of a proposal for a regional analysis of the I ematics. For example, the common approach to biologi- problem of the southeastern States, say those for W' cal proccsses mentioned above, that of dividing time into short units and considering what happens from one unit administration of Federal Aid funds is carried through the Atlanta office. At present there is no F of time to the next, is particularly susceptible to the progression of errors. That is, an error which might be pect of initiating such a study. But in the contex minor in one calculation is repeated with each calcula- this conference, a discussion of systems analysis tion and because there are so many calculations the best follow this proposal. error may grow to important magnitude. Accompanying growth of its deerherd, each State felt the need for added biological information and In spite of these cautions, the only path open for rnany carried on investigations at varying levels, supportec problems seems to be through computer simulation The State and Federal Aid funds. In some States, the am1 test wlll be how useful the method may be in the long of information compiled over the years has reac run major proportions. Most of the biological investigat After building a model wc must test it against reality. which would seem appropriate have been carried One way is to vary the inputs to the model and observe at least once. Now is the time to consider what is rc whether the results conform to nature. A better way is needed for management, and what opportunities to make a prediction based on the model and confirm exist for cooperation. this with an experiment, or by gathering information on Regional aspects are important. Ecological similar the particular point. A good model is one which makes occur over several States and social and economic fa( reliable predictions. are superficially similar, yet some management prob are more acute in some States than in others. A regional 1.2. Identification of the major subsystems making analysis could explore these differences. up the whole system, and the component parts making The management of the white-tailed deer is indeed up these subsystems will be required. For example, the an extremely complex activity made up of many inter- deerherd, its biology and population dynamics, comprise locking processes with multiple cause-and-effect path- the best known subsystem, and the sex-age subgroups ways. The whole problem is too large and complicated may be viewed as components of this subsystem. There to be mentally grasped as a whole, even though smaller are other subsystems, well known though lcss clearly parts are being handled very well on an intuitive basis. differentiated, which include, for example, the inter- There is great need to organize the available information action of the public with the wlldlife department, whlch into an understanding of the problem as a whole. The occurs on the one hand through educational activities methods of systems analysis seem to be an appropriate and on the other through pressures from Interest groups basis for such an organization. and through the legislature. Others are the regulation- The basic concept of systems analysis, old to biologists, producing system, the data acquisition systems (con- is that a process of great complexity can be attacked by cerncd with biological information and hunter statisticsi, breaking it into separate compone~ltunits, each of which and the system of laws and enforcement. This part of can be studied and understood at least as to the relation the investigation may be developed partly through study between inputs and outputs, whether or not all the inner of the hterature but mostly in discussioi~swith State workings are colilpletcly known. To this has been added personnel A model is needed for each subsystem an emphasis on the quantitative, a certain systematic I 3 Interactions and relationships among sub5ystems organization of the problem, and the conviction that any and components must be specifled An output of one complex process can be computer-simulated if useful component becomes an input of another (unless it is an approximations can be set up for the functional relation- output of the whole system) For example, what 1s thz ships. Further, for management processes there is em- effect of increasing enforcement effort on the biology phasis upon a clear definition of the desired outcome (or of the deerherd? Identification of the important path- "output of the system"), a point of obvious relevance ways of influence will be crltlcal, and must be developed to management of natural resources. Application of this through study of the system and in discussions. method may be discussed under the four elements of 1.4. There must be an identification of the decision Description, Analysis, Prediction, and Data Acquisition. points for deer management and the criteria for making A fifth element, Optimization, completes the analysis. decisions, including the influences bearing on the de- I. Description. The primary phase of this study cisions. These points must be explored in discussion would be a description of the system of deer management with State administrators. in each State. This would be carried out by a full-time 1.5. The important inputs into the whole system from specialist who would be what has aptly been described outside, and the outputs of managed deer hunting must as a "circuit-riding brain-picker." He would visit the be identified. This step also includes defining the boun- individual States and discuss deer management with daries of the whole system. Inputs from outside will in- technicians and administrators, developing from their clude, for example, the influences of weather on deer concepts and practices a statement of the whole problem and hunters and the effects of changing land use prac- in that State in the form of a system. He could start tices. discussions with a simplified version of a management system and elicit suggestions and changes to develop a better model of the local system. Eventually these State I1 1 This implies systematic study of each component models would be combined into a gel~eralizeddescription of the system, concerning what is now known of the of a State management system, with features left uilique relationships between Inputs and outputs, how ~tsfunc- to a State where necessary. This process of description tioning can be quantified, and what more needs to be known of its nature in order to build it into a simulation ! can be classified under five headings. model. Depending upon the component being studied, 1.1. Definition of all the objectives of deer manage- this step of analysis may lie anywhere between a desk ment will be needed to produce a clear statement of what study of literature and data, at the one extreme, and at deer management is expected to accomplish. The com- the