Bulletin of the New York State Vol. X, No. 2 College of Forestry at Syracuse University October, 1937 THE EFFECT OF DEER BROWSING ON CERTAIN WESTERN ADIRONDACK FOREST TYPES By John Pearce Roosevelt Wildlife Bulletin VOLUME 7 NUMBER 1 Published by the Roosevelt Wildlife Forest Experiment Station at the New York State College of Forestry, Syracuse, N. Y. SAMUEL N. SPRING. Dtan CONTENTS OF RECENT ROOSEVELT WILDLIFE BULLETINS AND ANNALS BULLETINS Roosevelt Wildlife Bulletin, Vol. 4, No. i. October, 1926. I. The Relation of Birds to Woodlots in New York State Waldo L. McAtee 2. Current Station Notes Charles C. Adams (Out of print) Roosevelt Wildlife Bulletin, Vol. 4, No. 2. June, 1927. 1. The Predatory and Fur-bearing Animals of the Yellowstone National Park Milton P. Skinner 2. Current Station Notes Charles C. Adams Roosevelt Wildlife Bulletin, Vol. 4, No. 3. July, 1927. 1. A Trout Survey of the Allegany State Park in 1922 William C. Kendall and Wilford A. Dence 2. A Preliminary Survey of the Fish Life of Allegany State Park in 1921 . . Thomas L. Hankinson 3. Current Station Notes Charles C. Adams Roosevelt Wildlife Bulletin, Vol. 4, No. 4. July, 1927. I. The Beaver in the Adirondacks : Its Economics and Natural History.. Charles E. Johnson Roosevelt Wildlife Bulletin, Vol. 5, No. i. March, 1928. 1. A Preliminary Wild Life and Forest Survey of Southwestern Catta- raugus Co., N. Y Victor H. Cahalane 2. A Preliminary Report on the Trout Streams of Southwestern Catta- raugus Co., N. Y Wilford A. Dence (Out of print) Roosevelt Wildlife Bulletin, Vol. 5, No. 2. February, 1929. 1. The Fishes of the Cranberry Lake Region W. C. Kendall and W. A. Dence 2. The Story of King's Pond F. A. Lucas 3. Its Fish Cultural Significance W. C. Kendall Roosevelt Wildlife Bulletin, Vol. 5, No. 3. September, 1929. 1. The Summer Birds of the Northern Adirondack Mountains Aretas A. Saunders 2. The Summer Birds of the Adirondacks in Franklin County, N. Y Theodore Roosevelt, Jr., and H. D. Minot (Reprinted. Original date of publication, 1877) Roosevelt Wildlife Bulletin, Vol. 5, No. 4. August, 1930. 1. The Biology of the Voles of New York Robert T. Hatt 2. The Relation of Mammals to the Harvard Forest Robert T. Hatt Roosevelt Wildlife Bulletin, Vol. 6, No. i. March, 1931. I. A Biological Reconnaissance of the Peterboro Swamp and the Labrador Pond Areas Charles J. Spiker Roosevelt Wildlife Bulletin, Vol. 6, No. 2. October, 1933. 1. The White-tailed Deer of the Adirondacks. Part I. Preliminary Survey of the White-tailed Deer of the Adiron- dacks M. T. Townsend and M. W. Smith Part 2. Ecology of the White-tailed Deer in Summer with Special Reference to the Adirondacks M. T. Townsend and M. W. Smith 2. Some Late Winter and Early Spring Observations on the White-tailed Deer of the Adirondacks Chas. J. Spiker Bulletin of the New York State College of Forestry at Syracuse University Vol. X. No. 2 October, 1937 THE EFFECT OF DEER BROWSING ON CERTAIN WESTERN ADIRONDACK FOREST TYPES By John Pearce Roosevelt Wildlife Bulletin VOLUME 7 NUMBER 1 Entered as second-class matter October 18, 1927, at the Post Office at Syracuse, N. Y., under the Act of August 24, 1912 ANNOUNCEMENT The serial |)ul)licati(>iis of the Koosevelt W ildlife ln>rest Exjjeri- iiK-iit Station consist of the followinj^ : 1. Roosevelt Wildlife Bulletin. 2. Roosevelt Wildlife Annals. The Bulletin is intended to include ]ia])ers of general and pojailar interest on the various ])hases of forest wildlife, and the Annals those of a more technical nature or having a less widesjiread interest. The editions f)f these puhlications are limited and do not permit of general free distribution. Exchanges are invited. Sale prices for the Station publications are based on the actual cost of printing and distribution in accordance with Chapter 220 of the Eaws of 1933. Price lists will be furnished on request. All communications con- cerning publications should be addressed to The Director axd Editor, Roosevelt Wildlife Forest Experiment Station, Syracuse. New York. Copyright. 1937. by Roosevelt Wildlife Forest Experiment Statio.n [2] Errata The follov/in^^ errors were made by the "printer in :ria,''cin^ the fin-.l chan-'^es in the te'-'t, Pa^e 39» F^^t perio'" after "Forest'' in lor-end r'or Fi^^, P-C» Pa-^e U9» "Sc'iool" should be capitalized in le-^end for Fi'^, Inside "back covir, Pat comria after "Jul '" in first line. Digitized by the Internet Archive in 2015 https://archive.org/details/rooseveltwildlif07unse TRUSTEES OF THE NEW YORK STATE COLLEGE OF FORESTRY Ex Ofkh io Dr. William P. Graham, Chancellor Syracuse University Dr. Frank P. Graves, Commissioner of Education Albany, N. Y. Hon. LiTHCiow OsRORNE, Conservation Commissioner... Albany, N. Y. Appointed in' the Governor Hon. M. ^\'ILLIAM Bkav, Lieutenant-Governor Albany, N. Y. Hon. Francis D. McCurn Syracuse, N. Y. Hon. Alfred E. Smith New York City Hon. William H. Kelley Syracuse, N. Y. Mr. Willis H. Michell Syracuse, N. Y. Mr. Charles A. Upsox Lockport, N. Y. Hon. J. Henry Walters New York City Mr. George W. Sisson, Jr Potsdam, N. Y. Mr. Reginald T. Titl s New York City Officers of the Board Hon. Alfred E. Smith President Hon. William H. Kelley Vice-President HONORARY ADVISORY COUNCIL OF THE ROOSEVELT WILDLIFE FOREST EXPERIMENT STATION Hon. Theodore Roosevelt New York City Mr. Kermit Roosevelt New York City Dr. George Bird Grinnell New York City Hon. GiFFORD Pinchot Harrisburg, Pa. Mr. Chauncey J. Hamlin Buffalo, N. Y. Dr. George Shiras, 3rd Washington, D. C. Dr. Frank M. Chapman New York City Dean Henry S. Graves New Haven, Conn. [3] STAFF OF THE ROOSEVELT WILDLIFE FOREST EXPERIMENT STATION Samuel N. Spring, M.F Dean of the College Ralph T. King, M.A Director WiLFORD A. Dence, B.S Ichthyologist and Ass't Director William L. Webb, B.S Junior Forest Zoologist H. Ruth Merrill Secretary Temporary Appointments * Myron T. Townsend, Ph.D Field Naturalist A. Sidney Hyde, M.A Field Naturalist MuRVEL R. Garner, Ph.D , Field Naturalist Charle.s J. Spiker, M..\ Field Naturalist Collaborator John Pearce, M.S Game Naturalist * Including only those who have made field investigations and whose reports are now in preparation. [4] Fig. 2. Composted horse manure heap upon which deer had chewed exteiiMVci} , Huntington Forest. Photograpli by C. E. John.-^on. THE EFFECT OF DEER BROWSING ON CERTAIN WESTERN ADIRONDACK FOREST TYPES By JOHN PEARCE, Collaborator Rooscz'clt Wildlife Forest Experiment Station, Syraeitse, A' civ York CONTENTS PAGE Introduction 8 Habits of White-tailed Deer Tending to Influence Forest Growth . 1 2 Seasonal Changes in Diet and Range 12 Browsing Procedure of Deer 15 Uncertain Tastes Developed by Deer 15 Sociability of Deer 16 A Method for Measuring the Influence of Deer Browsing on the Composition of Young Stands 16 Sampling 16 Tallying 17 Explanation of Terms Adopted for Sample Plot Tallying. . . 