other, gross speculat~on111 discussion wlth lnformed plexity of the deer problmi is apparent when one at- 1 persons With some of the components, we might be tempts to state the outcome expected of successful man- able to do no better than point out the need for compe- / agement. Even if one accepts some one of the different tent study For example, what is really known of the measures of hunting success, then what weight is to be / quantitative relationship between enforcement effort 1 given, for example, to tourists viewing the animals or and illegal hunting? ! to reduction of forest and crop damage? It may prove 1 impossible to set up a single objective or a set of alter- 11.2. A quantitative model of deer population dy- native single objectives, and the desired outcome may namics would be one of the principal objectives of the have to be stated as a compromise among several some- study. There is much information already available on what contradictory aims, expressed as a weighting of this subject, but no specific model general enough to the several objectives. But a specification of manage- include the features found in all States. One of the most / ment objectives will be necessary. In fact, it scarcely important questions to be answered here is what kinds I seems necessary to invoke the term "systems analysis." of data must be obtained on a continuing basis to actu- to point out the necessity of defining the objective of ally use a model of population dynamics in management. deer management. 111. Prcdiction. 111.1. A graphical rzpresentation or chart of the sys- in the future as routine information to support man tem of decr management in each State should be drawn ment decisions. up, based on discussions with the biologists and admin- IV.4. Cooperation among those States with sin istrators. This would be a first approach to simulation problems, climates and habitats promises to pro1 and could serve as a model for further discussion of efficiency, both in sharing the conduct of investigaf dctails. Such a chart would indicate the kinds of effects and in some of the routine data gathering. Explor: to be expected from changing a component and would of this possibility should be a principal aim of the st thus facilitate at least qualitative prediction. In summary, the process of systems analysis airr 111.2. Although computer simulation of the entire set up a model for such a complex entity as deer r system of deer management for a State must be the agement, by breaking it up into a number of sm, eventual objective of a successful systems analysis, it is units which can be individually studied. After a q doubtful that at the present time all factors can success- titative model has been set up for each of these sm~ fully be quantified well enough to permit such complete units, and the relationship between inputs and out simulation. But a minimum objective would be to dis- specified for each, then the whole can be reassem cover what factors we need to know more about in order from the parts. to attain this ultimate objective, and to set up a simu- A computer simulation may be set up if enoug lation for any subsystem whenever enough is known. known about all the parts. Then the simulated sy: 111.3. A model of deer biology could probably be may be tested to determine how well the predict programmed for computer simulation in a fashion general match the real world. If the goal is management, enough to be adaptable to the particular needs of each the objective will be to manipulate the system to pro1 State. Enough is known to allow reasonably successful the optimum output. simulation of the biology of a herd under management. Even if there cannot be a successful simulation of This could allow use of the computer in "experimental" whole system of deer management, the careful cons1 investigation of the effect of varying the different factors tion of models for subsystems and the attempt to q affecting the population dynamics, and investigation of tify their interpretation will yield worthwhile divid the outcome for different management plans. in the form of increased understanding. IV. Data acquisition. IV.l. One objective of the study would be to assemble LITERATURE CITED a list of what information now exists in the region. This could be carried out during visits to the States, when Churchman, C. W., Ackoff, R. L., and Arnoff, E. L. detailed notes could be made on the condition, accessibil- 1957. Introduction to operations research. 645 ity, and technical coverage of the stores of data previ- N. Y.: John Wiley and Sons, Inc. ously gathered and now in the file, as well as those now being gathered. Such a listing of information should be Davis, L. S. 1967. Dynamic programming for deer manager made available to all the States. planning. J. Wildlife Manage. 31: 667-675 IV.2. A check list of all reports on deer management and biology could be assembled, showing whether pub- Watt, K. E. F. (editor) lished or not, and whether available. 1966. Systems analysis in ecology. 276 pp. N. Y. IV.3. An important objective of the study would be London: Academic Press. to decide what further information will be needcd to Watt, K. E. F. understand both deer biology and the other factors of 1968. Ecology and resource management. 450 management, and also what data should be gathered N. Y.: McGraw Hill Book Co. Sociological and Economic Considerations in Management of White-Tailed Deer

E. L. Cheatum lnstitute of Natural Resources School of Forest Resources University of Georgia Athens, Georgia Lonnie L. Williamson Southeastern Cooperative Wildlife Disease Study Department of Pathology and Parasitology School of Veterinary Medicine University of Georgia, Athens A. Sydney Johnson Institute of Natural Resources School of Forest Resources University of Georgia, Athens

Many problems in deer management will develop ning public acceptance. It is a "people problem" and as human populations increase and social patterns thus a political one. He must operate in the context change. Some aspects of three problem areas are of traditional and strongly entrenched attitudes, and any discussed: the implications of urbanization, quality victory he achieves is likely to be a compromise. in recreational hunting, and the economics of re- There is ample evidence to support the thesis that the source use including conflicts of interest. Precise, accelerating urbanization of our society will have pro- objective studies of the value of deer and other wild- found effects on policies and their implementing regu- life resources are needed. Administrators and man- lations in future deer management. Attitudes of the agers are urged to take a broader view of the deer urban population will increasingly dominate legislative resource and its relevance to the needs of society and decisions and the policies and programs promulgated by to