19 Recording Terminal Browsing 20 Distinguishing Between Deer and Rabbit Browsing 21 The Influence of Deer Browsing in the Sucker Brook Yard 22 Description and History of the Yard 22 Description of Forest Cover Types in the Yard 22 Analyses of Sample Plot Data 24 Swamp Type 24 Spruce Flat Type 35 The Influence of Deer Browsing in Other Forest Types of the Cranberry Lake Region 41 Browsing in Bums 41 Browsing in Old Growth Types 42 Browse Chosen by Deer in the Cranberry Lake Region 43 Factors Influencing Browse Choice 43 Discussion of Browsed Species 44 Deer Preference for Shrubs 48 Reactions to Browsing Damage Noted Among Species on the Sample Plots 50 Factors Influencing Effect of Browsing Damage 50 Witch Hobble as a Key Species 52 Summary and Conclusions S3 Scientific Names of the Trees and Shrubs Mentioned in the Text which were Encountered in the Field 54 Bibliography 55 [7] 8 Roosevelt Wildlife Bulletin INTRODUCTION This study was inaugurated for the i>urp(jse of obtaining definite information relative to the quantitative efifect of animal browsing on the future character f)f the timber stand. The silvicultural effect of browsing must be clearly understood before the inhabita- tion of forests by herbivorous animals can be made compatible with most other forest uses. Studies to this end are indispensable to in- telligent forest-use planning. The writer has attempted to present, ])rimarily, the result of pre- liminary browsing investigations made in an area embracing two common western Adirondack forest types, i. e., the swamp and the spruce flat. This area had been previously cut over and subjected to deer yarding. Consideration has also been given to "preferences" shown by deer for the woody species encountered, the a])parent ability of these species to withstand the adverse effect of browsing, and the initial attempt to name an indicator or key species, which reflects the general intensity of deer browsing. Since this paper is but the outgrowth of a thesis submitted by the author as a require- ment for tiie Master of Forestry degree in silviculture at the New York State College of Forestry, it is not contended that final informa- tion is offered on any of the phases mentioned. The revised thesis material is herewith presented as a basis for further studies of a similar nature that are needed by foresters, biologists and wildlife technicians. The field data were obtained by methods which will have some degree of general usefulness. While this technique is treated at length, it will serve chiefly as a starting point for other investigators who seek to measure the influence of browsing on stand composi- tion. Certain drawbacks to the completeness of this study are acknowl- edged. Principally, the fact that the number of deer responsible for the browsing tabulated in the type studies is not known. Ignor- ance of this figure, however, is of less consequence in silvicultural studies on stand composition than in game management work. Fu- ture refinements of the procedure will undoubtedly call for studies on stands of known density influenced bv known numbers of deer. An area which lias l)een under close observation over a period of years also is desirable in order that the fullest ])ossible history may be presented. The white-tailed deer under consideration, Odocoilcns z'irginiamts borealis Miller, is the sub-species native to the Adirondacks. Its gen- eral browsing habits differ but sliglitly from those of other sub- 9 10 Fig. 5. Illustrating how inconspicuous hardwood browsing may be, even though fairly severe. Sucker Brook yard. September, 1934. Fig. 6. White pine with terminal and laterals repeatedly browsed. Sucker Brook yard. July, 1934. Deer Browsing in the Adirondack's II species in the East, although sharp differences may be found in the food chosen by tlie same sub-species inhabiting other sections of the countr}'. This fact must be recognized at the outset in order to avoid confusion from seemingly contradictory statements in the litera- ture. "Browsing" as used in this paper means "the feeding on browse," which Dixon ( 1934) defines as "shoots, twigs, leaves and fruits of woody plants." Reconnaissance work was carried out in June, 1934, to locate a suitable study area in the Cranberry Lake territory. During this period an examination was made of the forest about the State Ranger School, Cranberry Lake proper, Peavine Swam]), and a considerable area between Big Deer and Clear ponds (Map i;. These areas contained several types, including virgin, old-cut, new- cut, and burned-over forest, in a wide assortment of conditions. All of these supported deer. However, the effect of deer browsing was most sharply defined in the well-drained swamp type and in the lower sections of spruce flat t\ pe, which were heavily cut al)out ten to twent\- }-ears ago. Consequently, this study is limited mainly to these two types of forest growth. The locality finally selected lies along Sucker Brook about two and one-half miles east of Cranberry- Lake from Barber Point and is hereafter referred to as the Sucker Brook Deer Yard. Deer used this area as a wintering ground for several years. Detailed examination of the area was made in September, 1934, b}- the author, with the aid of Mr. Paul Zimmer as tally man. The data were gathered by recording the tree and shrub tally on milacre (one-thousandth acre) plots taken at regular inter- vals throughout the main portion of the yard. The common and scientific names of the trees mentioned in this report correspond with those of Sudworth (1927) ; of the shrubs, Rehder (1927). Only the common names appear in the text proper, but the corresponding scientific names appear at the end of the report on page 54. I wish to express my gratitude to the Xew York State College of Forestr}' for enabling me, by means of a graduate fellowship, to pursue the field research and subsequent analysis of data connected with this study. For valued assistance, courtesies or constructive criticism I am grateful to the following: Miss H. Ruth Merrill. Secretary of the Roosevelt \\'ildlife Forest Experiment Station ; Prof. James F. Dubuar, Director of the State Ranger School at Wanakena; Messrs. T. R. Phillips and O. W. Oja of the Huntington Forest at Newcomb and Mr. Wilford A. Dence of the faculty. I am particularl}- indebted to Prof. E. F. McCarthy, also of the faculty, for his many kindnesses and helpful counsel. 12 Roosevelt Wildlife Bulletin HABITS OF WHITE-TAILED DEER TENDING TO INFLUENCE FOREST GROWTH Certain habits attributed to the normal life history of the deer have a direct bearing on the findings. ^luch has been written on the subject by many able naturalists and it would be beside the point to present a thorough review of their work at this time. However, a brief summary of several pertinent browsing habits will undoubtedly prove helpful. Seasonal Changes in Diet and Range. — The autumnal change in the diet of deer from herbaceous plants to browse is induced chiefly by cold weather withering the succulent material. Coincident with snowfall comes a reduction of the daily range, which reaches greatest restriction during yarding periods enforced by deep snows. However, the actual diminution of the range may not l)e as great as is sometimes thought. The normal range of the white-tailed deer throughout the year is less extensive than that of most large mammals. Seton (1909) places it at less than any other species of native big game. Leopold (1933) sets it at about five miles. During extended yarding periods young trees may be subjected to considerable damage, particularly when the deer are confined for two or three months. Obviously the damage is more uniformly scattered during open winters when deer yard only for short periods. The term "yard" has not been standardized for all parts of the countrv. The eastern usage of the word generally means the imme- diate vicinitv to which deer are confined in heavy snows — an area, or occasionally several closely connected areas, used by a band of deer. Townsend and Smith (1933) define a yard as being, typically, a swampy area and its adjacent slopes. Deer herds indicate a strong desire to remain on a definite range ; they oftentimes continue to inhabit heavily-used, over-browsed areas rather than travel to easily accessible areas for better feed. Several individuals have noted this ^^eculiar ])hase of deer nature : Pearson (1925), the Pennsylvania Board of Game Commissioners (1930), Clepper (1931) and Ruhl (1932) have all remarked on it. The hidden factors which cause such seemingly illogical habits are significant, but beyond the scope of this investigation. This has a direct bearing on the use of deer yards, however, inasmuch as yards are frequentlv used season after season in spite of the greatly reduced food suppl}-. The only over-browsed areas encountered were former yards. 13 - .1 Fig. 7. Snowshoe rabbit skull. Note efficient upper incisors. Fig. 8. Skull of Adirondack buck. Note lack of upper incisors. 14 A B C Fig. 9. Twigs bitten ofi' by deer and rabbits. Those bitten off by rabbits are on the right in A and B. Compare these with the knife cutting on the right in C. Fig. 10. Typical rabbit damaged red spruce. Note that the top, being out of ordinary reach, was not damaged while the lateral branches were greatly affected. Osvvegatchie River, near Ranger School. June, 1934. Deer Broivsing in the . Idiroiidaeks 15 Browsing Procedure of Deer. — A very pronounced characteristic of deer is their unmethodical manner of feeding upon a plant. At the first browsing they seldom strip, completely, shrubs or saplings over, one or two feet in height. Their method of feeding has a certain daintiness which led Caton (1880) to comment on the "delicacy" of their feeding habits. Tourney (1928) and Townsend and Smith (1933) used the term "selective" to describe deer browsing. These remarks have been borne out b\- my own observations on deer feed- ing in several localities. Frequently they eat a few twigs or leaves at one point, stop and glance around, jiroceed rather nervously for a few steps, take a few more bites and move on — even when in no way alarmed. Consequently, plants are less affected and better able to continue growth than would be the case if hea\nly browsed. The accumulative effect of the light feeding damage over a wide area tends to prolong the life of the plants. As a result a deer herd may be supplied with food for a longer period of years. Uncertain Tastes Developed by Deer. — The uncertain choice of food by deer is sometimes a factor to be considered in the forest. Schenck (1909) says that the greatest damage is done to the rare species of plants which seem to arouse the curiosity of deer. Shepard (1934) mentions cases of European deer peeHng spruce bark — the reason being attributed to faulty diet. As an extreme example of what may be attractive to deer. Job (191 5) cites an established instance where thirty-eight deer died on a preserve from eating poisoned mice. Rutledge (1931) sums up the situation, paradoxi- cally, by saying that it is "much harder to discover what a deer will not eat, than what it wnll." The inconsistency with which deer feed on any one species of plant is even more surprising than their taste for strange food. Investigations by Clepper (1931) in Pennsylvania indicate that many species of browse plants are usuall\' more attractive to deer if grown on recently burned lands than browse of the same species grown on unburned lands. Tree planting experience in the same state lend weight to the idea that trees planted in fertilized nurseries are more attractive to deer than those reproduced naturally on unfertilized soil. This seemingly is due to the higher mineral content in plants grown on newly burned or fertilized areas. Coville (1929) reports a case where deer apparently showed a decided preference for western yellow pine, Piniis ponderosa, grown from Black Hills seed to that grown from seed from some other sections. Chemical variation in l6 Roosevelt Wildlife Bulletin the different stocks was advanced as a tlieory for the clioice. Mitchell and Hosley ( 1936) cite an example of a definite increase in hrovvsing due to increased nitrogen content hrought on by fertilizing. Clepper (1936) mentions investigations made by Prof. G. S. Perry, which indicate that increased mineral salts are found in forest soils follow- ing silvicultural thinnings. This may account for the well known deer preference for sprout growth. Sociability of Deer. — Deer congregate in loosely organized groujjs after the rutting season. Each group, averaging from five to twenty individuals in the Adirondacks, usually remains together until spring. Feeding proceeds with the whole group much the same as with solitary individuals. An average area is covered several times, and a network of trails is established before overbrowsed conditions become general. The exi.stence of these trails in the yards become increasingly useful as the snow gets deeper and the deer grow weaker. A METHOD FOR MEASURING THE INFLUENCE OF DEER BROWSING ON THE COMPOSITION OF YOUNG STANDS In considering methods of gauging browsing influence on forest growth one should bear in mind the wide diversity of natural stands. The minute dilYerences of two areas in the same forest type only serve to emphasize the greater variation ])etween sejmrate types. It is not likely, therefore, that one method can be used in ancrther section without deviation. Tlie method used in this study is described in detail because there is a scarcity of such information in forestry literature. Sampling. — A statistical summary, comprising the (|uantity. status and s])ecies of ])lants of the area, is the first requisite of a quantitative browsing study. Obviously a 100 j^ercent tally of all vegetation on even a small deer yard would be a forniidal)le task and not in accord with present dav knowledge of sampling procedure. Therefore, in order to arrive at a sound representation of the wofKly growth on the area it was considered l)est to use sample plots. In order to minimize errors in personal variation these plots were of uniform size and spacing. The "(luadrat-at-interval" method of Clements (1905), which is frequently called the "line-plot system," was chosen as the most efficient means to this end. For convenience, thousandth Deer Bro7Vsiiu/ in the . Idirointacks 17 acre Idiots were adopted. These were taken along cruise lines run at right angles from an estal)lished haseline (see sketch p. 23). This system included samples within the spruce flat type as well as the swamp type. The cruise lines were one-half a Gunter's chain (33 feet) apart. The plots were located 33 feet ai)art and forward of the half-chain point along these lines. The distance hetween the plots was established by pacing, while two straight poles, 79.2 inches in length, were used for laying off the sides of the plots. A tape measure could be used to check the diagonal distance of 112.005 inches. Tallying. — While it must 1)0 acknowledged that each i)lant species ])robably has some ecological influence on every other associated species, nevertheless in dealing with a study of forest types it is ob- vious that the major influences affecting tree species are the main factors to consider. The influence to be considered in this particular study concerns deer browsing, therefore the data is limited to the species of woody plants that affect the composition of the stand. In certain areas herbaceous plants may be sufficiently abundant to have a decided effect on woody plants, but that is not the case in the Sucker Brook yard. A specially designed form was used for tallying. The information contained thereon is outlined below : A. Hardwood tree species 1. Browsed a. Dead ( I ) Height class and number times browsed back (leaders) while alive b. Alive (i) Height class and number times browsed back (leaders) 2. Unbrowsed a. Dead (i) Height