diversify their management objectives to max- wildlife management agencies. Greater involvement in imize benefits and minimize conflicts with other resource issues by the urban-oriented populace will re- resource interests. sult in a broader base of interest in wildlife for its esthe---~ In considering how to treat the socioeconomic aspects tic and cultural values and an increase in antihunter of deer management in the Southern States, we have sentiment in general. determined to limit our discussion to three broad prob- The patterns being planned for interspersion of open lem areas. We doubt that any one State would claim it space, green belts or parks for recreation to accommodate had the answers to any more than a small fraction of the burgeoning urban and suburban communities will the multiple questions rearing their heads in each area, accelerate the loss of wildlife habitat on which any form I and we resisted the temptation to launch a "question- of firearms hunting wi!l be tolerated. na~re"to find who had the answers. As a matter of fact, we have concluded that for the purpose of this Scrublands, forest lands, and low grade agricultural panel, and for this assigned subject, we can perhaps be lands reverting to wild herbaceous and forest covers will of greatest service by questioning the validlty and utllity command the deer manager's major attention. Here he will continue to be most closely involved with rural of some of the conventional interpretations of economic data on deer hunting and carcass values, and pointing hunters and landowners. However, even here the de- up some sociological questions surrounding deer man- sires and needs of the urban populace will be influential, agement, answers to which are sorely needed by wildlife and the manager will have to learn to work with them i also. management agencies if their programs are to be more effective in the future. The wildlife agency that continues to depend almost entirely on revenue from the sale of licenses to finance The areas we have selected for discussion are: 1 its programs of acquisition, habitat management, law The implications of a rapidly urbanizing Amer- I. enforcement, and education finds itself on the defensive I ica to modern deer management il when recreational hunting is threatened. An increasing 2. 1f The challenge of achieving quality in recrea- problem in the cities will be one of convincing the ur- 1 tional hunting. banite of the value of hunting. In the United States, 1 3. Economics and resolving conflicts of interest only 3.4 percent of the population living in large cities i with regard to resource use. are hunters (U. S. Department of the Interior 1966). i i Many of the nonhunters see no great relevance to wild- i DEER MANAGEMENT AND URBAN SOCIETY life generally and regard hunting as cruel and even ata- There is probably no experienced wildlife manager vistic. The bulk of agitation for stricter gun control iI employed in a State or Federal regulatory agency who legislation comes from the urban centers where the dis- I will not agree that his major problem in implementing tinction between the criminal with a gun and a legitimate technically sound deer management programs is In win- hunter becomes fuzzy. Resource issues arc being settled increasingly on social, Beyond a point, quality of the hunting experience ethical, and moralistic bases. We must be able to show clines as the number of hunters increases. The ultir that the sport of hunting has broad social values as im- in low-quality hunting has already been reached portant for the nonhunter to understand as for the some of our public hunting areas. Excessive devc hunter. The economic importance of controlling deer ment has resulted in an atmosphere of artificia populations through regulated hunting seldom occurs Hunting pressure is so great that hunters can finc to the urbanite. He may view the wildlife agencies' refuge from other hunters. There is no opportunit special deer seasons as "gimmicks to sell more licenses apply knowledge of woodcraft and stalking skills. Ei and provide an outlet to a sadistic lust for killing." Are mentation of hunters, assigned stands, numbers, perr we doing a good job in answering this kind of charge and badges add to the artificial atmosphere. or preventing its gairiing credibility? In our information Disgusted, many an experienced sportsman hang: and education programs, it is necessary to emphasize that his gun unless he has hunting privileges on private la an important part of the wildlife agencies' responsibility He is replaced by the novice who accepts such condit is to control excess populations. Hunting is a manage- as part of the sport because he has never known anytl ment tool to this end, snd the recreational values may better. be only secondary. We should level with the public on Aldo Leopold (1949, 1953) eloquently appealed this fact. consideration of quality in game management and Man is by nature a hunter. Anthropological evidence what he called "split- values." I-Ie appealed for g indicates that man evolved from an apparently unique departments to take the lead in fostering "the distin group of predatory apes. Indeed, it appears that his American traditions of self-reliance, hardihood, wl predatory, aggressive nature was primarily responsible craft, and marksmanship." for the development of the brain, the use of tools and There are steps that State wildlife agencies can fire, and the general evolutionary success of the human should take to achieve greater quality in recreati~ spccies. Physiologically, man as a species is still equipped hunting. They should begin placing as much emph for life as a hunter. However, in the urban environment, on providing pleasant hunting experiences as they the hunting instinct may be expressed and adequately now placing on harvest. Through their public relat satisfied by most in their pursuits of making a living. programs game departments should conduct educati~ In others the hunting instinct is satisfied by ritualization efforts emphasizing sportsmanship and tradition in the form of sport hunting. Some hold that this con- placing less emphasis on the kill. tributes to the iiiental and physical well-being of the par- ticipants. Furthermore, codes of ethics, traditions, and General tax funds are needed to supplement f~ restraints have definite character-building influences. provided by hunters to eliminate the necessity of These character-building qualities must be preserved pending upon numbers of licenses sold to provide and nurtured if hunting is to survive as a sport. enue. Seasons should be as long as possible to avoid con In an increasingly violence-conscious, urban-oriented trating hunting effort in a short period of time. Pu society, we may be sure that antihunter sentiment and hunting areas should be diversified with some a demands for rigid gun control will increase. Is it possible developed to accommodate maximum use and some a for game managers and hunters to convince the public undeveloped and with limited access and contra that hunting has broad social values, that to some it is numbers of hunters (e. g., primitive weapons arc essential to mental and physical well-being, that it has Regimentation should be held to the minimum neces: character-building influences that pay off in reduced to preserve other values. Habitat on such areas shc crimes of violence and fewer social problems? There be managed in such a way as to preserve as nearl: is need for sound statistical data to support these claims possible a natural appearance with a diversity of spc if the antihunting public is to be convi~lced of their and a minimum of artificiality. validity. Administrators and biologists should give more sideration to local hunting traditions in recomment QUALITY IN RECREATIONAL HUNTING regulations concerning seasons and hunting meth As the pressures of mass use increase, the problem of Local traditions and codes enrich the sport of hun maintaining standards of quality in outdoor recreation and provide charm and color and diversity of hun will become more critical. Most of us will agree that opportunities. Unless there are sound biological the greatest social values of the white-tailed deer are administrative objections, local traditions should be recreational, cultural, and aesthetic. To many, the great- tured. est realization of these intangible values is through the More attention should be given to trophy values. il quality hunting expcriincc. agement on a maximum sustained yield basis does What is quality hunting? For purposes of this paper, allow animals to reach maximum size and develop trc we accept the statement of the Mississippi Flyway Plan- antlers. Research is needed to determine method. ning Committee 11961) that quality hunting is "charac- controlling deer numbers while allowing some bi terized by reasonable solitude. primitive surroundings, to reach trophy size. rugged exercise, suspense, excitement, and a chance to Also, it is generally recognized that prey species 1 pit the skill of the hunter against the innate cunning a genetic need for predation. In the South, hunters 1 of the prey, resulting in a hunt to remember with satis- largely replaced natural predators of deer, but the u faction whether or not a full legal bag is taken." regulations and harvest methods select against trc animals. American wildlife biologists have generally spokesman for the New Hampshire Motel Owners Asso- not been impressed by genetic factors in game manage- ciation, appearing before a legislative committee in ment. But European game managers place great import- opposition to proposed deer legislation, estimated that ance on selective harvest and deerherds. Funda- a 10-day shortening of the season would result in a mental studies are needed to evaluate genetic effects of minimum loss of $100,000-$200,000 to members of his harvest practices. organization. Another motel operator estimated his loss Private lands probably provide the best opportunities to be $500 per day in addition to the layoff of nine for quality hunting. It is axiomatic that "mass use im- employees. A restaurant owner at Colebrook estimated pairs quality" (Leopold 1949, 1953). Private property, his income from hunters to be $40-50 per day." functioning as a form of territoriality restricting use, Mangold (1965) stated: "Official reports indicate that averts destruction of quality values (Harttin 19681, and deerhunters in New Jersey legally harvested 8,029 deer 75 percent of the forest lands in the South are in private in 1964 with an average expenditure of $736.50 per dcer. ownership. Game departments should investigate means We have an estimated deer range of approximately 4,830 of providing technical advice for private landowners square miles or a little over 3 million acres, which aver- wishing to form game pools or cooperatives or clubs ages $2 per acre spent hunting deer." wherein quality hunting opportunities would be in- There are many other similar reports which show that creased. deer are economic assets to the community. Of all the benefits, however, how much is cancelled by crop dam- ECONOMICS AND CONFLICTS OF INTEREST age, tree damage, fence damage, vehicle damage, personal Deer populations in many of tile Southern States are injury, and disruption of other forms of hunting and now expanding rapidly and interest in hunting them outdoor recreation in general? Even though many people is growing. Studies on the economic impact of this situ- may think of deer as a priceless game resource, others ation are interesting, and they demonstrate that deer at the same time may view them entirely as pests. If hunting can make a very significant economic contribu- deer management is to successfully contend with these tion to an area. For example, a small six-county area varying attitudes, it may be necessary for management in Georgia receives an injection of more than '4 million personnel to become more familiar with these attitudes dollars into its economy each deer season, three-fourths and the people expressing them. The result may be that of this coming from hunters not residing in the area deer should be considered as more than just a hunter's (Almand 1968). Phillips (1965) reported that three par- quarry or a photographer's subject with positive aspects, ishes in northeastern Louisiana receive an influx of but also that they have negative inferences in certain more than $158,000 during each 5-day deer season. situations. Ramsey (1965), in making an economic comparison of Surveys of hunters in five Southern States (Alabama, deer to domestic livestock in the Edwards Plateau region Georgia, Kentucky, Tennessee, and Texas) indicate that of Texas, said: "Records from the Kerr Wildlife Manage- a majority of license holders never hunt deer and less ment Area ind~catethat the net return per an~malumt than 20 percent of the total hunting effort is for deer of deer can exceed that from livestock if the decrherd (Durell 1965, 1968; Kelly 1967; Legler [n.d.]; Marshall is adequately harvested " From Llano County, Texas, and Payne 1968; Wilke 1962). Most hunters, therefore, in the same region, Teer et al. (1965) collected data are not directly benefited by the South's growing deer whlch show that in 1961, 25 ranches conslstlng of 47,217 populations. How then are these nondeer hunters af- acres received a comblned income of $57,395 or $1 22 fected by expanding deerherds? per acre from deer hunters. The multitudes which hunt rabbit, fox, and raccoon Emerson (1968) reported hunters in the Tennessee generally require dogs for best enjoyment of their sport. Valley area to spend approximately $20 m1111on each year There are few hunting dogs which will not run deer. enjoying their sport. This creates