class h. Alive ( I ) Height class 3. Escaped a. Dead b. Alive i8 Roosevelt Wildlife Bulletin B. Softwood tree species 1. Browsed a. Dead (i) Height class and iiuiiiIkt times browsed back (leaders ) while alive b. Alive (i) Height class and nunilx-r times browsed back (leaders) 2. Unbrowsed a. Dead ( I ) Height class b. Alive ( I ) Height class 3. Escaped a. Dead b. Alive C. Shrub species 1. Alive a. Degree of abundance ( 1 ) Sparse (2) Common (3) Abundant b. Degree of browsing (1) Light (2) Moderate (3) Heavy c. Height class 2. Dead a. Al)undaiice as above b. Browsing sustained while alive (1) Light ( 2 ) Moderate (3) Heavy c. Height class 3. Escaped a. Alive b. Dead Specimens in the escaped class are not listed as browsed or un- Iirowsed. This classification was omitted because of the general impossibility of stating, after the lapse of a few growing seasons, the extent of browsing jirevious to escaping. Large trees were not tallied because the majority of them were culls which had only slight influence on the young growth in the two types. Deer Bro'a'siiui in the . Idiroinhieks 19 Explanation of Terms Adopted for Sample Plot Tallying. — - IleiyJit Classes ixcjiks in Class Heicht 1- foot I — 12 2- foot 13 — 24 3- foot 25—36 4- foot 37 — 48 It is practically impossible to state at what height a tree will 1)6 absoltitely safe from tip browsing, because many factors must be considered. The possibility of deer "riding down" trees by strad- ding them to get at the tops, the presence of large rocks, heavy snow weighing down the crowns, and packed and drifted snow — all permit the browsing of an occasional escaped tree at an abnormal height. Yarding sites in the Adirondacks are protected, ordinarily, from the wind force necessary to pack snow firmly enough to sup- port deer. Heavily-crusted snow usually occurs for short periods only. The escaping point was established by computing the lowest average height at which trees in the district were free of tip brows- ing. This was accomplished by determining the highest point at which browsing had occiu'red on escaped trees, foimd along several random lines. Care was taken to do this on areas which obviously had been browsed well during the time required for the trees to escape. The standard height as adopted (6 feet) is about the same as that arrived at by Hosley (1931), Bailey (1933) and others. Mr. .\. V. S. Pulling, Assistant Superintendent of the Upper Mississippi Wildlife Refuge, however, in a personal letter states that he has seen trees "much browsed" at seven feet, and some as high as nine feet where hard-packed snow occurred in Wisconsin. Degree of Browsing. The approximate intensity of browsing sustained on i)lants, including trees and shrtibs. was classified arl)i- trarily as ( I ) light, (2) moderate and (3) heavy. The following is a description of the three classes : Light Browsing — Damage so slight that it may be over- looked. Little efifect is apparent on the plants. Moderate Browsing — The plant visibily, but not seriously affected. A somewhat trimmed efifect is apparent. Heavy Browsing — Plant critically affected; decided inter- ference with height growth. An excessively trimmed appearance frequently noticed. Lnches in Class Height 5- foot 49 — 60 6- foot 61 — 72 Escaped 73 — 20 Roosevelt Wildlife Bulletin Degree of Abundance. An approximation is necessary where exact counts cannot l)c applied readily to the stems on a plot; for exani])le, cluni])cd sliruhs whicli fork from low stems. It so hap- pened that a loose classification fitted the refjuirements of this study — three dej^rees of ahundance were adopted as standards for the work. These arc descrihed as : Spak.se — One to six sinj^le stems, or a clump of about six stems per milacre. Common — .Seven to twelve single stems, or not more than three clumps each of about six stems per milacre. Abundant — Over twelve single stems, or more than three small clumps per milacre. Those who would have a more definite count of the shrubs may criticize this system, liut it is not sinij)le to devise one, especially after moderate or heavy browsing has caused i)rofuse sprouting of a species which suckers or layers easily (.See Fig. 23). Recording Terminal Browsing. I-lvidence of deer bnnvsing on woody stems usually remains for .several years. Due to this fact it was possible to determine definite years of yard occupancy. This was done by making growth-ring counts immediately below the browsed ends and comparing these with the subsequent growth re- sulting from the buds below the breaks. By close examination also it was possible to state at least the minimum number of times the terminal buds of unescaped trees had been browsed. Since the regular increment of additional height becomes a critical factor in young stands which must grow beyond a certain point to escape perennial danicige, the determination of shoot browsing is important. The ease and accuracy with which ;his "browsing back" can be determined depend upon the tree spe- cies browsed and the time elapsed since the feeding. The "number of times browsed" as herein used, refers to the apparent number of tunes the original leader (or the subsequent shoots assuming the lead) had been turned aside from regular growth by browsing deer. If based on visible evidence the minimum always will be the recorded number of browsings. For example, a shoot appears to have been broken off but once, although the stem is possibly more than a year old, and an older break is not visible. Again, evidence cannot be recorded unless the work is definitely attributed to deer. Rotting of browsed stubs is a common cause of uncertainty on this score. Deer BroivsiiKj in the Adirondacks 21 Maples, and other opposite-budded s])ecies, are exceptionally fav- orable specimens for collecting data on old browsing. In the winter when such species lose their tips the sections back to the first pair of buds die. In the following spring each bud of the pair develops into a branch forming a Y-shaped crotch with the browsed dead stub in the center. The age of the crotch may be determined during subsequent years b\- ring counts of a section near its base. It often happens that a series of such crotches may be traced out when each succeeding crotch is browsed, provided the stubs in the crotches remain to be identified as deer damage. Alternately budded species were somewhat less adapted to such studies, although on most stems it was possible to distinguish several browsings. When this type of shoot is broken the nearest bud generally develops into a new leader which grows at a slight angle to the stem. A zig-zag appear- ance is thus produced when such a branch has been browsed repeat- edly for several years. \\'hen the little stub disai)pears, only the angle in the twig or branch, and a slight disconformity in the bark mark the former break. This evidence is less conspicuous in later years than is the "crotching" mentioned in connection with opposite budded species. In either type of budding, the confirmed habit of the deer to feed only on the growth of the previous season, unless very hard-pressed, is a distinct help in ascertaining the dates of old damage. The browsing dates of such softwoods as white pine, tamarack and balsam can be determined in a similar manner. In these species a lateral branch frequently grows upright from the first whorl below the break to replace the lost terminal. Sometimes the most promising lateral is browsed before it is wholly upright. The effect of this check is actually equal to a browsing of the leader. Distinguishing Between Deer and Rabbit Browsing. Except in rare cases the ends of deer-browsed twigs can be dififerentiated from those clipped ofif by rabbits. Rabbits have an efficient set of upper and lower incisor teeth while deer have no upper incisors (Fig. 8) ; the former gnaw off twigs while the latter hold them against their upper jaws and break them off over the lower incisors. Therefore the ends of twigs severed by deer are nearly square across but quite rough (often with hanging shreds of bark and wood) while those severed bv rabbits are slanting but relativelv clean and smooth (Fig. 9). 