conflict, therefore, which is quite prevalent The Uwharrie Deer Restoration Project, which was in many areas where hunters spend great amounts of merely an idea in North Carolina in 1944, now enriches time trying to recall thzir dogs from a deer chase. These the local economy by more than $100,000 each gear people undoubtedly are left with a bad taste when they (Wilson and Thompson 1964). realize that a disproportionate amount of their money Deer hunters in Vlrgin~aharvested 24,934 dcer durlug may be spent on managing an animal which they consider the 1966-67 season (Cross, personal communscatlon ) It a pest. the figure of $400 per animal (Almand 19681 1s applied, Much has been said concerning possible detrimental dcer hulltlrlg in the State contrsbuted almost $10 million effects of hunting dogs on decr populations, yet little to the economy durmg that year hope is offered the hunters who utilize hounds and are Watson and Whitehead 11967) wrote: "Wildlife mali- increasingly plagued by what they collsidcr to be en- agernent in our town of Crossvillc (Tennessee ) is a prom- croaching deerhcrds. Other land management conflicts inent business." Dr. Watson is Mayor of Crossville. occur between deer and crop, timber, orchard, anti Mdny northern areas have for yeais enloyed the vzable flower garden interests. The critical attitudes of these Income produced by wild deer populationi In New groups should provoke serious corlsideration of whether Hampshire, 1962 hunt~ngvalues were asses5ed at over a high population of deer is desirable for all areas. $12 n~illion,of whlch deer accounted for at Iea\t 90 Would the exclusion of deer from selected areas help percent (Silver 1968) She stated "The ilnport,lnce of resolve somt conflicts? Are game management prograrrts decr huntll~gas a factor in the cconon1y of the non~ndu\- building a white-tailed deer monoculture in the South:7 trial 'North Country' was emphdsited 111 1963 A Are many game biologists criticizing forestry for pine plantations while at the same time establishing "deer Emerson, F. B. plantations?" 1968. Tennessee Valley wildlife: an outlook for It is good for deer managers to recognize these con- year 2000. TVA Div. Forest. Develop. Rep. 2' flicts and to expect that occasions will arise when they Ilardin, G. will be asked, "Why have deer at all?" McNeil (1962) 1968. The tragedy of the commons. Science stated: "Until we learn to make better assessment of 1243-1248. nonmarket values of deer, we will run the danger of Kelly, W. F. allowing deer populations to become so large that costs 1967. Game kill survey. In Ala. Dep. Conserv. A will exceed benefits." Progr. Rep. Fed. Aid in Wildlife Restora' pp. 95B-132B. It may be beneficial for deer managers to more criti- Legler, E., Jr. cally evaluate their economic data and better understand Tennessee survey of 1961 hunting and fis its full meaning. If a deer in area A, for example, costs [n.d.] activities. Tenn. Game and Fish Cornm., N the hunter $100 to harvest while a deer in area B costs $200, what conclusion is to be drawn? Are deer from ville. 79 pp. area B actually bringing more money into the commu- Leopold, A. nity than those from area A and thus are greater eco- 1949. A Sand County Almanac. 226 pp. N. Y.: Ox nomic assets? Are the hunters in area B paying more Univ. Press. for their sport? Leopold, A. 1953. Round River. 173 pp. N. Y.: Oxford Univ. P: In efforts to justify the costs of management, a game McNeil, R. J. manager may be tempted to use the cost figure for har- 1962. Population dynamics and economic impac vesting a deer in his State for calculating the economic deer in southern Michigan. Mich. Dep. Cons worth of the deerherd. This is a treacherous course Game Div. Rep. 2395, pp. 90-91. to take. Mangold, R. E. The cost, for example, of harvesting a deer from low 1965. What's your deer worth? N. J. Outdoors 16 1 density populations may be extremely high, whereas 12-16. the worth of the overall herd may be low. Conversely, Marshall, C. M., and Payne, R. L. as deer numbers increase, per capita harvest costs 1968. Wildlife harvest surveys and economic sur. could decrease, yet the value of the herd would become of sportsmen's demand by species and comp much higher. tive cost. Ga. Game and Fish Comm. Job C To further assure correct evaluation, meat value, pletion Rep. 12 pp. which is a rarely considered economic factor, must not Mississippi Flyway Planning Committee. be overlooked. Almand (1968) stated that solely from 1961. Waterfowl management and quality hunl a food standpoint, the meat from one deer represents Miss. Flyway Counc. 47 pp. at least $75 compared to an equal amount of meat bought Phillips, P. H. at a butcher's shop. Although most deer hunters may 1965. The economic impact of the Louisiana I view venison as being secondary to recreational benefits, hunter on the communities surrounding the the same venison is a primary stimulus for legal and cago Mills Game Management Area. M. S. ' illegal harvest in many underdeveloped areas. Deermeat sis. La. State Univ. Baton Rouge. 43 pp. can be an important quality protein source for people Ramsey, C. W. in such areas who survive on a subsistence diet. Further- 1965. Potential economic returns from deer as c more, this meat is produced on lands generally unsuitable pared with livestock in the Edwards Pla' for agriculture and domestic livestock production. region of Texas. J. Range Manage. 18: 247- It is evident that a comprehensive economic assessment Silver, H. of white-tailed deer must be derived by utilizing both 1968. Values of New Hampshire deer. In The wl positive and negative values. Future success in deer tailed deer of New Hampshire, pp. 174- management will be dependent upon a broad viewpoint Concord, N. H.: N. H. Fish and Game Dep in which deer are considered in relation to all other Teer, J. G., Thomas, J. W., and Walker, E. A. forms of land use and the overall social outlook 1965. Ecology and management of white-tailed I LITERATURE CITED in the Llano Basin of Texas. Wildlife Mon 15: 53-55. Almand, J. D. 1968. Wildlife: how valuable? Ga. Game and Fish United States Department of the Interior. l(4):10-12. 1966. National survey of fishing and hunting, 1 Durell, J. S. 76 PP. 1965. A study of Kentucky hunters who hunted only Watson, J. H., and Whitehead, C. J., Jr. in their home counties. Southeast. Assoc. Game 1967. Community economics and wildlife man and Fish Comm. 16th Annu. Conf. Proc. 1962: ment. Fontana Conserv. Roundup Trans. 8 171-175. Wilke, L. A. Durell, J. S. 1962. Texas survey of hunting and fishing. Tex. G 1968. A brief study of hunters and the owners of the and Fish 20 ( 1 ) : 24-25. land on which they hunt. Southeast. Assoc. Wilson, K., and Thompson, 0. Game and Fish Comrn. 21sl Annu. Conf. Proc. 1964. The economic importance of the Uwharrie 1967: 81-87. herd. Wildlife in N. C. 28(4): 18-20. Regulatory Legislation and Public Attitude on White-Tailed Deer Management