22 Roosevelt Wildlife Bulletin THE INFLUENCE OF DEER BROWSING IN THE SUCKER BROOK YARD Description and History of the Yard. The sketch map opposite this page shows the principal features of the Sucker Brook yard. Roughly, this yard comprises about a square mile of irregularly out- lined swamp land hemmed in by an esker (paralleling Sucker Brook J and slope formations — the latter supporting a spruce flat forest type. These features combined with the excellent cover make an attractive yarding ground for deer. F"igure 12 illustrates the general aspect of the cut-over spruce flat type. A small remnant of swamp growth appears in the foreground ; tlie esker in the left l)ackground. The area was cut some thirt\- \ears ago for softwood (red spruce and white ])ine) .saw logs; then in 1918 and again in 1923 for pulp (spruce and balsam fir). The second pulp operation included all softwoods down to five inches in diameter — virtually a clear cutting in the swamj). However, in the spruce flat type all the merchantable hardwood was removed, in addition to the ])ulp wood, leaving a very scattered stand of advanced reproduction and culled old growth. Both types seeded in rapidly and a heavy stock- ing of young growth was the result. This growth provided excellent winter deer browse. Mr. Warren Guinup. a local resident, states that deer have never been more than moderately abundant here in the winter time, the average population being from 8 to 12 deer. They make very little use of the area in the summer — a number of old work trails and haul roads serve as runways in crossing the yard. Description of Forest Cover Types in the Yard. The Hterature dealing with .Adirondack forest types is confusing because several authors have used different names for essentially the same types. The most widely used type designations appear to be those of Pin- chot (1898) and Graves (1899). Their classification is used for this study, although we are concerned with only two of the four types originally described ; namely, swamp and spruce flat. These may be described briefly as follows : Swamp typk — Low flats with wet soil (bogs not separated from drained swamps). Red and black spruce, balsam, tama- rack, soft maple and white pine are characteristic species. Spruce fl.\t type — Level and rolling "flats'* bordering lakes, streams and swamps ; soil fresh and frequently moist. Red spruce, yellow l)irch, red maple, white pine, hemlock, occasional sugar maple and balsam are the characteristic species. 23 24 Roosevelt Wildlife Bulletin In reality, the basis of separation is toijograj^hical location. The terms "swamp" and "s])ruce flat" have been used widely because of their convenience, but they are misleading since they may be assigned to areas which do not c(jnform to the mental jiictures sug- gested by tiie names themselves. The swam]) type may be well tim- bered and without standing water, while the s])ruce tlat may Ije (juite hilly. According t(j authoritative inf. has rightly called some of these "hidden variables." Discussion of Browsed Species. — The apparent preferences for the trees (Table 3) and for the shrubs (Tables 5 and 6) on the area are based upon the percentages of the total tally browsed. The tables merely indicate the actual conditions. They are sub- ject to the above-mentioned variables in some cases. The predomi- nant species in each type were abundant enough to furnish a sig- nificant tally, but the limited data of the remaining species were supplemented by published records — especially where such informa- tion was considered important from the standpoint of managing the forest. For example, Clepper (1931) has prepared an exten- sive list in connection with the browsing preference in Pennsylvania woodlands. These areas were over-populated, however, in many cases, so that the listed preferences dift'er in some respects from those in normally populated areas, ^ilaynard, ct al (1935) gi'^'C a preference list based primarily upon confined animals and, in addi- tion, include analyses of the nutritive value of the various plants. The following discussion of the trees and shrubs of the two types studied is presented by species, or groups of species. Maples. — The red maple was the only species of sufficient abun- dance in the sample plot data to warrant definite conclusions. Judging from the small amount left unbrowsed on the plots and its heavy utilization generally in the Cranberry Lake region, red Deer Browsing in (he Adirondacks 45 maple may be considered a winter staple for deer there. Striped, mountain and sugar maples were rather thoroughly browsed, but tiiese sjiecies were jioorly represented on the plots. These were also heavily utilized on several other areas in the Adirondacks. ;\Iaynard and his associates (1935) found maples, in general, to be good deer browse, but red maple was listed as being "readily eaten." yet not "best liked." However, their nutritional experi- ments (/or. cit.) showed this species to be an excellent deer food and capable of bringing the animals through the winter in good condition. Other writers also have found that the maples were readily accepted by deer in other parts of the East. Yelloiv Birch. — Based on the plot data, yellow birch heads the preferred species list ; the amount browsed totalled 98.6 per cent for both types. It appears to be quite generally utilized by deer throughout the Cranberry Lake country. I have been told by people from various sections of the East that the same choice prevails in other places. Maynard {op. cit.) classes yellow birch as a "best liked" species and because of its high palatability and food value lists it with red maple as a food for maintaining deer in healthy condition during the winter. Beech. — As stated previously, beech apparently is less appealing to deer than the other common hardwoods of the region. The reason is not evident, but taste or the toughness of the twigs may account for its unattractiveness. Certainly, anyone who has attempted to break a beech stem realizes how much more difficult it is to break than a maple or birch twig. Deer appear to use the species during the stress of yarding conditions, for the tally (Table 3) shows that in the spruce flat type a total of 36 out of a pos- sible 38 trees were browsed. This is in marked contrast to beech neglected in old growth located away from yards. Maynard (op. cit.) found that beech had little attraction to the confined deer. Mountain Ash, Pin Cherry, Black Cherry and Aspen. — ^With the exception of the mountain ash, only a few specimens of each were found on the plots. As a matter of fact less than a hundred moun- tain ash trees were tallied for both types. This species is subject to considerable browsing in the yards and furthermore many trees are girdled by deer feeding on its bark. As a result of this dam- age, mature mountain ash are usually scarce about deer yards. Black cherry and pin cherry w-ere not especially singled out by deer in the region. However, under yarding influence they appeared 46 Roosevelt Wildlife Bulletin to be acceptable. While Cie|)])er (1931) reports black cherry heavily browsed in Pennsylvania, this is not always the case. My personal observations in northwestern sections of the State were that areas of heavy deer concentration often showed very little use of tliis species. Black cherry leaves are known to be toxic to cattle, under certain conditions. ])ut I did iKjt find any record of deer being poisoned from this source. According to Chesnut (1898) and Gress (1935) the wilted or odierwise freshly damaged leaves, when eaten, give ofif a cyanic gas that results in paralysis of the lungs. The aspens (large-toothed and treml)ling) were browsed very sparingly by deer in the region. With the exception of occasional feeding on vigorous unlignified sjjrouts, slight damage was noted. Maynard considers this species as "poorly eaten." and Pulling (in correspondence) regarda it as of little attraction. But I have seen sprouts extensively browsed on several areas in Pennsylvania for- ests. Mr. Odell Julander, of the United States Forest Service, has stated tliat the aspens are considered as the key species for the summer range of the Kaibab National Forest deer. The Kaibab Deer Investigating Committee (1924), Pearson (1925) and Mac- Donald (1934) found it well utilized, while McAtee (1936) referred to it as one of the better browse species. Balsam. — Balsam i)resents a peculiar case in regard to its food value and choice by deer. This tree appears to be a choice food : furthermore, it is eaten so commonly in yards that many observers do not hesitate to call it a winter staple. However, Brandreth (1929) states that deer will die on a straight diet of it. Spiker (1933) cites examples of deer that apparently died from starva- tion although there was balsam brf)wse in their stomachs. In May, 1927, I examined appro.ximately two dozen deer skeletons and carcasses in a dense thicket of spruce and balsam near High Falls on the Oswegatchie River. These animals were trapped by deep snow during the previous winter with only spruce and balsam avail- able for food. While the l)alsam had been heavily browsed, the available suppl}' was by no means exhausted. Apparently the spruce was not browsed at all. S])iker ( 1933) says that in one observed case deer preferred white cedar and hemlock to balsam. Maynard and his colleagues finally i)roduced conclusive proof (1935) that balsam really is a starvation ration for deer. Although the animals readily accepted the balsam the\- were not maintained in Deer Browsing in llie .Uliroiidacks 47 good health. When unsuppleniented, they lost weight rapidly and finally died. Tiiis illustrates how an apparent choice may be very confusing. XortJicni Jl'liite Cedar. — Northern white cedar undoubtedly is the choice coniferous browse of deer. But it is not common in all sections of the western Adirondacks. Bartlett and Stephenson ( 1929) list it as a staple in Michigan yards. According to Mayn- ard {op. cit.) this species is highly preferred and capable of sup- porting deer throughout the winter because of its high nutritional value. The line of dead branches on the lower part of cedars which fringe many northern lakes and ponds is a much debated phenomenon. The sharp demarcation between live and dead twigs and the uni- form height are characteristic features of this line. No less an authority than Merriam (1&S4) attributes this condition to deer browsing. Many guides and hunters still accept this theory. John- son (1927), however, olifers convincing proof that it must be due to some otlier factor. As check I made a critical comparison of browsed cedar twigs with the dead branches of cedars forming the "line" about certain lakes in the Huntington Forest and I have concluded that Johnson's theory is correct. Characteristically, deer ])rowse cedar rather completely, yet the finest twigs on dead branches of many cedars which compose this "line" are often intact (see Fig. 20), Eastern Hemlock. — Hemlock was really too scarce in the sample plots to have any significance. However, it is an important browse plant in some parts of tlie region. Ehrhart ( 1936) points out that heavy browsing occurs on it in the Allegheny National Forest, a section where hemlock greatly outnuml)ers all other conifers. May- nard (1935) credits it with fair nutritional values. Red Spruce. — The sample plot data indicates that red spruce is very unpalatable to deer. This conclusion is upheld by statements of several authors: Brandreth (1929) states "deer do not like it" and Spiker (1933) says that deer "seldom use it for food." In at least one instance spruce (unspecified) is reported as an impor- tant browse \)\ant in New England (Hosley, 1936). However in some sections exotic and western species of spruce appear to be more acceptable than red. The Pennsylvania Board of Game Com- missioners (1930) list certain of these species as being eaten more than the red; Burnham (1928) expresses a similar thought. I have 48 Roosevelt Wildlife Bulletin found Norway spruce in sections of northwestern Pennsylvania sub- jected to frequent deer damage, but generally to a lesser extent than the associated tree species. The average deer population per square mile in this region is several times that of the Adirondacks. Other Conifers.— T\\t sample plot tally indicates that in most cases white pine and larch are eaten by yarding deer. The white ])ine alone is represented h\ an appreciable tally. It escaped brows- ing in only eleven out of a total count of eighty-six trees under six feet in height. Clepper (1931) records white ])ine as being heavily browsed and larch lightly browsed. Deer Preference for Shrubs. — Some shrubs appear to be highly I)rcf erred for browse. Summaries of shrub preference for the swamp and spruce flat types in the Sucker Brook yard appear in Tables 5 and 6. A discussion on the principal shrub species fol- lows. The information is based on plot data, general observations made during the field work and supplementary references to the literature. Witch Hobble. — This is probably the most generally utilized woody plant, particularly in old growth types, in the entire region. Deer feed on it during the dormant season, but I have noticed that before the advent of deep snow, only the large terminal buds are likely to be taken. Under the more stringent conditions of late winter most of the growth of the previous season is usually eaten. Since this plant is associated with shaded conditions characteristic of old growth forest, only a few were found on the sample plots. Mountain Holly. — This species appears to have quite definite site requirements not generally found outside of the swamp type. It was widely distributed through the swamp type of the Sucker Brook yard. This plant usually sustained moderate to hea\w brows- ing (95.6 per cent) and was an important browse source for deer. Wild Raisin, Chokebcrry and Honeysuckle. — These species, although not so abundant as mountain holly, were scattered through the swamp type and were moderately to heavily browsed. (In other regions I have found all of them browsed extensively.) Pub- lished data on the use of these species as deer browse are very meagre. Blueberry. — The low bush blueberry was quite generally but only lightly or moderately browsed. Ordinarily, when a certain plant species has a high percentage of browsed stems the damage will be either heavy or moderate. Tables 5 and 6 reveal that in the Fig. 25. Very light-browsed witcii hobble. Ranger school forest. March, 1935. 