Leslie L. Glasgow School of Forestry and Wildlife Management Louisiana State University Baton Rouge, Louisiana

Today, as zn the pust, econoinzcs and polztzcs large- in the face of long-held trad~tionsthat game was for the ly znfluence the shaplng of game n~anugenzentpolz- taking. czes Often, the sound recommendatzons of bzologzsts Since those bleak days enlightened management-and have been overruled by polztzcal pressure Tradz- circun~stance-has restored deer over much of their orig- tzons, land-use competztzon, recreutzonal needs, crop inal range, even adding thriving herds where none ex- damage, fee huntzng c~ndpolztzcal payoffs are among isted before. But political involvements have accornpan- the many factors that znfluence regulatory leqzsla- ied the transition and are with us today in even greater tzon and gume commzsszon polzcy Better cominzLnz- variety and complexity. In fact, I think it is a fair state- catzon and educatzon are needed to get a publzc un- ment to say that modern game management seems less derstandzng and support of programs that best meet a matter of biology and more a matter of allocating re- the needs of the people. sources among competing demands. This process of de- cision-making places deer management fully in the po- In the evolution of game management as a science, litical arena. some are inclined to view the growing involvement of socioeconomic and political factors as recent develop- I will refer again to Wisconsin, and the Lake States ments. And with it we look back a couple of decades area generally, to illustrate a point. Following logging to the good old days when biological facts seem to be and fire suppression in the late 1800's, the deerherds the main consideration in shaping the destiny of wild expanded rapidly. The era coincided with a return by things. But if affairs seem more complicated today- city dwellers to the out-of-doors. The backwoods became and they probably are--it is mainly in retrospect. In the a place to get away from it all. Old timber trails afforded game management equation, people have always been acccss to deserted logging camps, and hunters and fisher- prominent, and the frustrations they add through eco- nlen moved in. nomic and political pressures have always been with us. The next generat~onof sportsmen were accompanied In wildlife management, no other segment of th~sre- by the~rwlves and kids The crude camps and shacks source-unless IC IS waterfowl-- has felt the byplays, the were replaced by "civilized" hous~ng,waterways became economic haul~ngand pol~t~calpull~ng that have come scenlc spots for a profus~onof motels, hotels, and lodges to our deerherds And I say "deerherds" In a sweeping The summer resort ~ndustryhad arrlved For the deer, context because no State has been free of management and for those who would manage them, it brought prob- cons~derationsfar removed from good blology alone lems-political problems. Let me begin in Wisconsin-not because it is unique in More than being just venison, deer had esthetic values. having more or bigger problems, but mainly because it For a child of the pavement, the sight of a doe with her has done an unusual job in documenting its history of fawn is recompense enough to assure a successful vaca- deer management. We can see well enough that political tion. (The same could be said for most city-bound adults.) factors were present in Year One of beginning manage- Resort owners were quick to see the dollar values in a ment. flourishing deerherd. They wanted deer around--in quantity--and naturally enough they took to the stump Consider for a moment the attitudes of the first sett- and to the legislative halls to "save" the deer. In pur- lers, and of the succeeding gcnerations that followed for suing their cause, they had the enthusiastic support of at least 100 years. This was a new country then and tavern keepers, restaurateurs, gas n~erchants,and others nature had endowed it abundantly. Game was for the who benefit from a flourishing tourist trade. Eventually taking-not for sporting purposes-but as part of the there came the day when they had the immeasurable economic underpinning of a rural life of limited means. support of the late 's movie about Bambi. There was nothing- wrong- with that so long- as a seem- ingly bottomless supply could meet the demand of a Saving the deer meant restrictive regulations and, sparse human population above 311, no doe hunting--this at a time when the But the relationship didn't hold. Game supplies dwin- animals were in plague numbers and starving by the ed, and society stepped in with the doctrine that owner- tens of thousands every winter. Good biology dictated ip of resident wlldlife rests in the State and the State sweeping reduction of the herds if remnants of the range d responsibility for managing game in accordance with were to be saved, but more often than not politics ruled e supply. In Wisconsin, attempts at management began and a bad problen~grew worse. n the late 1800's. It was social and political in nature, In groping for a solution--one that would compromise aimed at parceling out remaining populations, not at between deer aplenty and winter starvatioil-a number development of herds. It's easy enough to imagine the of States have tried winter feeding. That sirnply adds political pressures that faced the early enforcement staff politics to politics to produce a sum of less than zero. Wisconsin tried it as far back as 1935, but it was not Wisconsin's Legislature tried to hush complai~ until the winter of 1912-1943 that they went all out. farmers by appropriating $40,000 annually to pay ( It was one of those "shakedown" winters, long and damage complaints. In one biennium, they shelled cold. The snow was deep, browse was in short supply, close to $60,000 in payment of 613 deer-damage cla and deer were everywhere. When the snow melted, the If such schemes were merely useless, it would be tr carcasses were everywhere, too. No one knows how enough, but they encourage dishonesty among the many starved, but estimates of the total loss ranged from zenry as well. I am not singling out Wisconsin farrr 50,000 to 200,000. One thing sure, more deer died that mind you. Any County Commissioner anywhere can winter than had been taken in any previous season by you that in the damage claims filed before his Coc the hunters. Board, the wild dogs always seem to kill only the 1 The next meeting of the legislature earmarked 50 cents ribbon livestock. fronl each deer liceilse for use in artificial feeding and Politics in Maryland are probably no worse than t the purchase of deeryards. In the 1950-1952 biennium, where-except when it comes to managing the deerh Wisconsin spent $154,000 from this fund to feed starving Then the professional's hands are tied. In some deer. And over an 