50 Roosevelt Wildlife Ihdleliii case of the blueberry only about 22 per cent was heavily browsed despite the fact that damage was noted on each of the 103 milacre ]jlots inhabited by the species. Under the system used blueberries are perhaps the least satisfactory of all species to classify for I)rowsing. The indication is, therefore, that this plant is not reliable as a staple. A deep cover of snow at critical periods may account in part for tlie light browsing. Other Shrubs Encountered. — This group includes shad bush, wil- low, speckled alder, raspberry, blackljerry and sweet gale. Most of these were too scarce to have any significance in regard to local preference studies. However, ncjne was refused completely by deer. Usually shad bush, the only common .species in this grouj) was browsed lightly. (See tables 5 and 6.j Deer evidently went to con.siderable trouble to get buds of the staminate aments of several speckled alder bushes located near the Ranger School in March, 1935 ; other parts of the shrub were neglected. REACTIONS TO BROWSING DAMAGE NOTED AMONG SPECIES ON THE SAMPLE PLOTS In certain sections more definite ' information concerning browse species in second growth stands is needed. Leopold (1933) recognized the need for such knowledge in order to coordinate game man- agement with silvicultural practices. \'arious authors have described the reaction of certain species of woody plants to browsing. Mann (1932) says that some species of trees and shrubs show increased vigor when browsed. On the other hand, the Pennsylvania Board of Game Commissioners ( 1930) reports that several species, such as mountain ash and rhododendron, die when persistently browsed. Also the influence of browsing on the form of the tree has been considered. Hill ( 1917) believes there is no relationship between browsing injury and the common deformities of western yellow pine. Mougin (1931) in his paper on forest browsing by domestic animals states that such activities sometimes result in trunk dam- age that renders the trees subject to fungi attacks. Factors Influencing Effect of Browsing Damage. — The deter- mination of browsing resistance is rather involved. Accordingly certain factors wdiich complicate the study of plant reactions to browsing will be discussed before the recorded reactions of the vari- ous species are considered. To begin with, it is usually impossible Deer Browsing in the Adiroiidacks to state definitely the cause for death of a hrovvsed tree. Like- wise, we do not know wlictlier hrowsin^- impairs or actuallv increases the vigor of certain trees or whether this response is in ])roportion to the browsing sustained. .Again, a host of other elusive variahles enter into the problem. It is obvious that browsed trees are more susceptible to di.sease and insect attack than unbrowsed ones. But consideration must be given to a number of other factors such as ( i ) exposure to excessive sunlight or frost (2) the presence of too much or too little moisture and (3) the type of browsing. The following list shows a few species which have persisted for several \ ears in spite of heavy, perennial browsing. It is based on the number of times browsed and the percentage of living browsed woody plants on the sample plots. Only those well represented in the tally are con- sidered. Trees Shrubs Yellow Birch Alountain Holly Red Maple Witch Hobble ' White Pine Wild Raisin Shad Bush ^laples (sugar, mountain, striped), hemlock, skunk currant, choke- berry, willow and honeysuckle have withstood browsing fairly well in other sections and some of these may be more hardy than those listed above. For the swamp type six times as many browsed balsams died in the few years preceding the investigation as unbrowsed. As explained previously the significance of these fig- ures in regard to browsing in general, cannot be definitely stated, but indications point to a low "browsing resistance." The form and growth of red maple, as illustrated by Figure 4, may be alYected for several years, if not pernianentl\\ Whenever groups of red maple escape after heavy browsing the escaping point is marked by a knot of dead stubs or an irregularity in each stem (Fig. 21). Some of these eventually sulYer from decay at the old escaping point, while others develop sharp angles or are broken by snow and wind. Although white pine persists in spite of heavy browsing, it sutYers considerably in form during the early vears (Fig. 6) — this handicap may last for many years. Addi- tional damage ma\- result from butt rot at the base of the dead browsed stubs. 52 Roosevelt Wildlife Bulletin Wlien lieavily browsed, some species, such as yellow birch, exhibit strong tendencies to sprout from the root collar. Certain species, such as red maple, apparently put the excess vigor from the browsed branches into shoots. Mountain holly becomes very compact and hedge-like when heavily browsed, but when an opportunity occurs to escape it has plenty of vigor to throw out several shoots. When wild raisin has been heavily browsed for several years it usually forms abnormally dense clumps. This indicates that browsing encourages the growth of sprouts from the root system. The reac- tion of witch hobble is discussed under a different heading. Experi- ments may eventually provide more exact knowledge, when the variables are controlled or measured and clipping is done mechani- cally to simulate browsing. Witch Hobble as a Key Species. — An important outgrowth of preference of browse and browse resistance studies is the selection of what has been termed a key species. That is, an indicator plant which will reflect by its own state the general intensity of browsing on an area. An ideal key species should have the fol- lowing qualifications : 1. Common to all parts of the feeding ground. 2. Availability when needed by deer. 3. Capable of reflecting various degrees of browsing by its reac- tion to damage. 4. Stability as a food. Used hal)itually until the supply is exhausted — not a "tid-bit." 5. Ability to survive although heavily brow.sed. The seasonal change in deer diet, with respect to the w^estern Adirondacks, obviously eliminates the possibility of finding a key species applicable at all seasons of the year. Therefore, consid- eration is given to a woody species that can be used to gauge only critical winter season browsing. No other species encountered in the field, either tree or shrub, conforms to the requirements listed so well as witch hobble. This plant is generally distributed throughout the Adirondack forest. However, it is absent in most swamps, certain heavily cut areas and young burns. The other qualifications are well met. It is especially useful in demonstrating several intensities and stages of browsing (figs. 22, 23 and 24). Specimens which have sustained very little browsing produce long slender laterals, while constantly browsed stems become stunted and have a knotty appearance. A normal Deer Browsing in the Adirondacks 53 witch liobble has buds or branches on the stem occurring several inclies to about a foot apart. When a bud is eaten the stem dies back the entire distance to the next bud. Figure 24 illustrates heavy browsing succeeded by a period of light browsing, followed again by heavy browsing. In general, it was observed that when little browsing occurred on witch hobble, damage was correspond- ingly light on other woody species. Conversely, where heavily damaged witch hobble occurred, a general increase was noted in the browsing sustained. SUMMARY AND CONCLUSIONS 1. At present western Adirondack deer usually browse woody growth during the dormant season only. This of course includes a portion of the fall and spring. 2. Evidence of deer browsing on young trees generally remains for several years. The date such damage occurred usually can be determined by careful examination of living specimens, often after a lapse of six or more years. 3. Deer do not usually browse on the leaders of trees six feet or over in height. 4. Deer feed on practically every woody plant except spruce, when in yards of the western Adirondacks. 5. Deer may enable red spruce to assume dominance by selective browsing on its competitors. This is especially true in drained swamp type where hardwoods, especially red maple and yellow birch, outgrow it ordinarily. 6. When red spruce is not dominant, as in most spruce flat re- generation, stand composition is nevertheless influenced because of damage to competitors. 7. In old-growth stands the composition of the understory is changed due to browsing of certain species in the undergrowth. This influence is cumulative. 8. Red maple, yellow birch, mountain holly and witch hobble have special appeal for deer in the western Adirondacks. 9. Red maple, yellow birch, mountain holly and wild raisin are particularly resistant to repeated browsing. Apparently balsam and mountain ash are not. 10. Witch hobble is the most satisfactory key species or indicator for the general degree of browsing sustained in old growth of western Adirondack forests, 54 Roosevelt Wildlife Bulletin SCIENTIFIC NAMES OF THE TREES' AND SHRUBS^ MENTIONED IN THE TEXT WHICH WERE ENCOUNTERED IN THE FIELD Trees Aspen, Trembling Populus treinidoides Michx. Balsam Abies balsamea (Lj Miller Beech Fagus grandiftora Ehrhart Birch, Gray Betula populifolia Marsh. Birch, Yellow Betula liitea ]Michx. Cedar, White Thuja occideii talis L. Cherry, Black Pniiius serotina Ehrhart Cherry, Pin Fruuus peimsylvanica Linnaeus filh Hemlock Tsuga canadensis (L.) Carriere Maple, Mountain Acer spicatuin La Marck Maple, Red Acer rubrum L. Maple, Striped Ace'r peuHsylvanicnui L. Maple, Sugar Acer saccharuin Marsh. Pine, W'iiite Pinus Strobus L. Spruce, Black Picea mariaiia (Miller) Britton, Sterns, and Poggenherg Spruce, Red Picea rubra Link Tamarack Larix lariciua ( Uu Roi ) Koch Shrubs Alder, Speckled Alnus iucaiia Moencli. Blackberry, Mountain. . . . Rubus allcghcniciisis Porter Blueberry, Sour-top J'acciniuiii canadeuse Kalm. Chokeberrv 4ronia niclanocarpa Elliott Hardback Spiraea tomenlosa L. Honeysuckle, Fly Lonicera oblotigifolia Hook Labrador Tea Ledum groeulandicuni Oed. Leather Leaf ChainaedapJine calyculata Moench Meadow-sweet Spiraea latifolia Borkh. Mountain Ash. Sorbus americaua ^l^Lrsh. Mountain Holly Xeiiiopauihus mucronata Trel. Raspberry, Red Rubus idaeus L. Shad Bu.>;h Amelanchier ccniadeusis Med.^ Skunk Currant Ribes gland ulosum Grauer Sweet Gale Myrica Gale L Viburnum Jlburnuui sp.^ Wild Raisin Viburnum cassinoides L. \Mllow. Bebb's Salix Bebbiana Sarg. \\'itch Hobble Viburnum alnifolium IMarsh. Yew, Canada Taxus canadensis ^larsh. 'From Sudwortli (1927I. -From Rehder (iqj7). .\ few specimens of A. taeris Wieg.. were found in the tally. They were included as shad bush. * Probably J', optilus. Deer Bro-a'siiu/ in flic . IdiroiKlacLw 55 BIBLIOGRAPHY Alli;n, W. E. 1929. 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Some Factors Influencing the Reproduction of Red Spruce, Balsam Fir. and White Pine. Jour. For., Vol. 15. pp. 827-853- Deer Bro7K's'ni(] in the Adiroiidaeks 59 MouGix, M. p. 1931. La Restauration dos Alpcs. Direction Gencrale Des Eaux et Forets, Eaux et Genie Rural, Ministerie De L'agricul- ture, pp. 1-584. Paris. (Translations of Selected Por- tions No. 41, A. H. Krappe, Div. of Silvics, U. S. Forest Service. July 10, 1934, Wash., Y). C.) MowRY, Jessk B. 1920. The Nature and Development of Forest Types. Rhode Island Forestry Comm., Providence. 18 p. Murphy, L. S. 1917. Red Spruce: Its Growth and Management. U. S. D. A. Forest Service Bull. 544. 100 p. 1935. Planning for Wildlife in the United States. Part 9 of the Supi)lenientary Report of the Land Planning Committee to the National Resources Board. U. S. Govt, print. of¥. NiCHOL, A. A. 1936. The Experimental Feeding of Deer. U. S. Govt, print, oft'. Proc. North American Wildlife Conference, pp. 403-410. Pearson, G. A. 193 1. 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Some Late Winter and Early Spring Observations on White-tailed Deer of the Adirondacks. Roosevelt Wild- life Bull. Vol. 6. No. 2, pp. 327-385. Deer Browsing in the Adirondacks 6i SUDWORTH, G. B. 1927. Check List of the Forest Trees of the United States, their Names and Ranges. U. S. D. A. Misc. Circ. 92. 295 p. Taylor, W. P., and McGinnies, W. G. 1928. The Bio-ecology of Forest and Range. Sci. Monthly, Vol. 27, No. 2, pp. 127-182. TouMEY, James W. 1928. Foundations of Silviculture upon an Ecological Basis. John Wiley and Sons, New York. 438 p. TowNSEND, M. T., and Smith, M. W. 1933. The White-tailed Deer of the Adirondacks. Roosevelt Wildlife Bull. Vol. 6, No. 2, pp. 161-325. United States Senate 1931. Wild Life Conservation. Wash. D. C. Rpt. Spec. Comm. on Conservation of Wild Life Resources, Rpt. 1329, pursuant to Senate Resolution 246, 71st Congr., 3rd Sess. Westveld, Marinus 1 93 1. Reproduction on Pulpwood Lands in the Northeast. U. S. D. A. Tech. Bull. 223. 52 p. Roosevelt Wildlife Bulletin, Vol. 6, No. 3. July 1935. I. A Popular Account of the Bird Life of the Finger Lakes Section of New York, with Main Reference to the Summer Season. .Chas. J. Spiker Roosevelt Wildlife Bulletin, Vol. 6, No. 4. January, 1937. 1. Part L Preliminary Reconnaissance of the Land Vertebrates of the Archer and Anna Huntington Wild Life Forest Station Charles E. Johnson 2. Part IL Preliminary Reconnaissance of the Waters of the Archer and Anna Huntington Wild Life Forest Station and their Fish LiJiabitants Wilford A. Dence ANNALS Roosevelt Wildlife Annals, Vol. 2, No. i. January, 1929. I. The Red Squirrel : Its Life History and Habits, with Special Reference to the Adirondacks of New York and the Harvard Forest. R. T. Halt Roosevelt Wildlife Annals, Vol. 2, No. 2. October, 1929. 1. The Ecology of Trout Streams in Yellowstone Park Richard A. Muttkowski 2. The Food of Trout Stream Insects in Yellowstone Park Richard A. Muttkowski and Gilbert M. Smith Roosevelt Wildlife Annals, Vol. 2, Nos. 3 and 4 (Double Number). January, 1932. I. Ornithology of the Oneida Lake Region ; With Reference to the Late Spring and Summer Seasons Dayton Stoner Roosevelt Wildlife Annals, Vol. 3, No. i. January, 1932. I. Parasites of Oneida Lake Fishes. Part I. Descriptions of New Genera and New Species H. J. Van Cleave and J. F. Alueller Roosevelt Wildlife Annals, Vol. 3, No. 2. October, 1932. 1. Parasites of Oneida Lake Fishes. Part II. Descriptions of New Species and Some General Taxonomic Considerations, Especially Concern- ing the Trematode Family Heterophyidae Justus F. Mueller and Harley J. Van Cleave 2. Trichodina renicola (Mueller, 1931), a Ciliate Parasite of the Urinary Bladder of Esox niger Justus F. Mueller Roosevelt Wildlife Annals, Vol. 3, Nos. 3 and 4 (Double Number). July, 1934. I. Parasites of Oneida Lake Fishes. Part 3. A Biological and Ecological Survey of the Worm Parasites H. J. Van Cleave and Justus F. Mueller Part 4. .Additional Notes on Parasites of Oneida Lake Fishes, in- cluding Descriptions of New Species. . . .Justus F. Mueller Roosevelt Wildlife Annals, Vol. 4, No. i. December, 1935. I. Studies on Some of the Small Mammals of Central New York M. T. Townsend Roosevelt Wildlife Annals, Vol. 4, No. 2. May, 1936. I. Studies on the Bank Swallow, Riparia riparia riparia (Linnaeus) Dayton Stoner