11-year perioti 11913-1953) they put counties of the State a burgeoning herd has all but a total of nearly $5CO,000 in the grand experiment. some orchardists out of business. As you might ex1 a few operators have acted on their own by shoo While substantial, such an outlay of sportsmen's funds deer on sight, or hiring it done, in season and out. would not be objectionable if favorable results could be orchardist, without apology, stated he counted in ex claimed. Rut decr feeding programs have been tried of 75 dead deer in his orchard 1 year. Since he around the country and the unanimous feeling of trained shooting them with a .22, the odds are the total was wildlifers is that, in general, feeding aggravates the above that figure. problem. The complexitits don't cnd with the resort trade Other Reporting to the Sixteenth Annual Conferencr intcirests have other reasons for managing deer in a Southeastern Association of Game and Fish Commiss different fashion There's the forester whose stands of ers, Maryland biologists Flyger and Thoerig, said t serdlings are browsed to the ground by the ravenous "Feelings run high between orchardists and hun hordes This can add to a substantial loss, in some areas, ers in this region. The orchardist wants deer numl decr are taking off I out of 5 acres of forest rcproductlon ers reduced and together with the Game and Inlar Then there's th~.orchardist whose fruit trees are chewed Fish Commission have tried to establish antlerle baclr to misshapcned snags And farmers everywhere seasons. The more vociferous hunters in this art bemoan the loss of soybeans, corn, mclons, tomatoes, have been able to prevent all but one small ineffe peppers, alfalfa, winter pasture, and other crops too tual antlerless season. On one occasion the Gan numerous to mention Commission, wishing to control the killing of de~ It matters llttle if as often happens that the orchard and lteep it out of the hands of the public, assignc or patch iarrn was grubbed from a cutover forest and so wardens to shoot decr in the orchards. This did n( had a full blown depredations potentla1 built in The please the hunters and within a week a large o fact is that for these people there are indeed too many storage bullding ln one of the orchards mysterious deer and, hunter liitercst\ aside, the anrmal~are little caught fire and burned to the ground, just as one I morc than vcrinin to bc clim~ilateden masse the hunters earller had predicted would happen In the West the dileinma takes a new form Here the They went on to say: cattlemc1-1 and woolgrowers want deer off--more live- "The deer situation in western Maryland is nl stock on Sportsmen, as expected, argue for reduced only unpleasant and wasteful but needless. Effe grazing pressure to put more forage in the bellies of big tive management of the herd is obstructed by a sma gamc. I hasten to emphasize that 1 am not painting either but influential portion of local hunters. The Gal1 as the bad guy. A cattlenian or a woolgrower can also be and Inland Fish Comniission is caught in the midd a sportsman. I suppose this illustrates the political im- of this situation with its hands tied by legislatic plications of inoderil deer oratory. and public opinion. On the one hand farmers d I11 niosl areas the conflict between drer and agriculture niand action threatening to take matters into the grows worse as the adaptable white-tail learns to relish own hands. If the Con~missionpermits such actio the products of the farm. Would-be solutions have come open lawlessness is encouraged with loss of respe not from bicilogists, but from legislative halls. The re- for hunting laws However, if a farmer is prosecute sults, as you kilow, have bce~jless than satislactory for shoot~ngdeer 111 defense of hls property the cou One of the early palliatives was Pennsylvania's Decr- is likely to favor the farmer, thereby settlng a prec Proof Fence Law of 1923. Through this expedient, dis- dent for other landowners and farmers to follow tressed farmers were furnished 8-foot wire fencing to On the other hand, hunters want more deerhuntir hold back the hungry herds, providing they would meet but refuse to permit kin antlerless season believii h'tlf the cost of con\truction Sm,ill landowners moat that they call build up their herd." 111 need of help couldn't diiord lt -and fortunately so While I have dclved into the past and into probl for the gaine dep,irtmciit I3ccausc fencing out dcei- of faraway States In the course of exempllfylng poli "iierc" s~ml,ly put5 mr,re ol them "there," anti when considerations that have entered into deer manager followed to its logical end the scheme would bai~kru~~tit should not be inferred that thcse problems have s any gallie f~md. been solved. To the contrary, these and more are much with us and some tend to be aggravated by still as the camp cook. The solution-no agent should be per- another relatively new development on the scene. I refer mitted to work in his home parish. to the trend away from free hunting as we have known A few elected public officials fail to take action against it and towards commercialized gunning and private pre- deer poachers or go very light on them with the expecta- serves. tion that the offenders will round up several votes during Over much of our northern country the number 9 wire the next election. Some feel sorry for the violator be- has come to identify the private deer club. In Michigan's cause he may be poor and have a large family. But in better deer range one can now drive for miles and see this day and age no one need be dependent on game for on both sides of the road the single strand of wire with a meat supply. It should be pointed out that most sher- signs hanging at measured intervals that caution the iffs, district attorneys, and judges are more enlightened unattached hunter to stay out. It started in the Lake today than ever before and that they assume their right- States area, but has since spread to many parts of the ful responsibility by taking proper action and imposing country, including the South. Today deer clubs are stiff fines. To most of them, game laws are no longer prevalent throughout the range of the white-tail. And a joke. in the course of reserving hunting rights for a relative In Louisiana, the parish governing body in 37 parishes few, the membership has opened up a new area of politi- supposedly has the authority to veto doe seasons in their cal maneuvering. parish. There have been many instances in which these I can speak firsthand of situations that have appeared police juries have not permitted the Wild Life Cornmis- in Louisiana. sion to have an any sex deer season when it was badly needed. These same governing bodies have enacted re- In Louisiana a few clubs prevail upon prominent strictive camping ordinances and trespass laws designed political figures to take meinbership. This gives them especially to harass out-of-parish deer hunters. a certain amount of prestige and a lot of political lever- age. During years when permits are issued for the taking Some Wild Life Commission members are politically of extra deer, those clubs having the most influential motivated and insist on a politically oriented program. political leaders exert the most pressure and receive They still believe in the "spoils system" in which em- the most permits. However, many progressive clubs ployees are rewarded for their political activity. These leased land having a low deer population, protected it political employees have no knowledge of deer problems and have no interest in them. at their own expense and harvested deer as recommended Some are complete "dead- heads" and some spend most of their time in political by State game biologists. Because of good management, they harvest many more deer annually per unit of area activities. Under these conditions, management of our deer reaches its lowest point. This form of wildlife ad- than is taken on surrounding lands that are open to ministration is not only the least productive but it is also public hunting. Since most club members invite friends the most costly. to hunt, they spread the kill among a large number of people. We know well enough what happens to biological management when dollar values appear for any entity of A few tax assessors have been influential in deer man- the outdoors. I have already alluded to the power play agement by threatening to raise taxes when forest land- of summer resort operators who, in part, are capitalizing owners demanded a reduction in deer numbers because on the assured prospect of their clientele seeing deer. no forest reproduction could be obtained. In other areas (particularly the West) similar commer- Wherever deer come in contact with soybeans, there cial interests are catering to free-spending, nonresident is no escape from damage. Tensas Parish, Louisiana, has hunters. These interests seek more liberal seasons and many acres of beans and a high deer population; there- bags and, above all, a "fair and reasonable" nonresident fore, farmers suffer crop depredations. For the past 2 license fee that will encourage a good influx of outside years farmers have killed many deer in bean fields dur- hunters. Again, it pits dollar interests with local hunters, ing the crop growing season. They were left to rot or and management decisions often involve a little biology given to farm employees. Wildlife agents filed many and a lot of politics. charges against these farmers, but they were not brought And there's more. Few people outside the South can to trial. The police jury, sheriff, and the district attorney appreciate fully the intensity of feelings engendered by maintained a list of people who would not be prosecuted. devotees of the fox chase. Some fox hunters hold deer The Wild Life and Fisheries Commission contended that in genuine disdain. To a man with a , Heaven is the deerherd should be reduced by licensed hunters a township crawling with red foxes but free of grays during the regular season to a level that damage could and white-tails and in which dogs can be run every night be tolerated by the farmer. of the year. And they make themselves heard, but not In Louisiana, legislators often exert heavy pressure without a great clamor from deer hunters who seek on the Wild Life Commission to employ a particular laws permitting the shooting of "free-running hounds." person as a local agent. He then becomes "my" agent If that's not enough, you can add archers versus gun and may be requested to give special protection to fav- hunters, riflemen versus the advocates of slugs and buck- ored lands during the nonhunting period and to look shot, doe hunters versus buck-only supporters, quail hunt- "the other way" during the deerhunting season. EIe often ers who want no deer gunners out during the fall and condones violations by local hunters and harasses outside winter. And I will add the far-out instance of a Michigan hunters. In some instances he may be used as a guide, Commissioner (there are probably others from other driving a State vehicle to take hunters into the woods States) who wanted an earlier deer season so it would and to carry deer out. He may even be requested to serve not conflict with his Florida vacation. All of them are being heard-and far too many being heeded-in caucus the problem--if indeed it has one-will have to fol rooms well removed from the laboratory and checking on sound biology. station. We have the facts but not the following. No ma In retrospect, we have not always managed our deer- how valid the basis for biological management, i herds along sound biological lines. This most popular useless in the face of public resistance. And, so, a and abundant of our big game animals has a way of commonly the case, our job now is one of communica touching the lives of many people of diverse interests. and education. Not to be entirely pessimistic, we h witnessed good progress in some States, but it's not lil. A majority of our landowners in the range of the to stay that way unless we keep on with our educatic white-tail are aggrieved by its presence. Foresters, graz- efforts. As has been remarked, in conservation educa ers? truck crop producers, diversified farmers, orchard- we are not addressing ourselves to an audience bu ists, and others all bemoan their presence as they see a passing parade. Each new generation brings its ( their livclih?od penalized by this omnivorous beast. doubters and self-styled experts and so we must p The conflict occurs to some extent even when herds for public understanding and support for a long tim~ are maintained within the carrying capacity of the na- come. tural range. And the strife increases when populations As of now, the name of the game is still politic: are allowed to expand beyond the capability of the most areas, and in terms of maximum sustained y natural habitat to support them. On the other side of it's not the best way. But by other measurements the ledger are still other interests who see economic gain, may approach "the greatest good for the greatest num and sporting satisfaction in seeing the herds at saturation for the longest period of time." And as profession level. It often adds up to a wasted resource through we live by that credo too. In the meantime, the consul starvation and impaired reproduction in the herds. audience, which adds to 6 million, gains 40 million re, ation days from the sport while decorating 2 mil Too often solutions have been sought in trying to fence the animals out, payments for damages, artificial feed- fenders with this greatest of game, or worst of verr ing, trapping and removal and other expedients that depending on how you view it. wasted funds and accomplished no good. A solution to And that's not bad.